Front Cover
 Back Cover

Group Title: Bulletin of the Florida State Museum
Title: The streams of Florida
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00000096/00001
 Material Information
Title: The streams of Florida
Series Title: Bulletin of the Florida State Museum
Physical Description: 92-126 p. : ; 23 cm.
Language: English
Creator: Beck, William M ( William Maser ), 1918-
Publisher: University of Florida
Place of Publication: Gainesville
Publication Date: 1965
Subject: Rivers -- Florida   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Bibliography: "References cited:" p. 123-126.
General Note: Cover title.
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Bibliographic ID: UF00000096
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0253
notis - ACK4301
alephbibnum - 000443510
oclc - 05067857
lccn - a 66007411

Table of Contents
    Front Cover
        Page 89
        Page 90
        Page 91
        Page 92
        Page 93
        Page 94
        Page 95
        Page 96
        Page 97
        Page 98
        Page 99
        Page 100
        Page 101
        Page 102
        Page 103
        Page 104
        Page 105
        Page 106
        Page 107
        Page 108
        Page 109
        Page 110
        Page 111
        Page 112
        Page 113
        Page 114
        Page 115
        Page 116
        Page 117
        Page 118
        Page 119
        Page 120
        Page 121
        Page 122
        Page 123
        Page 124
        Page 125
        Page 126
        Page 127
    Back Cover
        Page 128
Full Text




Volume 10 Number 3


William M. Beck, Jr.


lished at irregular intervals. Volumes contain about 300 pages and are not nec-
essarily completed in any one calendar year.



Consultants for this issue:

George K. Reid

Selwyn Roback

Communications concerning purchase or exchange of the publication and all
manuscripts should be addressed to the Managing Editor of the Bulletin, Florida
State Museum, Seagle Building, Gainesville, Florida. 32601

Published December 8, 1965

Price for this issue $.50O


"I !V



SYNOPSIS: The various classifications of Florida streams proposed in the litera-
ture are reviewed and a revised classification offered. This is examined both
statistically and with regard to factors controlling the distribution of aquatic in-
vertebrates as outlined in Berg's Susaa study. Finally, the stream types proposed
are delineated chemically, physically, and biologically.


Historical Review -- 93
Statistical analysis 96
FACTORS ---- 105
Velocity ---------_ 105
Substratum --107
Vegetation --- 109
Temperature 109
Oxygen -- 111
Water hardness 112

Geographical distribution ---- 114
Pollution .....----------- 115
Discussion ------------- 115
The Sand-bottom Stream ----- 116
The Calcareous Stream ------ 117
The Larger Rivers ------- 117
The Swamp-and-bog Stream 120
The Canals of Southeastern
Florida -------- 120
SUMMARY --------- 122
REFERENCES CITED --..... --- 123

1 The author is senior biologist of the Bureau of Sanitary Engineering, Florida
State Board of Health. He is particularly interested in the ecology of stream
invertebrates. This paper was supported in part by Research Grant No. AI-
4098-04, Institute of Allergy and Infectious Diseases, National Institutes of
Health. Manuscript submitted 29 April 1965.-ED.

Beck, William M., Jr. 1965. The Streams of Florida. Bull. Florida State Mus.,
vol. 10, no. 3, pp. 91-126. ;



Florida is abundantly supplied with varied and beautiful inland
waters. Except for a few papers noted below, most limnological in-
formation on these waters is to be found in various faunal studies in
which limnology is of importance mainly to help explain the ecologi-
cal and geographical distributions of single groups of organisms. This
has resulted in a variety of classifications which, when examined care-
fully, present a certain uniformity of considerable interest.
Population growth in Florida in the past 10 years has been little
short of phenomenal, and industrial growth has been correspondingly
great. This has placed a great demand on the natural waters of the
State which, though abundant, are not always equitably distributed in
time and space. Engineering alterations of the State's natural water-
ways have been extensive and are certain to increase in the near fu-
ture. While many people consider these changes both necessary and
beneficial, the fact remains that many of our most interesting water-
ways will soon cease to exist as natural entities.
The primary aim of this paper is to propose a uniform classifica-
tion of the lotic habitats of Florida based on various published re-
ports and my own investigations of the streams of Florida and ad-
jacent states. Except for certain comparisons, no attempt is made
to include lenitic habitats; several academic institutions in Florida
are now conducting extensive lake studies. A second purpose is to
record what the streams of Florida are like chemically, physically,
and biologically while many virtually undisturbed examples still exist.
It is hoped that this description of the streams of the area will be of
use to the many biologists in North America who are helping to iden-
tify the previously unknown components of our aquatic fauna.
Throughout this discussion a stream is defined as a body of water
with unidirectional flow of measurable velocity. This definition is
manifestly a loose one, but such looseness-or flexibility-is necessary
at present. Pollution is not a factor in any of the streams or reaches
of streams on which this study is based.
The study of stream limnology in the United States has lagged
far behind lake study. The basic reason for this is simple-the pio-
neering studies in North American limnology were made in universi-
ties located in lake regions. As these same institutions have remained
dominant influences to the present time, the lag in stream studies is
hardly surprising.
Most of the chemical, physical, and biological data for streams


throughout the nation are to be found in the files of state and fed-
eral regulatory agencies, and they are seldom published in other
than mimeographed form. This is strikingly true in Florida where
the files of the Game and Fresh Water Fish Commission, the Conser-
vation Department, the Geological Survey, and the State Board of
Health contain extensive data. My own work for the past 17 years
has involved water quality studies with major emphasis on streams.
The biological data available consist of an estimated 180,000 distri-
bution records for the macroscopic invertebrates.

The first serious study of aquatic biology in Florida was begun by
Rogers in 1922 and published in 1933. He was followed shortly by
Byers (1930). Subsequently the program was enlarged by their stu-
dents, Carr (1940), Hobbs (1942), Picice (1947), Berner (1950), Her-
ring (1951), Young (1954, 1955), and Beck (1958). Other information
on the streams of Florida may be found in the writings of Van Der
Schalie (1940), Odum (1953a, 1953b, 1956, 1957a, 1957b), Wurtz and
Roback (1955), Yount (1956a, 1956b), Sloan (1956), Whitford (1956),
Yerger (1960), and Reid (1961). Of the aquatic invertebrates, major
studies have been published thus far on the craneffies, dragonflies
and damselflies, mayflies, aquatic bugs, chironomids, crayfishes, and
plankton of selected habitats.
Currently under study in Florida are the amphipods (E. L. Bous-
field, National Museum of Canada), freshwater shrimps (H. H. Hobbs,
United States National Museum), stoneflies (A. R. Caufin, University
of Utah), caddislies (H. H. Ross, Illinois Natural History Survey),
ostracods (E. Ferguson, Jr., Lincoln University), blackflies (J. E.
Burgess, Jr., Florida State Board of Health), and freshwater sponges
(W. A. Moore, Loyola University). The mosquito fauna is one of
the most intensively studied in the world.
Modest knowledge exists of the Florida faunas in the following
groups: flatworms, bIyozoans, mysids, isopods, water mites, dobson-
flies, biting midges, and molluscs. The oligochaetes are rather poorly
known, as are the leeches, alderfies, spongillaflies, and aquatic moths.
Physical aspects of Florida's streams have been covered in the
publication by Smith et alii (1954).
It is impossible to acknowledge fully my indebtedness to the great
number of people who have contributed in one way or another to the
background of this paper. I wish especially to thank George A. Pur-
cell, Statistician, Florida State Board of Health, for help with the


statistical aspects of this study, and Lewis Berner and Oliver L.
Austin, Jr., of the University of Florida, for reviewing the manuscript.


Table 1 summarizes and compares five previously proposed stream
classifications. Their differences are largely semantic, except for
Carr's and Berner's recognition of canals and Bemer's emphasis on
vegetation. The semantic differences are significant in such a term
as "flatwoods river", which might actually be a sand-bottomed stream,
a calcareous stream, or a swamp-and-bog stream. The latter more
specific and descriptive terms are adopted for the present statistical
examination of stream type. This study does not include estuaries,
though they are mentioned briefly in the discussion of the St. Johns
Methods used for biological stream surveying published elsewhere
(Beck, 1954, 1955, 1957) are summarized here for reference. Selected
macroscopic invertebrates in Florida have been classified with regard
to their observed reactions to organic pollution in streams. The ap-
proach differs from most other proposed methods in that it uses
selected invertebrates to prove a stream clean on the basis of the pres-
ence of sensitive organisms, rather than the obverse approach that
proves a stream polluted by the dominance of tolerant taxa. It fur-
ther requires that each taxon used as an indicator organism be dis-
tributed widely both ecologically and geographically in the state.
A discussion of the mayflies may help clarify these points.
Florida species of the genus Stenonema are almost totally con-
fined to clean streams (Berner has recorded the genus from at least
one lake). As they are distributed rather widely in the state and their
reactions to organic pollution arc known, they are valuable as indi-
cator organisms. The species Callibaetis floridanus and Caenis di-
minuta, though much more widely distributed ecologically and geo-
graphically, are tolerant of gross pollution and are, therefore, of little
value to the program. Isonychia pictipes, common in the flowing
waters of northwestern Florida and sensitive to organic pollution, can-
not be used because of its restricted range. The several species of
the burrowing genus Hexagenia have both ecologic and geographic
restrictions in their distributions that make them almost valueless in
a statewide program.
Identifications are reported at several taxonomic levels, necessi-
tated by the state of knowledge of the various groups, as many
groups of aquatic organisms have been studied inadequately in Flor-

Vol. 10


J. S. Rogers A. F. Carr H. H. Hobbs L. Berner J. Herring This Paper

small streams small streams sand-bottomed vegetation sand-bottomed sand-bottomed

calcareous spring runs spring runs calcareous calcareous calcareous

swamp-&-bog with canals small rivers vegetation swamp-&-bog swamp-&-bog

lower streams larger streams flatwoods rivers slow-flowing larger rivers larger rivers

n. r.* canals n. r.* canals n. r.* canals

small rills -springs stagnant-rivers

* not recognized


ida. The oligochaetes are not identified beyond class, the simuliids
beyond family, and only a few caddisflies beyond genus. Although
this falls short of being completely satisfactory, at least the taxonomic
reporting is as consistent as possible throughout.

Thirty-five collections representing each of the live stream types
listed in Table 1, plus an equal number of collections from lakes
and ponds, were tabulated in order of decreasing frequency of taxa.
The 10 most frequently collected taxa were tabulated for compara-
tive purposes. Each fauna thus delineated was then compared with
every other one in the manner illustrated in Table 2. This table of
two colunms, A and B, is divided into four quadrants, 1, 2, 3, and 4.
Quadrant 1 lists the 10 most frequently collected taxa from the sand-
bottomed stream and quadrant 2 lists the numbers of records for the
same taxa in the swamp-and-bog stream. Thus, the most consistent
inhabitant of the sand-bottomed stream is a midge of the genus Tany-
tarsus, occurring in 33/35 collections, while the same genus occurred
in only 12/35 collections from the swamp-and-bog stream. The com-
parison here is not of rank, but of records per 35 collections. Quad-
rant 3 lists the remainder of the 10 most frequently collected taxa
from the swamp-and-bog stream and quadrant 4 gives the occurrence
per 35 collections from the sand-bottomed stream of those taxa listed
in quadrant 3. This was done for each possible pairing of water types.
The null hypothesis was used and the premise "There is no sig-
nificant difference between the faunas of the several water types
listed" was tested by chi-square.

chi2 = N( ad bc -N)2
(a+b) (c+d) (a+c) (b+d)
The formula includes Yates' correction (see Snedecor, 1956). Re-
sults of these tests are presented in Table 4.
Returning to Table 2, note that quadrant 1 lists five taxa repre-
sented by zeros in quadrant 2. The five taxa in question are all rhe-
ophilous taxa that would not predictably be found in waters with
extremely low velocities typical of the swamp-and-bog stream. They
are, therefore, typical or "character" (Berg, 1948) taxa when these two
water types are compared. It was decided to retest the pairings fol-
lowing the removal of all taxa represented by a zero in any cell and
use a revised null hypothesis as follows: "There is no significant
difference between the faunas of the several water types listed when

Vol. 10



Sand-bottomed Swamp-and-bog

1. Tanytarsus sp. 33 .-- --------- -- 12
2. Cheumatopsyche sp. 31 ---------------- 0
3. Mayfly n. 31 1. .. --------- 23
4. Stenonema exiguum 30 --- 0
5. Ceratopogonidae 28 7. --- ---------- 15
1. 2.
6. Oligochaeta 27 2. ..--- ----- -- 22
7. Corydalis cornutus 27 --... -------- 0
8. Damselfly n. 26 4 ...-... --- 19
9. Simuliidae 26 ..----- 0
10. Elmidae 25 .----------- 0

-.-- ...-------- 21 3. Beetle ad. 19
.. .. 20 5. Palaemonetes paludosus 16
9 6. Hyalella azteca 15
4. 3.
S23 8. Cryptochironomus fulvus 14
...- ..--- .. 12 9. Beetle la. 14
9 10. Chironomus attenuatus 13


Sand-bottomed Swamp-and-bog

1. Tanytarsus sp. 33 --.----------- --- 12
3. Mayfly n. 31 1. ------- 23
5. Ceratopogonidae 28 7 .. --- -- 15 2.
6. Oligochaeta 27 2. .... .--- ---- 22
8. Damselfly n. 26 4. -- - --------- 19

.--- - 21 3. Beetle ad. 19
20 5. Palaemonetes paludosus 16
9 6. Hyalella azteca 15
4. 3.
S23 8. Cryptochironomus fulvus 14
...-- --- 12 9. Beetle la. 14
9 10. Chironomus attenuatus 13


all "typical" taxa are deleted." The data from Table 2, revised ac-
cordingly, are presented in Table 3. Results are presented in Table 5.
Table 4 shows that the original null hypothesis must be rejected
unquestionably in every case. This resulted in a decision to remove
all taxa represented by a zero in any cell in any pairing and to recal-
culate chi2 and P values taking smaller degrees of freedom into con-
sideration. Three categories (calcareous X larger rivers, larger rivers
X swamp-and-bog, and swamp-and-bog X ponds) had no zeros in
the original pairings and were not recalculated. Table 5 shows that
even the revised null hypothesis had to be rejected in all cases. Ac-
tually, partly because of the decreased degrees of freedom, P values
were even lower than in the initial chi2 tests.


Pairings x2 d.f. value 0.95 C.L.

Sand-bottomed X Calcareous 194.57 15 0.005 Significant
Sand-bottomed X Larger Rivers 171.93 16 0.005
Sand-bottomed X Swamp-&-bog 311.29 16 0.005
Sand-bottomed X Canals 366.92 16 0.005
Sand-bottomed X Ponds 305.15 15 0.005
Sand-bottomed X Lakes 506.13 17 0.005
Calcareous X Larger Rivers 99.54 17 0.010
Calcareous X Swamp-&-hog 153.08 17 0.005
Calcareous X Canal 340.87 18 0.005
Calcareous X Ponds 214.48 17 0.005
Calcareous X Lakes 870.47 19 0.005
Larger Rivers X Canals 194.62 16 0.005
Larger Rivers X Swamp-&-hog 44.58 15 0.001
Larger Rivers X Ponds 77.65 15 0.005
Larger Rivers X Lakes 245.78 18 0.005
Canals X Swamp-&-bog 175.24 14 0.005
Canals X Ponds 203.45 14 0.005
Canals X Lakes 581.45 19 0.005
Swamp-&-bog X Ponds 28.99 13 0.007
Swamp-&-bog X Lakes 246.22 17 0.005
Ponds X Lakes 237.17 16 0.005

Rejection of both null hypotheses points out that significant simi-
larities are lacking. It would be possible at this point to resort to
other statistical methods that would determine the significance of
the existing differences. Such, however, does not appear to be neces-
sary, as data already prepared are sufficient to point out that distinct

Vol. 10


faunal differences exist between the water types listed, and typical
as well as ubiquitous taxa are listed for each stream type.


Pairings x2 d.f. value 0.95 C.L.

Sand-bottomed X Calcareous 84.13 12 0.003 Significant
Sand-bottomed X Larger Rivers 136.09 15 0.005
Sand-bottomed X Swamp-&-bog 79.93 11 0.002
Sand-bottomed X Canals 99.72 10 0.001
Sand-bottomed X Ponds 48.52 9 0.0005
Sand-bottomed X Lakes 103.05 4 0.0005
Calcareous X Swamp-&-bog 31.33 13 0.005
Calcareous X Canals 154.71 12 0.003
Calcareous X Ponds 58.77 12 0.003
Calcareous X Lakes 24.83 5 0.0005
Larger Rivers X Canals 138.90 14 0.005
Larger Rivers X Ponds 59.41 14 0.005
Larger Rivers X Lakes 81.10 8 0.0005
Canals X Swamp-&-bog 139.40 13 0.005
Canals X Ponds 178.18 13 0.005
Canals X Lakes 113.83 5 0.0005
Swamp-&-bog X Lakes 25.58 5 0.0005
Ponds X Lakes 105.85 7 0.0005

In Table 6 are listed the non-common faunal elements for each
pairing of water types. Those marked with an asterisk are rheophilous
taxa. The differences in taxa reveal differences in physical and
chemical features of the stream types in question. In the pairing of
sand-bottomed X calcareous streams, for example, the differences are
attributable to chemical, not physical factors, due to the fact that both
streams support rheophilous taxa, but the calcareous stream is popu-
lated by greater snail populations, one genus being confined to that
stream type.
In comparing the sand-bottomed stream with the swamp-and-bog
stream the only distinctive elements of either fauna are confined to
the sand-bottomed stream (within the limits of this particular pairing)
and all five distinctive taxa are rheobionts. The comparisons are dis-
cussed more thoroughly below.
The general approach used herein is hardly a new one, as the
following quotation from Curtis and Greene (1949: 83) shows:




Elmidane Goniobasis spp.*
Campeloma spp.

Elmidae* O

Cheumatopsyche spp.' O
Stenonema exigutum
Corydalis cornutus'

Cheumatopsyche spp.' Chlamydotheca
Stenonema exiguum*
Corydalis cornutus*

Cheumatopsyche spp.* Glyptotendipes lobiferus
Stenonema exiguum*
Corydalis cornutus*

Cheumatopsyche spp.' Chaoborus sp.
Mayfly nn. Coelotanypus spp.
Stenonema exiguum' Procladius spp.
Corydalis cornutus* Unid. chironomids
Damselfly mn. Hexagenia sp.
Simuliidae* Glyptotendipes parpies


Goniobasis spp.* O
Stenonema exiguum'
Corydalis cornutwu
Cheunatopsyche spp.'

Vol. 10

1965 BE

Goniobasis spp.*
Stenonema exiguum*
Corydalis cornutaus
Cheumatopsyche spp.*
Tanytarsus spp.



Goniobasis spp.' Glyptotendipes lobiferus
Stenonema exiguum*
Corydalis cornutus*
Cheumatopsyche spp.'

Goniobasis spp.' Chaoborus sp.
Stenonema exiguumn Coelotanypus spp.
Beetle larvae Unid. chironomids
Mayfly nn. Hexagenia sp.
Campeloma spp. Glyptotendipes paripes
Corydalis cornutus*
Beetle adults
Cheumatopsyche spp.*

Argia spp.' Chlamydotheca


Sphaeriidae O

Mayfly nn. Unid. chironomids
Damselly nn. Hexagenia sp.
Polypedilum illinoense
Argia spp.'
Pentaneura manilis gp.
Beetle ad.

Glyptolendipes lobiferus





Damselfly nn.
Hyalella azteca
Palaemonetes paludosus
Physa pumilia
Mayfly nn.
Beetle ad.
Dragonfly nn.
Ferrissia sp.


Coelotanypus spp.
Procladius sp.
Unid. chironomids
Hexagenia sp.
Glyptotendipes paripes

0 0

Mayfly nn.
Beetle la.
Damselfly nn.
Palaemonetes paludosus
Hyalella azteca
Cryptochironomus spp.
Beetle la.

Mayfly nn.
IIyalella azteca
Damselfly nn.
Cryptochironomus spp.
Physa pumilia
Beetle ad.

Chaoborus spp.
Coelotanypus spp.
Procladius spp.
Unid. chironomids
Hexagenia sp.

Coelotanypas spp.
Unid. chironomids
Hexagenia sp.

"The second method involves the study of as many stands as pos-
sible for each type of assemblage in the area. Only simple charac-
ters are examined in each stand, since the time and expense of de-
tailed studies are prohibitive when used on this scale. Presence is
the character usually recorded in the method (Braun-Blanquet, '32;
Dansereau, '43). If a given species occurrs in only a few stands, it
obviously is not an important member of the community, regardless
of possible prominence in any one stand chosen at random. The pres-
ence method gives a valuable picture of the range of variation be-
tween the stands of each community and may indicate interrelation-
ships between communities more readily than is possible with the
single stand technique. It is easy to apply in the field and is particu-
larly valuable in the detection of indicator species for land-use studies
and other practical conservation endeavors."

Vol. 10


The non-common faunal elements listed in Table 6 are potential
indicator organisms of the several stream types. This must, however,
be handled conservatively. In the pairing of sand-bottomed X larger
rivers, for example, it would appear that the elmid beetles are con-
fined to the sand-bottomed stream and are, therefore, indicative of
that particular stream type. Examination of all records for elmid
beetles reveals that they are occasionally found not only in the larger
rivers but in calcareous streams as well. They were simply not
present in the 35 collections from larger streams used in this study.
In all pairings involving canals the ostracod, Chlamydotheca uni-
spinosa, is the single apparently typical taxon, except in the pairing
with ponds in which this ostracod is also found. In lotic environ-
ments, therefore, this organism is indicative of canals.
Table 6 also shows that the relationships between sand-bottomed
streams, calcareous streams, and larger rivers are close. This is as it
should be, for all three are strictly lotic habitats, as opposed to two
other types which must be considered sublotic (see below). Snails
of the genera Goniobasis and Campeloma appear to be typical of the
calcareous stream, though only Goniobasis is truly typical, and the
faunal distinctions between calcareous and sand-bottomed streams
appear to be the result of chemical rather than physical differences.
When compared with each other neither calcareous streams nor larger
rivers have any distinctive taxa. This is hardly surprising as part of
total flow of the larger rivers originates from calcareous streams as
well as other stream types-hence the lengthier term "larger rivers
of mixed origin." It is interesting to note that when compared with
each other neither the swamp-and-bog streams nor the larger rivers
possess any truly distinctive faunal elements despite the strong re-
jection of the null hypothesis in this pairing. It is also of interest that
the 10 most frequently encountered taxa of the larger rivers include
not one rheobiont, although rheobionts are a definite part of the nor-
mal fauna of the larger rivers.
Table 7 lists 10 taxa considered ubiquitous in order of decreasing
ubiquity index. Mayfly nymphs include all mayflies other than the
genera Stenonema, Baetisca, and Hexagenia, these being the only may-
flies identified beyond order in our routine work. Oligochaetes as
reported here include only those forms either belonging to or similar
to the tubificids. Damselfly nymphs include all members of the sub-
order except species of the genus Argia. All beetles with the excep-
tion of the elmids are unidentified. Tanytarsus as here reported in-
cludes only unidentified species of the chironomid subgenus Atany-
tarsus. Only the bottom-dwelling, serpentine Ceratopogonidae are



included in the family-level reporting. The short bodied genera such
as Atrichopogon and the elongate inhabitants of algal masses are
Although this ubiquity index is an artificial device consisting sim-
ply of the average frequency of occurrence of the 10 most frequently
listed taxa universally encountered in the five stream types, it does
present some interesting facts. Most ubiquists so delineated are
simply unidentified representatives of larger taxonomic groups, thus
making the ubiquity index, within limits, a reflection of lack of identi-
fication beyond the higher categories. Of the three taxa listed that
represent specific identifications, each is actually not only the most
widely distributed member of its genus in Florida, but the most
widely distributed member of its order. Of the normally unidentified
mayflies at least two, Callibaetis floridanus and Caenis diminuta, may
well be true ubiquists (See page 94). Certainly the same may be
said for several oligochaetes, damselflies, and beetles.
This same approach may be used in another way by calculating
such an index for the stream types (Table 7). The fact that the highest
figure obtained is that for the canals, while the second highest is for
the sand-bottomed stream, appears to have some value. The canals
are carefully designed artificial bodies of water having quite uniform
physical conditions spatially, if not temporally. This merely indicates
the rather monotonous uniformity of the fauna, a fact easily deter-


Comparative Frequencies by Taxon
Stream Type Ubiquity
Taxa SB Cal. Lr S&B Can. Index

Mayfly nymphs' 31 24 22 23 31 26.2
Oligochaetes 27 14 21 22 32 23.2
Damselfly nymphs 26 13 18 19 35 22.2
Beetle adults 21 19 16 19 31 19.4
Tanytarsus (Atany.) sp. 33 17 12 12 17 18.2
Ceratopogonidae 28 9 16 15 22 18.0
Hyalella azteca 9 15 13 15 33 17.0
Physa pumilia 12 16 13 11 31 16.6
Palaemonetes paludosus 20 11 4 16 31 16.4
Beetle larvae 12 24 12 14 13 15.0

Stream type ubiquity index 21.9
SSee page 94

16.2 14.7 16.6 27.6

Vol. 10


mined by observation. The second highest index, that for the sand-
bottomed stream, is also logical, as this stream type is the easiest to
define of all the stream types found in Florida, the only great variable
being the degree of development of vegetation (Berner, 1950). The
three remaining types possess much greater variability and are, there-
fore, more difficult to delineate.

Although the literature on stream limnology is not extensive-
at least when compared with that on lake limnology-it is desirable
to compare material presented in this paper with certain published
observations from elsewhere. For this the excellent study of the
River Susaa by Berg (1948) is used as a model, particularly his outline
of factors controlling the distribution of aquatic organisms (p. 285).
Each of these factors is discussed below in detail with regard to the
proposed classification of the streams of Florida and to selected pub-
lished material.
A great deal has been written about the influence of velocity on
the composition of stream faunas, of the causes of this influence, and
of morphological adaptations to the lotic environment. That velocity
exerts a major influence on the distribution of aquatic organisms is
unquestioned, but no attempt is made to review the literature on this
subject completely.
Shelford and Eddy (1929: 382) state: "Our hypothesis is that per-
manent stream communities exist, undergo successional development,
reach and maintain a quasi-stable condition, and manifest seasonal
and annual differences, as do terrestrial and marine communities."
While I agree with this statement in general, some limitations should
be imposed on it. The fact that seasonal variation is of minor con-
sideration in Florida is not a fault. The main problem lies in the
complex variations in minor habitats within short reaches of streams.
Berg (1948: 285) states it thusly: "A water course changes its char-
acter more or less during its course from its rise to its discharge, and
the reaches that succeed each other may differ so much that they
have merely ecological features of minor importance in common.
"The consequence of this fact is that it is not possible to undertake
an ecological grouping of water courses but merely of reaches of
water courses. The unit of the system will not then be an entire
water course, but a reach of one within which the life conditions are
in the main uniform."


Thus the stream types proposed here are in a sense merely "reaches
of water courses." Beck (1954: 216-7) points out that the Suwannee
River in its flow across Florida is successively a swamp-and-bog
stream, a sand-bottomed stream, a calcareous stream, and, according
to some authors, a larger river of mixed origin. This succession is
due to velocity, substrate, and chemical alteration of the "river" water
by the discharge of numerous calcareous springs. In addition, no
mention has been made of estuaries, a stage through which most
rivers pass and which is largely ignored in the present paper. There-
fore we have a choice of considering the Suwannee either as four
rivers, or as a single river with four recognizable and widely dupli-
cated reaches. Obviously the latter is the only possible choice.
Ruttner (1953: 199) makes the following statement: "Early work-
ers were inclined to attribute these specific effects of swiftly flowing
water to its higher oxygen content. It is easy to demonstrate, how-
ever, that even cascading water never has an oxygen content higher
than will correspond to the momentary saturation equilibrium with
respect to the air, whereas in standing water supersaturations occur
commonly. The effect of strongly agitated water in promoting growth
and respiration must, therefore, have some other basis. In quiet or
in weakly agitated water, the organisms are surrounded by a closely
adhering film of liquid, which speedily forms around the animal or
plant a cloak impoverished of substances important for life. In a
rapid current, however, the formation of such exchange-hindering in-
vestitures is prevented, and the absorbing surfaces are continually
brought into contact with new portions of water as yet unutilized.
In this manner moving water promotes respiration and the getting
of food much more than quiet water of the same content; it is not
absolutely but rather physiologically richer in oxygen and nutrients.
A current consequently promotes respiration as well as eutrophica-
These statements are firmly supported by empirical data in a
paper by Whitford and Schumacher (1961: 423) who report: "An
'inherent current demand' by this lotic species was indicated (a) by
x10 increase in P32 uptake at current equivalents of 18 cm/sec;
and (b) by a 70% increase in respired CO2 in dark bottles at current
equivalents of 15 cm/sec. In a current this demand is satisfied by
the creation of a steep diffusion gradient." The reader is also referred
to Nielsen's (1950) excellent review of morphological adaptations to
velocity and of the specialized feeding habits of rheobionts.
In summary it may be said that faunal responses to lotic environ-
ments involve specialized feeding habits, a minor amount of morpho-

Vol. 10


logical adaptation, and an unknown amount of physiological adap-
In my work I have found a descriptive terminology convenient for
reporting velocities. Swift flow is that velocity in which Plccoptera
are found in Florida. Moderate velocity is any velocity sufficient to
maintain a population of Simuliidae (these are confined to running
water in Florida). Any velocities below these two loosely defined
levels are termed low. We have only recently obtained and started
using velocity meters in our biological work. When the results of
this work are available it will be interesting to see how well the actual
measurements support the observed qualitative terminology.
From the standpoint of velocity, the lotic habitats as proposed
herein must be divided into two groups: lotic and sublotic. Despite
the difficulty of measuring the velocity of a swamp-and-bog stream,
it is nevertheless flowing water with a transport of water from head
to mouth. In the canals the seaward flow may be quite swift for defi-
nite periods, may be reversed completely, or may remain stagnant
for days at a time. They must be considered lotic habitats because
the long-range flow is seaward and because the visiting observer may
encounter flowing water at any given time. This latter reason gains
strength in light of early work with these canals, when biologists first
realized that the fauna was a strange one mainly in that no stream
species were present despite the obvious flow. It might be logical
to change the above terminology to permanently lotic and seasonally
lotic, because rainy season flows in the canals are always seaward and
in dry seasons the flow may ease entirely.


The relationship between velocity and bottom deposits (or sub-
stratum) is directly reciprocal, in that the substratum determines the
velocity and the velocity determines the substratum. For instance,
bottom deposits of mud do not occur under swift water, nor are bare
rocks and rubble typical of sluggish water. As Hesse, Allee, and
Schmidt (1937: 304) point out: "Division into lower river (with a
minimum of erosion and a maximum of deposit), middle river (with
a balance between erosion and deposit and a more noticeable side
erosion), and upper river (with a maximum of deep erosion and a min-
imum of deposit) is also frequently inapplicable." The fact that such
a breakdown of stream courses is frequently inapplicable does not
preclude the fact that it is often applicable.


The swifter portions of a stream system are generally found in
its headwaters (the head-streams and highland brooks of Carpenter
(1928), the torrential streams of Nielsen (1950)). While we have no
torrential streams in Florida, many of our streams, the Suwannee,
Withlacoochee, and Chipola rivers for instance, have well-developed
rapids. Here the substratum is frequently eroding limestone, often
well supplied with growth of moss. Generally downstream from
these stretches (the Chipola is an exception) are areas of a typical
sand-bottomed nature and, though the Suwannee farther downstream
becomes a calcareous stream with many limestone outcroppings, it
eventually becomes a slower river with little erosion and greater dep-
The substrata of the streams of Florida may be divided into two
main categories-hard and soft. The hard materials may be sub-
divided into limestone and clay. The fauna of the limestone bottoms
is largely a function of velocity, which must be high enough to pre-
vent the deposition of finer soft materials. The faunas of clay bot-
toms are a function of position, a matter of whether the exposed
face of the clay is horizontal or vertical. For example, two Florida
chironomids, Xenochironomus taenionotus and X. rogersi, are com-
monly found burrowing in clay; the former species occupies vertical
clay banks through which a stream has cut, the latter burrows in hori-
zontal beds. Another occupant of the horizontal clay beds, Glypto-
tendipes meridionalis, has not been found coexisting anywhere with
X. rogersi.
Soft bottoms divide less distinctly into two basic categories-
organogenic and minerogenic-, and all degrees of intergradation
exist between them. In general it may be stated that the lower the
velocity, the finer (or lighter) the bottom materials. Thus a basic
difference between the sand-bottomed stream and the swamp-and-bog
stream lies in coarser sand bottom with a minor amount of organo-
genic material in the former, and finely divided organic detritus with
a minor amount of fine sand in the latter. This difference in bottom
deposits is demonstrated faunistically by a bottom community in the
swamp-and-bog stream resembling what some have termed a "pollu-
tional" fauna. This consists of tubificid worms, red chironomid larvae
(Chironomus, Cryptochironomus, etc.), and serpentine ceratopogonid
larvae. In contrast the sand-bottomed stream, while supporting many
of these same groups, has them represented by smaller numbers and
is also populated by larger burrowers such as dragonfly nymphs (Gom-
phus, Progomphus, Aphylla), mayflies (Hexagenia, Ephemera), cad-
disflies (Molanna, Phylocentropus), and lumbriculid oligochaetes.

Vol. 10


Berner (1950) was the first worker in Florida to emphasize strongly
the influence of vegetation on the distribution of aquatic insects.
This and his recognition of canals as a definite stream type are the
major differences between his classification and the basic one Rogers
(1933) proposed. The importance of vegetation is best demonstrated
by the sand-bottomed stream. In the western panhandle of Florida
are many examples of this steam type, apparently quite young, and
almost totally lacking in aqautic vegetation. Bottom deposits consist
mainly of course, shifting sand. Invertebrate faunas are meagre. In
one such, Canoe Creek in Escambia County, a rich invertebrate fauna
lives in accumulations of dead leaves held against the upstream sides
of small stumps by the high velocity of the water. The remaining
minor habitats are virtually sterile. Similar streams in the same basin
more adequately supplied with submerged vegetation support much
richer faunas in a greater variety of minor habitats.
Streams such as Canoe Creek naturally lack all species that re-
quire vegetation for food or shelter. Missing groups include the
pyralidids, leaf-mining or browsing chironomids, numerous mayflies,
caddisflies, Odonata, and most macro-crustaceans.
Florida has a rich flora of tropical aquatic plants, some of which
such as Eichhornia crassipes, Pistia stratiotes, Alternanthera philox-
eroides, and Salvinia rotundifolia, have become major nuisances. It
is interesting that only one of these, Alternanthera, has had a species
of aquatic insect, Nanocladius alternantherae, described with that
plant an apparent host. The date of the introduction of the water
hyacinth (E. crassipes) is the only one known with certainty (Goin,
1943: 143). It appeared near New Orleans in 1835 and was estab-
lished in Florida in 1840. Goin also notes that the salamander Pseudo-
branchus striatus axanthus is restricted to the water hyacinth habitat
m Florida and is the only aquatic vertebrate so to be. It is surprising
that no invertebrate has as yet become closely associated with this
Aquatic vegetation is, therefore, a major distributional factor in all
types of aquatic situations in Florida.

Temperature extremes in Florida do not prevent the year-round
occurrence of the aquatic stages of most indigenous invertebrates.
Stream quality surveys may be run in January just as effectively as


in June. This is not to suggest that temperature is unimportant; the
minimized variation actually can be a source of difficulty.
Residents of Florida have been concerned for many years about
the introduction of noisome tropical plants. Florida is subject every
few years to a severe winter (by Florida standards) with freezes as
far south as the northern Everglades. These occasional periods of
severe weather limit the spread of most terrestrial and aquatic species
of tropical origin. As an example, the midge, Metriocnemus abdom-
ino-flavatus, described from the bromeliads of Costa Rica, was fre-
quently found inhabiting the bromeliad, Tillandsia utriculata, as far
north as Vero Beach prior to the severe winter of 1957-58. Subse-
quent repeated samplings did not yield a single specimen of this
midge north of Ft. Lauderdale until May 1964.
The constant temperatures of the springs and, to a lesser degree,
the spring runs of the State pose a definite problem with regard to in-
troduced aquatic organisms. One such stream, located in an area of
tropical fish hatcheries, is known to support some 10 species of exotic
fishes, at least some of which seem to have become established (J. E.
Burgess, personal communication). No list of introduced species of
plants and animals found in the canals of Broward and Dade Counties
has been compiled, and the list might well be a long one.
Recently John H. Davis of the University of Florida (personal
communication) pointed out that although southern Florida has ap-
pealed to many workers as a place to study the effects of high mean
annual temperatures on the distribution of organisms, actually, this
area is the best place in the continental United States to study the
biotic effects of cold. My own observations support this strongly.
The above paragraphs are not meant to imply that all organisms
occurring in tropical latitudes require a high mean annual tempera-
ture. Mention has been made of the large ostracod, Chlamydotheca
unispinosa, which is so prominent in the canal fauna. This ostracod,
described from the cenotes of Yucatan, has been reported also from
Jamaica, Maryland, Ohio, and Louisiana, hardly a tropical distribu-
tion (Tressler, 1959: 704).
The thermal effects of the springs of Florida are not inconsider-
able. Near the spring heads temperatures remain nearly constant
all year. Sloan (1956: 86) reports that during the period from Novem-
ber 1952 to February 1954, the temperature in the boil of Weeki-
wachee Spring varied from a minimum of 23.20C to a maximum of
24.0C, with a total variation of only 0.80C. Many of the chemical
components are almost as constant, making these springs and spring
runs virtual outdoor laboratories. The northern Withlacoochee River,

Vol. 10


a major tributary of the Suwannee River, rises in Georgia and in its
upper reaches in Florida is a typical sand-bottomed stream. During
a two-year study of these streams the following temperature varia-
tions were found:
Temp. C Temp. OC Annual
Minimum Maximum Range

Withlacoochee 9.5 30 20.5
Suwannee 15 28 13

The temperature differences in the two rivers are due mainly to
the discharge into the Suwannee of great quantities of spring water,
the thermal effects of which are obviously considerable.

Although dissolved oxygen is one of the most thoroughly studied
factors in the aquatic environment, it is frequently a misused and
misunderstood determination in stream work. At least part of this
is explained by the above quotation from Ruttner.
In an area as richly supplied with anaerobic (or nearly so) springs
as are many parts of Florida, the effects of spring discharge on the
oxygen relationships are considerable. The Fenholloway River in
one area was found to have a dissolved oxygen concentration of less
than 1.0 mg/L in times of low flow. This resulted largely from
the increased proportion of anaerobic spring water in the reduced
quantities of "river" water. A rich fauna containing a number of
pollution-sensitive species was present. Lowering the dissolved oxy-
gen from a normal summer concentration of about 7 mg/L to the
observed level would predictably destroy all but the most resistant
organisms, had this been due to organic pollution such as domestic
sewage. In pollution studies the effects of minor quantities of toxic
decomposition products are probably often disregarded and total
responsibility for faunal alteration placed on lowered oxygen concen-
Perhaps the most surprising oxygen relationships in any of the
waters of the state are those of the canals. Summer values are almost
unbelievably low. During August 1959, the North New River Canal
had oxygen concentrations ranging from a minimum of 0.5 mg/L to
a maximum of 3.8 mg/L, with the mean value for no station reaching
2.0 mg/L. Biochemical oxygen demand values never reached 3.0
mg/L. Pollution was not a factor, for good populations of several


sensitive sunfishes were present and invertebrate communities were
balanced and diverse.
Spot samples were taken from Ft. Lauderdale all the length of
the canal to and including Lake Okeechobee. Oxygen values in the
lake were not a great deal higher. The explanation of the low oxygen
figures in the canal proved to be geological. These canals are cut
through a thin overburden of soil into an extremely porous limestone.
Careful observation revealed a flow of canal water into this limestone
in some areas and out of it in others. Thus canal water and ground
water were continuous and interchangeable.
Dissolved oxygen figures, much like isolated pH values, are seldom
worth much by themselves. It matters little that the midge, Chirono-
mus attenuatus, has been found in waters having oxygen concentra-
tions ranging from 0.0 to 33.7 mg/L, but it is of interest that another
midge, Trichocladius extatus, has never been found in a stream with
less than 3.0 mg/L of oxygen. Largely because of Florida's geologi-
cal history, it is necessary to know an entire stream, its geology, soci-
ology, hydrology, and uses before its oxygen relationships may be

Just what Berg means by hardness is not clear, for he does not
define it. This discussion is based on the assumption that he means
not hardness alone, but the entire CO2 cycle.
Determination of hardness is often neglected in limnological work.
If we list the lotic water types of Florida in order of ascending hard-
ness we have listed the order of ascending faunal importance of mol-
luscs. This holds true within limits for other invertebrate groups, but
for less obvious reasons.
In the Suwannee basin in Lafayette County two small streams flow
under a highway within 100 yards of each other and join within 100
yards downstream from the bridges. The northernmost, slightly the
larger of the two, is a typical sand-bottomed stream. In July 1953
the southern stream had the following characteristics: pH 7.9, alka-
linity 164 mg/L, hardness 162 mg/L, CaCOs, 88 mg/L, MgCO5,
74 mg/L, and temperature 24"C. Color and turbidity were both low.
Calcareous characteristics were unquestionable and the spring origin
of this stream was subsequently confirmed. Below their confluence
the effects of the sand-bottomed stream predominate for approxi-
mately a mile to the point where this stream flows into the Suwannee
River. The Suwannce in this area is heavily populated with snails of

Vol. 10


the genera Coniobasis and Viviparus, the former the most typical in-
habitant of calcareous streams, as was pointed out above. Neither
occurred in the two small streams in question, although both could
have occupied the southern stream. Apparently a chemical barrier
between these streams and the Suwannee existed.
In March 1961, these streams were visited again. Though no
chemical sampling apparatus was available, it was immediately ap-
parent that the southern stream had changed completely, for it now
had the appearance of a typical sand-bottomed stream. Its color was
high, a condition never before seen here, and its discharge had in-
creased an estimated 25%, although the northern stream was low.
In the southern stream's narrow and well-defined bed, the increased
discharge was accompanied by a major increase in velocity.
Table 8 compares the species of chironomids present in 1953 with
those found in 1961. Only one species was present during both sam-
pling periods. An examination of all records available for each chiron-
omid listed for either period indicated that it was impossible to justify
the presence or absence of a single midge species on the basis of the
altered carbon dioxide cycle. What had formerly been a rich growth
of many different aquatic plants now consisted of scattered patches
of Hydrocotyle and Ludvigia, neither of which makes a good habitat
for chironomids. Their replacement of former growths of submerged
aquatic plants with ligulate leaves such as Vallisneria doubtless caused
the disappearance of at least the first two species in the left hand
column of Table 8. Former areas of quiet water with bottom deposits
of fine sand and organic detritus had disappeared, and only areas of
coarse sand remained. The four species in the left hand column
marked with an asterisk were gone simply because their proper minor
habitat no longer existed. The rest of the list reveals very little of the
possible factors responsible for these changes in the fauna.
The above suggests that alteration of the carbon dioxide cycle
within the limits that may take place naturally in Florida waters has
little direct effect on the chironomids.
The swamp-and-bog stream is an unusual body of water when the
carbon dioxide cycle is considered. With pH values ranging as low
as 3.8 (this minimum has been determined both colorimetrically and
potentiometrically) and figures as low as 4.0 not uncommon, one can
only wonder about the physiological strain this must place on the
buffering system within an organism's body. Theoretically, waters
with this low a pH value should be equally low in alkalinity, hardness,
and, consequently, buffering capacity.



Vol. 10

July 1953 March 1961

Cricotopus bicinctus
Thienemanniella xena
Tanytarsus guerla
Cryptochironomus fulvus*
Chironomus stigmaterus*
Chironomus attenuatus*
Tanytarsus sp. A*
Polypedilum sp. A
Pentaneura carnea
Tanytarsus politus
Tanytarsus sp. B Tanytarsus sp. B
Polypedilum halterale
Trichocladius extatus
Polypedilum tritum
Cricotopus sp. C
Unid. Orthocladiinae


It should, perhaps, be unnecessary to mention this as a factor in
the consideration of the fauna of a stream. Frequently, however,
such a factor is overlooked in studies of single streams, probably be-
cause it is so obvious.
The use of indicator organisms in my routine work keeps this
factor a matter of constant consideration in that a state-wide program
must be based on organisms of state-wide distribution or else consist
of a series of regional programs. Although the former approach is
used at present, much serious thought has been given to the possi-
bility that the latter might well yield a more definitive program. The
regional approach would definitely yield greater knowledge of the
individual taxa involved. This is especially true of the western por-
tion of Florida from the Apalachicola drainage to the Perdido River.
One aspect of distribution is the influence of man on former nat-
ural distributions. In southwestern Florida numerous canals dug for
drainage purposes differ from the canals of southeastern Florida in
that they have unrestricted flow to the Gulf of Mexico and, conse-
quently, unidirectional flow. They have, in essence, become streams.
Gradients have been produced that contribute velocities high enough
to support rheophiles and rheobionts. These canals (or ditches) have
not been considered in the stream classification because of their


ephemeral nature, though they have extended the ranges of such
stream-inhabiting forms as Rheotanytarsus exiguus, Cricotopus bicinc-
tus, hydropsychid caddisflies, and others to a considerable degree.
Work is now underway by several state agencies to control this
indiscriminate removal of surface and ground waters. If such con-
trol is not instituted, we can only expect accelerated salt water en-
croachment in an area of Florida already experiencing some water
shortage. When proper engineering controls are instituted ranges of
those species extended by the canals of unrestricted flow should again
be the natural ranges of the species in the streams of Florida.

Although it was pointed out above that pollution is not a factor
in the present study, it is certainly a major factor in the distribution
of aquatic organisms and, while not one of the factors listed by Berg,
is nevertheless carefully considered in his work.

The five types of lotic habitats described for Florida, while dis-
tinct entities, exhibit varying degrees of intergradation. Many sand-
bottomed streams contain varying amounts of spring water and, con-
sequently, vary in their chemical characteristics. Few, if any, cal-
careous streams do not have some contribution of acid waters. Only
the canals of southeastern Florida fail to intergrade with the other
water types. These, then, are the only truly distinctive aquatic en-
vironments in the state.
A certain amount of objection to this last statement is anticipated.
It may be argued, for example, that our many springs of the first mag-
nitude constitute a distinctive feature in Florida. Yet these springs
exhibit a wide variety of chemical and physical characteristics (Fer-
guson, et alii 1947; Whitford, 1956; Sloan, 1956). In addition, Florida
has several rivers that disappear underground, most reappearing not
too far away (Santa Fe, Chipola, St. Marks). The reappearance of
one, the Alapaha, has not positively been discovered, although several
small springs along the Suwannee River, into which the Alapaha dis-
charges during high water, have distinct color, a rare feature in Flor-
ida springs. These may well represent the reappearance of the
Another aquatic feature slighted in this paper is the roadside ditch.
These ditches, formed when highways are constructed, are of two



types. The first type contains runoff water and seepage water, has
little or no true flow, and is chemically, physically, and biologically
similar to a pond. The second type has constant unidirectional flow,
is supplied by permanent seepage, and is basically a sand-bottomed
The main point in this discussion is that it is possible, by selecting
odd and virtually unique examples, to propose an almost endless list
of possible lotic habitats. It is the purpose of the present paper to
delineate a simple, basic classification in which major stream types
are recognized and, perhaps, may be made recognizable to others.

The following five chemically, physically, and biologically distinct
stream types exist in Florida.

This is the most widely distributed and most frequently encount-
ered type of stream in the state. It has been the most typical lotic
feature of the area and is the one disappearing most rapidly with the
alteration of drainage patterns. The sand-bottomed stream is a prom-
inent feature of the Central Highlands (see Cooke, 1939, for a discus-
sion of the topographic regions of Florida), Coastal Lowlands, Mari-
anna Lowlands, and Western Highlands. The fauna is dominated by
rheophiles and rheobionts. Typical faunal elements are hydropsychid
and philopotamid caddisflies, mayflies of the genera Stenonema and
Isonychia, simuliid larvae, Plecoptera, orthocladiine chironomids,
elmid beetles, and Corydalis cornutus. Of all lotic habitats this is the
most typically so.
Chemically and physically the sand-bottomed stream is mildly acid
to circum-neutral (pH 5.7-7.4), has alkalinity ranging 5 to 100 mg/L,
hardness from 5 to 120 mg/L, color moderate to high, and of mod-
erate to swift velocity. Bottom deposits consist of fine sand with vary-
ing amounts of leaf and other organic detritus in the quieter reaches.
Areas of limestone outeroppings are frequent. In the western pan-
handle the sand-bottomed streams are usually swifter and have coarser
bottom deposits. Shifting sand bottoms are common. Plant growth
may be slight to quite dense and of great variety.
Typical examples are the Black Creek Complex, both Withlacoo-
chee Rivers, the Yellow, Blackwater, and Shoal Rivers.

Vol. 10


These streams are predominantly of spring origin and many of
the finest examples have been carefully protected because of their
beauty. Visitors to Florida find these a major attraction; a number of
the larger springs and their runs have been developed commercially,
and the natural aspects of most have been carefully preserved. These
are, indeed, a striking sight with their cool, very clear waters, dense
and varied growths of submerged plants, and banks shaded by large,
moss-hung trees. The beauty of these streams is actually a limnologic
feature in that it is the result of two factors; the clarity of the water
itself and the high concentrations of phosphorus.
Widely distributed in Florida, the calcareous stream is found in
the Central Highlands, Coastal Lowlands, Relict Areas (Beck and
Beck, 1959), Marianna Lowlands, and the southern portion of the
Tallahassee Hills.
-The fauna of these streams is less definitely rheophilous than that
of the sand-bottomed stream. The most obvious feature is their high
mollusc populations (Goniobasis, Campeloma, Viviparus, and Poma-
cea). Rheophiles and rheobionts are normally represented by hydro-
psychid caddisflies, mayflies of the genus Stenonema, a great variety
of chironomids, Corydalis cornutus, and occasionally simuliids and
The waters are alkaline (pH 7.0-8.2), the alkalinity ranging from
20 to 200 mg/L, hardness from 25-300 mg/L (omitting the oligohaline
and mesohaline springs of Whitford (1956)). The water is normally
clear (some examples have a slight turbidity from small amounts of
Montmorillonite clay in suspension) and generally low in color. Ve-
locity ranges from low to swift.
Bottom materials consist of sand, clay, limestone, and quite heavy
deposits of organic detritus in the slower reaches. Submerged plant
variety appears to be a function of bottom material.
Typical examples are the Suwannee, Silver Crystal, St. Marks, and
Wakulla Rivers. In size they range from small rills to large rivers.

This is a category of convenience, for no two of these streams
have many features in common. These are what Rogers (1933) re-
ferred to as "lower streams." Of the four examples in Florida, three
(Apalachicola, Choctawhatchee, and Escambia) are interstate, rising
in the hills of Georgia and Alabama. All three normally carry a sig-
nificant amount of clay and silt and are always turbid. The fourth,


the St. Johns River, lying entirely within the state, is quite clear but
with fairly high color. As Pierce (1947: 2) describes it:
"The St. Johns River rises in the Kissimmee Prairie, west of Mal-
abar and flows north for almost 320 kilometers (200 miles) to empty
into the Atlantic Ocean 34 kilometers (21 miles) east of Jacksonville.
An outstanding feature of this river is its low gradient, which is less
than 6.1 meters from source to mouth. A number of interesting phe-
nomena characterize this very small change in water level along the
course of the river. One is the distance from the mouth at which
daily tidal effects can be felt. Records of the Coast and Geodetic
Survey show mean tidal ranges of 1.4 meters at Mayport (inside the
mouth of the river) and 0.15 meters at Welaka (166 kilometers, or 103
miles, up the river). Other characters of the river which are associ-
ated with the low gradient are the slow current, the shallow depth,
and the relatively broad basin for most of its length.
"These features give the St. Johns River an older appearance than
its geological record (Cooke, 1939: 109) indicates. The valley of the
river, which dates from the Pleistocene period, was formed in part
from wave-built sand bars; its basin was formed from lagoons, streams,
and solution lakes."
This states accurately many of the unusual features of this most
fascinating river, which some maintain is not a river at all, but a series
of connecting lakes. Anyone who has gone around the Devil's Elbow
near Palatka in a small skiff would question both statements concern-
ing low velocity and non-river characteristics. Other features of this
stream are of equal interest. One of the most unusual, and perhaps
also unique, factors is the behavior of its salinity. Progressing from
the mouth upstream (southward) the salinity quite properly drops
until a minimum is reached downstream from Palatka, the exact point
varying with the season. From this point upstream the salinity rises
again from the discharge of a number of mesohaline springs (see Whit-
ford, 1956) in Marion County until the vicinity of Lake George is
reached, above which it suddenly drops. This produces, in effect,
a double estuary and permits the presence of a commercial crab
(Callinectes sapidus) fishery some 125 miles above the mouth. The
mesohaline springs are inhabited by several species of marine isopods,
and beach-hoppers may be found around the periphery of Lake
Obviously this stream cannot be included in any one of the des-
ignated types, as it has reaches of swamp-and-bog characteristics,
others that have sand-bottomed characteristics, and stretches not com-
parable to either.

Vol. 10


The chemical characteristics for this river defy summarizing, for
anything reported for one reach would be untrue of reaches a few
miles upstream or downstream. Suffice it to say that perhaps no river
in North America so warrants thorough study as does the St. Johns.
It is hoped that a complete and careful study of this river may be
made cooperatively within the next few years.
In contrast to the above probably no stream in Florida has been
as carefully studied over as many years as has the Escambia River.
This material has recently been summarized (Robert F. Schneider,
unpub. M.S. thesis) and, it is hoped, will be published. It will suffice
for the present to state that pH values for the upper Escambia River
range from 6.5 to 6.9, with an increase in pH in the lower reaches
from the influence of salt water penetration in the estuarine portion
of the stream. The upper stream is normally well supplied with dis-
solved oxygen, seldom falling as low as 75% saturation. Both alka-
linity and hardness are normally below 40 mg/L and chlorides below
20 mg/L. Regardless of flow this river always has a slight turbidity.
The third of the larger rivers, the Apalachicola, has also received
considerable attention. It is formed by the confluence of the Flint
and Chattahoochee Rivers near the junction of the states of Alabama,
Georgia, and Florida. At Chattahoochee, just below the confluence,
the 25-year average discharge of 22,240 cfs., is exceeded in Florida
only by the St. Johns River.
The following chemical and physical data are from two surveys
of this river, one during the summer and one during the winter. The
area covered is from below the junction of the Flint and Chattahoo-
chee Rivers to a point well above any possible estuarine effect, a dis-
tance of 76 miles. Ranges of the several determinations are as fol-
lows: pH 6.5 to 7.4, alkalinity 12 to 31 mg/L, hardness 22 to 56 mg/L,
sulfates 6.0 to 9.0 mg/L, color 10 to 32 units (USGS), and turbidity 13
to 48 units (Jackson turbidimeter). The banks in the area covered
are rather high and steep, and the stream has few shallow places and
few aquatic plants. Bottom deposits consist of coarse sand and lime-
stone and are singularly unproductive. Logs lodged against the
banks constitute the most productive habitat. The invertebrate fauna
is dominated by hydropsychid caddisflies, mayflies of the genus Sten-
onema, chironomids of the genera Polyprdibun and Xenochironomus,
the damselfly genus Argia, and molluscs. An outstanding feature of
this river is its monotony, abetted somewhat by dredging and water
level control.



These highly acid, suggish streams are most typical of the Coastal
Lowlands but occur occasionally in the Central Highlands. They
originate in swamps, sphagnum bogs, and marshes. They show a defi-
nite relationship to the sand-bottomed streams in that all chemical
differences are functions of the one significant physical difference,
velocity. An increase in gradient would convert them to the sand-
bottomed type by increasing turbulence, which in turn would increase
reaeration, reduce carbon dioxide, and increase pH and alkalinity,
and, finally, by removing the finer bottom sediments of organic silt
and replacing them with sand. The swamp-and-bog stream is, then,
a sand-bottomed stream with lowered velocity and the chemical and
physical attributes that accompany this lowered velocity. This is
suggestive of a successional relationship (see discussion of the Su-
wannee River above) but such is not proposed at present. While
examples of succession in space are available, as in the Suwannee,
streams in Florida have not been studied long enough as yet to per-
mit any extensive discussion of temporal succession beyond the prin-
ciples of base-leveling.
The swamp-and-bog stream has the following characteristics: pH
3.8 to 6.5, alkalinity and hardness both normally well below 40 mg/L,
color sometimes as high as 750 units, turbidity low, and carbon dioxide
at times above 100 mg/L. These streams support a surprising fish
fauna, with many species normally considered sensitive to high car-
bon-dioxide values such as sunfishes and darters.
Faunistically these streams differ little from an acid pond. Rhe-
ophilous forms are universally lacking, molluscs are represented only
by Physa pumilia, and the general fauna gives the impression to those
in pollution abatement of being composed almost totally of species
highly resistant to organic pollution, though the fishes are an ex-
In this study the term "canal," unless otherwise specified, always
refers to canals of the southeastern quadrant of the State.
From a point north of the St. Lucie Canal to a point south of
Homestead, a distance of some 140 miles, not a natural stream of any
significance remains along the east coast of Florida. All are now
canals. We have no records of the invertebrates that once inhabited
the natural waters of this area, with the exception of the mayflies,
beetles, and one or two other groups. That these former streams no
longer exist is perhaps unfortunate, but the canals replacing them

Vol. 10


represent a very interesting and complex series of habitats whose
faunas are uniform enough to permit fairly sensitive ecological differ-
entiating. In addition the canals probably represent the only lotic
aquatic habitats, insofar as streams are concerned, of the Coastal Low-
lands of southeastern Florida. As a result they become extremely im-
portant to the aquatic biologist and to limnology in general.
Chemically these waters are most unusual. Their oxygen rela-
tionships have been discussed above. Other factors are: pH 7.1 to
8.1, alkalinity 135 to 250 mg/L, hardness 150 to 295 mg/L, turbidity
is low, except in the vicinity of dredging, and color is about the same
as found in the sand-bottomed streams with the exception of some
of the southern Dade County canals, in which it is surprisingly low.
The most significant and misleading factor in the limnology of the
canals is velocity. A given canal may be observed to flow eastward
at quite a high rate; 24 hours later the same canal may have no water
movement at all. The over-all effect is to produce a sublotic environ-
ment. These may be classified as streams for the reasons outlined
above with one additional reason; they must be considered with re-
gard to what they actually are, a present-day substitute for the former
streams of the area.
The fauna is lenitic. It bears repeating that not a single rheoph-
ilous species has been found in these waters. The wide-spread pres-
ence of the large Central American ostracod, Chlamydotheca uni-
spinosa, the recently introduced and rapidly spreading Colombian
snail, Marisa cornuarietis (Hunt, 1958), and the presence of the exotic
Belonesox not only are, at present at any rate, typical of thelcanals,
but they suggest something of the future of these subtropical waters.
The chances are excellent that the faunas of these canals in the near
future will be composed to a significant degree of exotic, mainly trop-
ical invertebrates accidentally introduced by the aquarium industry.
Another aspect of these waters worth discussing is their relative
constancy with regard to chemical characteristics. This makes them
excellent waters for the study of the effects of pesticides and com-
mercial fertilizers used in the rich agricultural areas within their drain-
age (see Davis, 1943, 1946). Such studies have recently been insti-
tuted by the Dade County Department of Public Health, but it will
be many months before results of this work will be available.



1. In Florid6 five basic types of streams, or reaches of streams, are
chemically, physically, and biologically distinctive. These are: (1)
the sand-bottomed stream, (2) the calcareous stream, (3) the larger
rivers, (4) the swamp-and-bog stream, and (5) the canals of south-
eastern Florida.

2. Of these five types, three (1, 2, and 3) are unquestionably lotic
and two (4 and 5) are sublotic.

3. One type, the larger rivers (of mixed origin), is a category of con-
venience covering four larger rivers, no two of which are alike, yet
all differing markedly from the other proposed stream types.

4. After many years of study by a number of workers, the proposed
classification does not differ materially from the classification pro-
posed many years ago by J. Speed Rogers. This paper brings to-
gether observations based on several different groups of aquatic in-
vertebrates, instead of past classifications reflecting the distribution
of single family, order, or class.

5. Of the factors controlling the distributions of aquatic invertebrates
in Florida waters, the most important appear to be physical (velocity,
substratum, temperature, engineering alteration). Of secondary im-
portance are the chemical factors (oxygen, carbon dioxide cycle). In
tertiary position are the biotic factors (plant growths and parasitiza-
tion). A single factor outside the limits of the above listed is the
geographic distribution factor, often neglected.

Vol. 10


Beck, Elisabeth C.
1952. Notes on the distribution of Culicoides in Florida. Fla. Ent. 35 (3):

Beck, Elisabeth C., and W. M. Beck, Jr.
1959. A checklist of the Chironomidae (Insecta) of Florida (Diptera: Chirono-
midae). Bull. Fla. State Mus. 4 (3): 85-96.

Beck, William M., Jr.
1954. Studies in stream pollution biology. I. A simplified ecological classifi-
cation of organisms. Quart. Journ. Fla. Acad. Sci. 17 (4): 211-227.
1955. Suggested methods for reporting biotic data. Sew. Indust. Wastes. 27
(10): 1193-1197.
1957. The use and abuse of indicator organisms: in Biological problems in
water pollution. Public Health Service, Cincinnati. pp. 175-177.
1958. A study of the interrelations of selected chemical and physical factors
in the Suwannee River. Quart. Jour. Fla. Acad. Sci. 21 (1): 12-24.

Berg, Kai
1948. Biological studies on the River Susaa. Folia Limnologica Scandinavica.

Berner, Lewis
1950. The mayflies of Florida. Univ. Fla. Stud., Biol. Sci. Ser. 4 (4): 1-267.

Byers, C. Francis
1930. A contribution to the knowledge of Florida Odonata. Univ. Fla. Publ.,
Biol. Sci. Ser. 1 (1): 1-327.

Carpenter, Kathleen E.
1928. Life in inland waters. Sidgwick and Jackson, London. 1-267.

Carr, Archie F., Jr.
1940. A contribution to the herpetology of Florida. Univ. Fla. Publ., Biol.
Sci. Ser. 3 (1): 1-118.

Cooke, C. Wythe
1939. Scenery of Florida, interpreted by a geologist. Bull. Fla. Geol. Surv.
17: 1-118.

Curtis, J. T., and H. C. Greene
1949. A study of relic Wisconsin prairies by the species-presence method.
Ecol. 30 (1): 83-92.

Davis, J. H., Jr.
1943. The natural features of southern Florida. Bull. Fla. Geol. Surv. 25:
1946. The peat deposits of Florida. Bull. Fla. Geol. Surv. 30: 1-247.


Furguson, G. E., C. W. Lingham, S. K. Love, and R. O. Vernon
1947. Springs of Florida. Bull. Fla. Geol. Surv. 31: 1-196.

Goin, Coleman J.
1943. The lower vertebrate fauna of the water hyacinth community in north-
ern Florida. Proc. Fla. Acad. Sci. 6 (3-4): 143-152.

Herring, Jon L.
1951. The aquatic and semi-aquatic Hemiptera of northern Florida. Part 4:
Classification of habitats and keys to the species. Fla. Ent. 34: 141-146.

Hesse, Richard, W. C. Allce, and Karl P. Schmidt
1937. Ecological animal geography. John Wiley and Sons, New York: 1-597.

Hobbs, Horton II., Jr.
1942. The crayfishes of Florida. Univ. Fla. Publ., Biol. Sci. Ser. 3 (2): 1-179.

Hunt, Burton P.
1958. Introduction of Marisa into Florida. Naut. 72 (2): 53-55.

Nielsen, Anker
1950. The torrential invertebrate fauna. Oikos, 2: 176-196.

Odum, Howard T.
1953a. Factors controlling marine invasion into Florida fresh waters. Bull. of
Marine Sci. of Gulf and Caribbean, 2 (2): 134-156.
1953b. Dissolved phosphorus in Florida waters. Fla. Geol. Surv., Rept. of
Invest., Misc. Studies 9, Pt. 1: 1-40.
1956. Primary production in flowing waters. Limnol. Oceanogr. 1 (2): 102-
1957a. Trophic structure and productivity of Silver Springs, Florida. Ecol.
Monog. 27: 55-112.
1957b. Primary production measurements in eleven Florida springs and a ma-
rine turtle grass community. Limnol. Oceanogr. 2 (2): 85-97.

Pierce, E. Lowe
1947. An annual cycle of the plankton of four aquatic habitats in northern
Florida. Univ. Fla. Studies, Biol. Sci. Ser. 4 (3): 1-62.

Reid, George K.
1961. Ecology of inland waters and estuaries. Reinhold Publishing Corpora-
tion, New York. pp. 1-375.

Rogers, J. Speed
1933. The ecological distribution of the crane-flies of northern Florida. Ecol.
Monog. 3 (1): 1-74.

Ruttner, Franz
1953. Fundamentals of limnology. (transl. by D. G. Frey and F. E. J. Fry).
Univ. Toronto Press: 1-242.


Schneider, Robert F.
1964. An ecological survey of the Escambia River, Florida. unpub. thesis.
Illinois State University, pp. 1-229.
Shelford, V. E., and Samuel Eddy
S 1929. Methods for the study of stream communities. Ecol. 10 (4): 382-391.
Sloan, W. C.
S1956. The distribution of aquatic insects in two Florida springs. Ecol. 37
(1): 81-98.
m. sT.. David B., J. W. Wakefield, II. A. Bevis, and E. B. Phelps
S1954. Stream sanitation in Florida. Fla. Engineering Ser. 1: 1-149.

Snedecor, G. W.
1956. Statistical methods applied to experiments in agriculture and biology.
Iowa State College Press. Ames. pp. 1-534.
Tressler, Willis L.
1959. Ostracoda: in Fresh-water biology, ed. W. T. Edmondson, John Wiley
and Sons, New York. pp. 657-734.
Van Der Schalie, Henry
1940. The naiad fauna of the Chipola River, in northwestern Florida. Lloydia,
3: 191-208, Pls. 1-3, 1 map.

Whitford, L. A.
1956. The communities of algae in the springs and spring streams of Florida.
Ecol. 37 (3): 433-442.

Whitford, L. A., and G. J. Schumacher
1961. Effect of current on mineral uptake and respiration by a fresh-water
alga. Limnol. Occanogr. 6 (4): 428-425.
SWirth, Willis W.
1951. A new biting midge of the genus Leptoconops from Florida, with new
records of other American species. Proc. Ent. Soc. Wash. 53 (5): 281-
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1953a. Biting midges of the heleid genus Stilobezzia in North America. Proc.
U. S. Nat. Mus. 103 (3316): 57-85.
1953b. American biting midges of the heleid genus Monohelea. Proc. U. S.
Nat. Mus. 103 (3320): 135-154.
1962. A reclassification of the Palpomyia-Bezzia-Macropeza groups, and a re-
vision of the North American Sphaeromiini (Diptera, Ceratopogonidae).
Ann. Ent. Soc. Amer. 55 (3): 272-287.

Wirth, W. W., and R. W. Williams
S 1964. New species and records of North American Monohelea (Diptera: Cera-
J topogonidae). Ann. Ent. Soc. Amer. 57 (3): 802-310.



Wurtz, Charles B., and Selwyn S. Roback
1955. The invertebrate fauna of some Gulf Coast rivers. Proc. Acad. Nat.
Sci. Philadelphia. 107: 167-206.
Yerger, Ralph W.
1960. Etheostoma okaloosae (Fowler), a percid fish endemic in Northwest Flor-
ida. ASB Bulletin, 7 (2): 41.
Young, Frank N.
1954. The water beetles of Florida. Univ. Fla. Stud., Biol. Sci. Ser. 5 (1):
1955. A preliminary survey of the water beetle fauna of Glen Julia Springs,
Florida. Quart. Journ. Fla. Acad. Sci. 18 (1): 59-66.
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1956a. Factors that control species numbers in Silver Springs, Florida. Lim-
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1956b. Productivity of Florida springs. Third Annual Report to Biology
Branch, Office of Naval Research.

/ ,,

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