The fishes of the Santa Fe River system

Material Information

The fishes of the Santa Fe River system
Series Title:
Bulletin of the Florida State Museum
Hellier, Thomas Robert, 1928-
Place of Publication:
University of Florida
Publication Date:
Physical Description:
40, [6] p. : illus., map. ; 23 cm.


Subjects / Keywords:
Fishes -- Florida -- Santa Fe River ( lcsh )
bibliography ( marcgt )
non-fiction ( marcgt )


Thesis (M.S.)--University of Florida.
"Literature cited:" p. 35-37.
General Note:
Cover title.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright held by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, images and other media are for nonprofit, educational, and personal use of students, scholars, and the public. Any commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum. For permission or additional information, please contact the current editor of the Bulletin at
Resource Identifier:
AAA0845 ( LTQF )
ACK4304 ( NOTIS )
027874832 ( AlephBibNum )
05069179 ( OCLC )
a 67007280 ( LCCN )

Full Text







Volume II

Number I


Thomas R. Hellier, Jr.





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

WALTER AUF FENBEG, Managing Editor
OLIrvm L. AusrnN, JR., Editor

Consultants for this issue:
Warren Freiberg
Carter S. Gilbert

Communications concerning purchase or exchange of the
manuscripts should be addressed to the Managing Editor of
State Museum, Seagle Building, Cainesville, Florida. 32601

publication and all
the Bulletin, Florida

Published January 18, 1967

Price for this issue $.80



SYNOPSIS: This paper correlates the fish populations of the Santa Fe River with
such physical aspects as its underground channel, substantial subterranean drain-
age, and low chlorinity. Recorded from the river and its tributaries are 60
species of fishes representing 21 families and 13 orders. Pertinent life history
notes are given for many of the species, including one species nearly endemic
to the river, Micropterus notius.
The derivation of the ichthyofauna is suggested; 25 species comprise an cle-
ment widespread to the north, 18 to the southeast; 8 species are typically Florid-
ian, and 8 species are from marine waters. Euryhaline species in the Santa Fe
are generally restricted to the portion of the river downstream from the under-
ground channel. A new term is proposed, dihaline, to describe those species
that maintain breeding populations in both fresh and marine waters.


Introduction ................ 2
Description of the river .... ...... ............ . 2
Description of the stations ... . ...... ............ ........... ..---- --... - 6
Materials and methods ..... ....... .....-- 12
Annotated list of fishes ...... ......... 14
Discussion ........-------..-.... ---------. --...--....-..-- ...--------- 30
General fish fauna ---.. .............. ............. .... 31
Derivation of the ichthyofauna - --------- 33
Literature cited 35

SThe work reported on here was submitted to the Graduate School, University
of Florida, in partial fulfillment of the requirements for the Master of Science.
The author is now associated with the Department of Biology, Arlington State
College, Arlington, Texas. Manuscript received 3 August 1963.-ED.

Hellier, Thomas R., Jr. 1966. The Fishes of the Santa Fe River System. Bull.
Florida State Mus., vol. 11, no. 1, pp. 1-46.


The Santa Fe River, one of the three major tributaries of the
Suwannee, is unique in that it is the only river in Florida that leaves
its surface channel and flows underground for a considerable distance
before reappearing on the surface. The underground channel of
the Santa Fe measures more than 2 miles in length, but can be
traced with ease on aerial photographs which mark its subterranean
course by a series of sink holes separated by intervening natural
bridges (Stubbs, 1941). Many other streams in the limestone regions
of Florida disappear underground, but they either reappear within a
few hundred yards, or do not return to the surface as the same
stream. The long underground section of the Santa Fe divides the
river into two distinct portions and forms a possible barrier to up-
stream migration of some fishes.
Although a number of fish population surveys have been made
in North America, none has dealt with a clear-water river like the
Santa Fe, in which direct observations of the fishes can be made in
the main channel. Furthermore, none has been made on a river that
is divided into two portions by a long subterranean channel. In the
Southeast the fishes of only two rivers have been reported on with
any degree of thoroughness. The Escambia in western Florida was
the object of an investigation by Bailey, et al., (1954) and the St.
Johns was studied over a 10-year period by McLane (1955). Neither
of these studies, however, were in places that afforded a consistent
opportunity for direct observation of the fishes, or that presented a
possible physical barrier to the ascent of marine invaders.
The present paper with modifications was submitted as a masters
thesis to the University of Florida in August, 1957. I wish to thank
my advisory committee, John D. Kilby, Chairman, E. S. Ford, and
J. N. Yount for guidance during the study. I am indebted to the
many students and faculty who assisted in the field work and pro-
vided advice, particularly David K. Caldwell and William L. Jen-
nings. J. C. Briggs, and Clark Hubbs were especially helpful in
suggesting revisions of the manuscript. My wife, Evelyn, provided
encouragement and assistance in all facets of the work.

The Santa Fe River drains the central portion of the upper pen-
insula of Florida (Fig. 1). It arises in the northeast corner of Alachua
County, Florida, from Lake Santa Fe, a typical, oligotrophic, sand-


hill lake bedded in sands of Pleistocene origin. From Lake Santa Fe
the river follows the northern Alachua County line westward for
approximately 40 miles to O'Leno State Park where it disappears
underground through the Santa Fe Sink. The major tributaries of
this section of the river are the New River, which forms the boundary
between Bradford and Union Counties, and Olustee Creek, which
separates Union and Columbia Counties. The watershed area for
the Santa Fe above Santa Fe Sink includes the northern portion of
Alachua, all of Bradford and Union, southwestern Baker and south-
eastern Columbia Counties. This area is principally agricultural land
and pine flatwoods underlain by the Miocene Hawthorne formation,
whose imperviousness encourages drainage into the Santa Fe through
small surface streams.
The river itself after leaving Lake Santa Fe flows through Pleisto-
cene sands, which lie on top of the Hawthorne formation, until it
reaches a point just west of Station I (see Fig. 1). As might be ex-
pected, the depth of the river in this section varies considerably with
the amount of rainfall, and often during periods of dry weather no
water flows in the river bed for several miles below Lake Santa Fe.
The aquatic plants found in this part of the river are generally very
sparse and consist principally of figwort (Hydrotrida caroliniana),
spikerush (Eleocharis), water-purslane (Ludwigia), water-pennywort
(Hydrocotyle umbellata), aquatic moss (Fontinalis), maidencane (Pan-
icum hemitomon), and some floating filamentous algae. Several small
streams collect rainwater from the surrounding flatwoods and empty
into this portion of the Santa Fe; the water in them is generally more
brownish than that of the Santa Fe proper and their emergent veg-
etation is more extensive.
Just downstream from Station I the Santa Fe cuts down into the
Hawthorne as evidenced by outcroppings of this formation along the
banks, particularly at the mouth of Olustee Creek. The character
of the river changes considerably from this point on to the Santa Fe
Sink. It becomes wider and slightly deeper with occasional rapids.
The current in the main river channel increases, but more backwater
reaches become evident and the aquatic plants change. Tape-grass
(Vallisneria), naiad (Najas), muskgrass (Chara) and much attached
filamentous algae dominate the areas with current, while cattail
(Typha) and spatterdock (Nuphar) are found along the banks in areas
of lessened flow. In the shallow rapids the rocks are covered with
aquatic moss (Fontinalis), and some figwort (Hydrotrida caroliniana)
groWs in sandy patches between the rocks. The bottom of this por-





--. -.L




Fig. 1.-The Santa Fe River area of Florida. The dotted line indicates
the boundary of the drainage basin and the eight numbered circles show the
locations of the regular collecting stations.

Vol. 11


tion of the river is still generally sand. The vegetation along the
river from Lake Santa Fe to Santa Fe Sink is of a mixed river swamp
and low hammock type with bald cypress (Taxodium distichum),
willow (Salix longipes), river birch (Betula nigra), water oak (Quercus
nigra), white bay (Magnolia virginiana), titi (Cyrilla racemiflora),
black gum (Nyssa sylvatica), and sweet gum (Liquidamber styraciflua)
as the dominant trees. Where the river cuts through sand bluffs the
mixed river swamp and low hammock type vegetation along its banks
is replaced by live oak (Quercus virginiana), slash pine (Pinus elli-
ottii), and occasionally longleaf pine (Pinus palustris). The Santa
Fe River east (upstream) of Santa Fe Sink will be referred to as the
area "above the natural bridge."
After disappearing into the sink, which penetrates the pervious
Ocala Limestone of Eocene origin, the Santa Fe follows an under-
ground course southwesterly for two miles and then reappears as a
surface stream. Thereafter it follows the northern boundaries of
Alachua and Gilchrist Counties as a surface stream for approximately
30 miles to enter the Suwannee River 9 miles southeast of the town
of Branford, Florida. This confluence is about 70 miles upstream
from the mouth of the Suwannee on the Gulf of Mexico.
The Santa Fe west of (downstream from) the natural bridge re-
ceives most of its drainage from a series of limestone springs. Some
of the larger springs emptying into this part of the river are Poe and
Lily Springs in Alachua County, Blue Springs in Cilchrist County,
and Ichetucknee Springs in Columbia County. The run of Ichetuck-
nee Springs receives the water of several smaller springs, and its
average flow of 335 cubic feet per second is the third largest for
springs in Florida (Ferguson, et al., 1947). The pervious Ocala Lime-
stone that underlies this last section of the Santa Fe accounts for the
large amount of subterranean drainage into the river. This portion
of the Santa Fe receives very little surface drainage except during
periods of heavy rain. The principal aquatic plants that occur here
are: Tape-grass (Vallisneria), naiad (Naias), muskgrass (Chara), water.
purslane (Ludwigia), Sagittaria with narrow strapshaped leaves, aqua-
tic moss (Fontinalis), and filamentous algae. Coontail (Ceratophyl-
lum demersum) is locally abundant in some of the larger springs and
spring runs. The vegetation along the river on its western end is
also of a mixed river swamp and low hammock type with the same
species of trees dominating as listed before, with the addition of the
oveicup oak (Quercus lyrata) which occurs sporadically in several
places. (The aquatic plants are after Eyles and Robertson (1944) and

1 fnf>

_~~__ ~I___^ ^_ _I__ __ ___ __


the trees are after West and Arnold (1956). The above species of
plants are referred to in the remainder of this paper by generic name
only.) Downstream from the natural bridge the current is generally
swift, the water clear except after heavy rains, and the bottom is
usually of sand on rock or bare rock. The Santa Fe west of the
natural bridge will be referred to as "below the natural bridge."
Odum (1953) lists the Santa Fe as one of the rivers with the
lowest chlorinity in Florida. The chloride content is ten parts per
million or less both above and below the natural bridge. Tempera-
tures are generally higher in the summer and lower in the winter
above the natural bridge than those below it (Table 1).


Above Below
Months Natural Bridge Natural Bridge

January 20.5 22.0
February 17.5 20.5
April 22.0 22.0
May 25.0 25.0
June 27.0 25.5
July 26.5 25.0
August 29.0 25.0
September 28.0 24.0
O-.tober 25.0 23.5
November 24.5 23.5
December 21.0 22.5


The collecting stations shown on Figure 1 were selected to in-
clude the major habitats occurring in the Santa Fe River. The reg-
ular stations included locations above and below the natural bridge
that were as nearly comparable to each other as possible.

STATION 1. This station is at Cox Bridge where State Highway 241
crosses the Santa Fe. Here the river is a shallow, sand-bottom stream
with a moderate current and sparse aquatic vegetation. Hydrotrida
and Eleocharis are the dominant plant genera and cover approxi-
mately 10 per cent of the bottom in small mats. An aquatic moss
Fontinalis covers some of the logs and stones'in the river bed. These

Vol. 11


three species are the principal submerged plants; the only emergent
plant present in any significant amount is Ludwigia. The depth of
the water varies considerably; no water was flowing in April 1956,
but by July the river covered its flood plain at Cox Bridge to a depth
of approximately 3 feet.

~;;p~~s~= ~:-~....~

Fig. 2-Station 1. Santa Fe River at Florida State Highway 241. The
water is about two feet deep, slightly above average for the period covered by
this study.

STATION 2. The Santa Fe River flows over a series of rapids in
O'Leno State Park. The last of these rapids, about 250 yards up-
stream from the Santa Fe Sink, was selected as Station 2. Here the
rapids extend from bank to bank across the river, which is about 12
yards wide. During the present study the depth at this station varied
from a few inches to about 3 feet according to rainfall. The heaviest
rains and the deepest water occurred in summer. The vegetation is
principally of the submerged type; Najas, Vallisneria and Chara
occur in patches over the rapids, Fontinalis and several green algae
grow on many of the submerged stones covering the bottom of the



STATION 3. In O'Leno State Park 20 yards downstream from Sta-
tion 2, a slough about 20 yards wide extends approximately 50 yards
northeasterly from the Santa Fe; its average depth is 2% feet. Four
major types of aquatic vegetation occur here. The bottom is nearly
covered by large beds of Naias and Chara interspersed with smaller
patches of Vallisneria. The pleuston is made up of the duckweeds
Lemna minor and Spirodela polyrhiza, the aquatic fern Azolla caro-
liniana, and small patches of green algae, Spirogyra. The floating
leafed Nuphar and the emergent Typha occur around the margins of
the slough. The bottom of the slough is fine silt on sand except di-
rectly under the Nuphar where moderate deposits of mud cover the
rhizome systems of these plants.

'I.'~ ~

Fig. 3-Station 2. Santa Fe River at O'Leno State Park showing the rapids
approximately 250 yards upstream from Santa Fe Sink.

STATION 4. Santa Fe Sink is roughly circular in outline and about
70 yards in diameter. The depth, measured in the center in March
1956, was 105 feet. The sides of the sink slope off very rapidly and
the only aquatic plants present in any substantial quantities are the
pleuston forms Lemna, Azolla, and Spirodela which, together with

Vo. 11


floating logs, tree branches, leaves
the sink's entire surface.

and other debris, usually cover

Fig. 4-Station 3. Slough on the Santa Fe River at O'Leno State Park about
200 yards upstream from Santa Fe Sink. The water depth is approximately
three feet at the boat in the slough.

STATION 5. This station is mile upstream from Duncan's Landing,
Columbia County, at the mouth of a small unnamed spring on the
north bank of the river. The bottom is sand and the only vegetation
is a submerged moss, Fontinalis, attached to the few logs present.
The spring and its short run have large beds of Najas and Ludwigia
along the banks.
STATION 6. Station 6 is on the north bank of the Santa Fe River at
Duncan's Landing, which is 2M miles upstream from the confluence
of the Ichetucknce and the Santa Fe. The current is much reduced
by a curve in the river and by a rock outcrop from which two springs
flow. The bottom is silt on sand with an occasional spot of silt on
rock. The plants consist of large beds of Najas, Chara and Vallisne-
ria. Lemna and Spirogyra occur in small quantities and Ludwigia
grows close to and on the bank. The rocks just below the station are


Fig. 5-Station 4. Santa Fe Sink at O'Leno State Park. The material cov-
ering the surface is principally Lemna and assorted debris.

Fig. 6-Station 5. North bank of the Santa Fe River 3 miles upstream from
the confluence of the river and Ichetucknee Spring Run. A small unnamed spring
run is visible on the left.

Vol. 11


covered with Fontinalis and filamentous algae. Some Ceratophyl-
lum is attached to the rocks in the direct flow of the springs. The
depth of the water varies from about 5 feet on the river side to a
few inches at the shore.

Fig. 7-Station 6. Duncan's Landing on the Santa Fe River 2 miles up-
stream from the mouth of Ichetucknce Spring Run. A spring may be seen roiling
the surface of the water.

STATION 7. Downstream 30 yards from Duncan's Landing is a small,
nearly round depression in the river bed near the north bank of the
river. About 10 yards in diameter, the depression slopes off rapidly
to a depth of approximately 5% feet. Here current is insignificant
and the only vegetation is Vallisneria along the upstream edge of the
hole. The bottom and sides of this station are rock, except where the
depression is open to the river on the south side. Quantities of dead
plant debris float on the surface of this area.

STATION 8. This station is at a broad stretch of rapids 1 miles
above the mouth of the Ichetucknec. Throughout the study water
covered these rapids to an average depth of from 2 to 2V2 feet.
The bottom is rock with patches of sand in the depressions. Many


large beds of Najas, Sagittaria and Vallisneria occur throughout the
rapids. Small patches of Fontinalis grow on some of the submerged
stones and green, non-filamentous algae cover most of the remaining

Fig. 8-Station 7. A hole in the river bed just downstream from Duncan's
Landing on the Santa Fe River.

OTHER STATIONS. Many collections were made at other widely spaced
localities in the Santa Fe and its tributaries, but as these places
were not visited regularly, they are not numbered. Except those
along small, flatwoods streams and in one pond, these other sites
generally resembled one of the numbered stations.

Collections were made monthly at the eight regular stations and
at many other sites along the Santa Fe including representative habi-
tats throughout the drainage basin. Active field work was conducted
from June until October 1955, and from January 1956 through Jan-
uary 1957.
Most collections were made with either a 15-foot knit seine with
/4-inch mesh on a 4-foot drop and with a 4-foot bag in the center,

Vol. 11


or with an 8-foot common-sense seine also with /-inch mesh and a
4-foot drop. Other types of collecting gear occasionally used in-
cluded gill nets, wire traps, trotlines, hook and line, poison, spears,
and explosives. Underwater observations were made with a face
mask and scuba gear.

Fig. 9.-Station 8. A broad exposure of rapids in the Santa Fe River one
and one-half miles upstream from the junction of Icctucknee Spring Run with
the Santa Fe. The water depth at the boat is approximately 18 inches.

All specimens were preserved in the field in 10 per cent formalin,
transferred in the laboratory to 35 per cent isopropanol, and are
deposited in the University of Florida collections. Measurements
were taken in a straight line to the nearest millimeter, and all
length measurements given are standard lengths.
Water samples were analyzed for chlorinity in the laboratory by
the Mohr method of silver nitrate titration.


The following list of fishes from the Santa Fe River, Florida, en-
compasses 13 orders, 21 families and 60 species. Tables 4, 5, and 6
are a part of this list, and much of the data, such as specific catches
by months and by major habitats both above and below the natural
bridge, have been consolidated in the tables for ease of reference
and to free the annotated list of these details. The sequence of spe-
cies and the scientific and common specific names follow Bailey
et al. (1960).
1. Acipenser oxyrhynchus desotoi Vladykov. In the fish collec-
tions of the University of Florida is a specimen of the southern sea
sturgeon taken from the Santa Fe River at High Springs. No indi-
viduals of this species were seen or caught during the study.
2. Lepisosteus osseus (Linnaeus). The longnose gar was the
largest fish found in the river system. One specimen was collected
below the natural bridge in July, and three above it-one each in
July, September, and October. Single individuals of this species
were observed below the natural bridge on nine occasions in April,
May, and July. Invariably they were seen in midstream 1 to 3 feet
below the surface. One specimen 54 mm long was collected in July
1955 and one gravid female of 1,016 mm was taken in July 1956.
3. Lepisosteus platyrhincus DeKay. The Florida spotted gar is
the most abundant gar in peninsular Florida. It occurs in all major
freshwater and some brackish habitats, but populations are heaviest
in still waters with an abundance of submerged aquatic plants and
a rich food supply of small fish (McLane, 1955). Although plentiful
in Florida, only 3 individuals were observed in the Santa Fe during
the study. Two of these were above the natural bridge in Septem-
ber and one below in July.
4. Amia calva Linnaeus. The bowfin is a widespread but sel-
dom abundant fish of quiet waters, especially those with shallows,
some cover, and a silty bottom. Places that meet these requirements
are more plentiful above than below the natural bridge and, as ex-
pected, more individuals were seen above (8) than below it (1).
5. Alosa alabamae Jordan and Evermann. The only specimens
of this anadromous species found in the Santa Fe were 4 collected
at Station 8 in October 1956, which represent a range extension for
the species (Berry, 1964). John D. Kilby and J. C. Dickinson, Jr.

Vol. 11


(personal communication) collected several dozen Alabama shad
from the Santa Fe near Station 8 also in October 1956, but unfor-
tunately the specimens were lost. Thus the species may be locally
abundant in the Santa Fe at certain times.
6. Umbra pygmaea (DeKay). McLane (1955:63) records two
eastern mudminnows in the University of Florida collections taken
in the Santa Fe drainage at Ocean Pond near Olustee, Baker County.
The species is rare in Florida, and none were taken during the pres-
ent study.
7. Esox americanus Gmelin. During the study two redfin pick-
erel were taken above the natural bridge in small, acid tributaries
of the Santa Fe. One of these caught in September was a 175 mm
female containing ripe ova.
8. Esox niger Lesueur. The only chain pickerel taken was a 58
mm individual caught in April 1956 at Station 1.
9. Hybopsis harper (Fowler). The redeye chub is the most
abundant vertebrate inhabitant of subterranean waters in Florida
and occurs most frequently in springs and their surface runs. Second
only to Lucania goodei in numbers collected below the natural bridge
(1,082 specimens), we collected none above it. The fact that the
lower river is fed principally by subterranean streams and the upper
portion principally by surface drainage may explain this distribu-
tion. It is significant that 39.6 per cent of the fish were taken at
Station 7 where a large spring enters the river and another 29.8 per
cent from springs or spring runs. Gravid females were caught in
January, February, May, June and July. The only specimen under
25 mm collected was a 20 mm fish taken in February.
10. Notemigonus crysoleucas (Mitchill). We took 425 golden
shiners above the natural bridge and only 9 below it. Though we
found them throughout the year and in all habitats examined, 94.3
per cent of the specimens came from under patches of floating
Nuphar at Station 3. We collected few fingerling or smaller indi-
iiduals, but took one gravid female of 165 mm in June 1956.
11. Notropis chalybaeus (Cope). As McLane (1955) emphasizes,
the ironcolored shiner occurs most regularly in small hammock
streams that drain pine flatwoods. In the Santa Fe the species is
not abundant. All but 2 of the 30 specimens collected were from
small flatwoods streams tributary to the river above the natural



bridge, and all were found in association with aquatic vegetation.
Males with breeding tubercles were taken in June and July, well
within the mid-April to early September breeding season Marshall
(1947) reports for the species in Alachua County.
12. Notropis hypselopterus (Giinther). Over 99 per cent of the
123 sailfin shiners collected were from the headwaters of the Santa
Fe where the river has many of the characteristics of a flatwoods
stream, including much sandy bottom and a moderate current. Here
a collection in September 1955 yielded 65 specimens at a point where
the river is almost completely shaded by the hammock type vegeta-
tion along the bank. We collected only one specimen below the
natural bridge, but in the University of Florida fish collections are
several from springs tributary to this part of the river. Collections
from July and September contained tuberculate males and gravid
females. The single specimen collected in August was a tuberculate
male of 39 mm. No sailfin shiners were collected from October
through June.
13. Notropis maculatus (Hay). I found the red minnow only
above the natural bridge and took all the 15 specimens from unveg-
etated sandy or silt)' bottomed still water habitats. No one collec-
tion contained more than a few individuals; sizes ranged from a
minimum of 23 mm in September to a maximum of 45 mm in July,
and gravid females were noted in July and in September.
14. Notropis petersoni Fowler. The coastal shiner is by far the
most widespread cyprinid in the Santa Fe, occurring in all types of
habitats above and below the natural bridge. This species was most
abundant at Station 7 (28.7 per cent) where it occurred in close as-
sociation with aquatic vegetation. Observations made with scuba
gear indicate that it also occurs commonly in midstream around
patches of Najas, Vallisneria, and Chara as well as over small sandy
areas in water up to 20 feet deep. We found this cyprinid frequently
in association with harper and the two species are very
difficult to distinguish in the field. Tuberculate males and gravid
females collected in all months except March, June, and October in-
dicate year-round spawning in the Santa Fe.
15. Opsopoeodus emiliae Hay. The pugnose minnow is prin-
cipally a fish of the still shallows and it occurred both above and
below the natural bridge. One collection made in July and two in
August from backwaters above the natural bridge contained a total

Vol. 11


of 60 specimens ranging from 25 to 41 mm, and all contained gravid
females. In the University of Florida fish collections are 4 individ-
uals taken below the natural bridge at Poe Springs.
16. Erimyzon sucetta (Lacepede). All but one of the 17 chub-
suckers taken, 15 above the natural bridge and 2 below it, were
caught at stations having submerged or emergent vegetation. Sizes
ranged from a 22 mm individual caught in July to one of 165 mm
taken in June.
17. Minytrema melanops (Rafinesque). The spotted sucker seems
to be more of an open water form than E. sucetta, and the Suwannee
River drainage is its eastern limit in Florida (Carr and Goin, 1955).
We saw 7. 1 in January, 3 in July, and 3 in December, drifting or
swimming along the bottom of the river channel at depths up to
15 feet, and collected 8 specimens above the natural bridge and
5 below it. Two taken in Ichetucknee Spring Run during April
1956 were large males (420 and 380 mm) with breeding tubercles on
the snout and anal fins. One individual of 33 mm was caught in the
river one-half mile north of Louise, Alachua County, in July 1955.

18. Ictalurus catus (Linnaeus). Seventeen white catfish were
caught on trotlines, 2 below and 15 above the natural bridge; 14 of
the latter were taken at Station 4, which furnishes perhaps the best
catfish habitat in the Santa Fe. Catfish living in this sink have only
to wait for food to be washed downstream to them. Gravid females
were collected in June and February. McLane (1955) found young
in the St. Johns River during May, June, October, and December
which indicates a prolonged breeding season for the species. Our
largest catfish from the Santa Fe was 522 mm long and weighed
approximately 6 pounds.

19. Ictalurus natalis (Lesueur). 1. natalis was taken principally
in the same type of habitat as I. nebulosus. However, more yellow
cats were caught in small tributary streams (38 per cent) than were
bullheads (14 per cent). Our 29 specimens ranged from 16 mm long
in September to 156 mm in August.

20. Ictalurus nebulosus marmoratus (Lesueur). We found the
brown bullhead principally in heavily vegetated areas and around
debris. I saw this catfish several times lying under large beds of
Vallisneria and the masses of debris on the upstream side of tree trunks
in the river. The four specimens we caught on trotlines in midstream


over a sand bottom at night show that I. nebulosus leaves its con-
cealment to forage after dark.
21. Ictalurus punctatus (Rafinesque). Although no channel cat
was taken or observed during the present study, Coleman J. Goin
informs me that he has caught this species in the Santa Fe River.
The University of Florida collections have specimens from Fannin
Springs on the Suwannee, but none from the Santa Fe.
22. Ictalurus sp. This is an undescribed species to be named
elsewhere by Yerger and Rclyea. Yerger (personal communication)
states that it is more closely related to Ictalurus catus than to the
flatheads I. platycephalus and I. brunneus. We caught 27 specimens
below the natural bridge ranging in size from 36 mm in December
to 155 mm in April. We took none above the natural bridge.
23. Noturus gyrinus Mitchill. The tadpole madtom is the most
common representative of its family in the Santa Fe, occurring in
practically every major habitat above and below the natural bridge.
However, it is most plentiful inside large beds of Najas along the
margins of the river below the natural bridge. We collected gravid
females in all months except January, March, May, October, and
November and found individuals under 15 mm long in April, June,
August, and September. Each of 70 specimens examined was para-
sitized by Acanthocephala. The largest specimen caught was a
heavily parasitized female 79 mm long.
24. Noturus leptacanthus (Jordan). The speckled madtom is re-
stricted almost entirely to streams having appreciable currents, some
sandy or rocky bottom, and cover in the form of rocks, logs, or other
debris. Relatively common in some parts of the Santa Fe, of 132
specimens collected, 75.8 per cent were from the rapids at Stations
2 and 8. Gravid females were found in April, June, July, August,
and September. Specimens under 20 mm were collected in August
and September; no specimens were taken in March or May. Of 50
fish examined, 32 had Acanthocephala parasites.
25. Anguilla rostrata (Lesueur). I caught one specimen of this
well known catadromous eel above the natural bridge and 12 be-
low it. The smallest was one 122 mm long taken in December. The
digestive tracts showed crayfish remains in all specimens collected.
26. Strongylura marina (Walbaum). The needlefish maintains
breeding populations both in coastal marine and in the fresh waters

Vol. 11


of Florida, and is thus completely euryhaline. It is a surface form
usually found where little or no current flows, but in spring runs
and in some rivers it feeds actively over bars where currents are
sometimes swift. Found only below the natural bridge, needlefish
were seen most often swimming at the surface in small schools of
up to 8 or 10 individuals. Only 3 specimens were collected. One
was taken at dusk in a common sense seine over a bed of Vallisneria,
and two were taken at night in a 15-foot bag seine close to the
springs at Station 6. Sight records indicate the species is more
abundant in the Santa Fe during the spring and summer than at
other seasons.
27. Fundulus chrysotus (Ginther). One golden topminnow was
collected below the natural bridge and 54 above it. Fish in breed-
ing condition were found in July, August and September.
28. Fundulus cingulatus Valenciennes. Four collections, one in
June, two in August, and one in September, all from above the
natural bridge, contained a total of 35 banded topminnows. These
fish were found on the surface in shallow marginal water over a
sandy bottom and in close association with emergent vegetation.
August and September collections contained adults in breeding con-
29. Fundulus notti (Agassiz). Except for its occurrence in quiet
waters of calcareous streams, this species is practically restricted to
clear sandhill lakes. Six of the collections above the natural bridge
yielded a total of 22 specimens, and two collections below it one
specimen each. As elsewhere the starhead topminnow in the Santa
Fe is a surface water form. When pursued with nets it sometimes
retreats into stands of emergent vegetation, but I have never seen
it seek the protection of deep water. The largest and smallest speci-
mens taken were both from July collections, and measured 74 and
16 mm respectively. Gravid females were found June and July.
30. Fundulus seminolis Girard. The seminole killifish we found
only below the natural bridge, where 188 specimens were collected.
Sizes ranged from 18 to 122 mm. The most common habitat of this
species in the river was over sandy bottom around patches of vege-
tation, principally Vallisneria and Najas. Unlike most other species
of Fundulus, this killifish was never seen on the surface for any length
of time, but usually from near the bottom up to half way to the
surface. Again unlike most other fresh water Fundulus in Florida,


F. seminolis frequently occurs in loosely organized schools of from
30 to 40 individuals, particularly in the spring and early summer.
Gravid females were taken in April only. Ova examined in one
female varied from approximately 0.75 to 2.0 mm in diameter. Of
50 specimens examined, 45 were found parasitized by Acanthoceph-

31. Jordanella floridae Goode and Bean. The flagfish is most
abundant in Florida in roadside ditches and in similar natural situ-
ations. I collected a single specimen 22 mm long in June 1955 from
shallow water over a sandy bottom below the natural bridge.

32. Leptolucania ommata (Jordan). The pygmy killifish inhabits
shallow, quiet waters sufficiently vegetated to provide either surface
or bottom cover. I took 8 of these fish from the Santa Fe, all from
one small flatwoods stream near the headwaters. The fish were near
the bank in shallows with Hydrotrida, Ludwigia, and Sphagnum
on the bottom. Three gravid females were found in a July collection
and the smallest individual (13 mm) was taken in September.

33. Lucania goodei Jordan. The bluefin killifish was by far
the most common cyprinidontid encountered, with 560 specimens
collected above the natural bridge and 1,343 below it. Although
found in every habitat, it occurred most frequently in and around
beds of Najas, Chara, and Vallisneria. These fish, together with
Syngnathus scovelli and Noturus gyrinus, were often found deep in
the center of the beds of vegetation. Females with ripe ova were
taken in every month and fish under 20 mm in all months except
January and May. One specimen collected in April was heavily
parasitized by metacercaria of the fluke Clinostomum marginatum.

34. Gambusia affinis holbrooki Girard. The mosquitofish was one
of the more abundant fish in the river with 966 specimens collected
above and 400 below the natural bridge. A shallow-water, marginal
dweller found generally in association with emergent vegetation and
filamentous algae, the species may also be found in small tributary
streams, springs, and flood plain ponds where it is often very numer-
ous. One such small pond when poisoned yielded only G. affinis
holbrooki, Elassoma zonatum and Heterandria formosa. Females
containing eyed embryos were collected in every month, but a much
lower percentage of these ripe females was'found from November
through February than during the rest of the year. A large disparity

Vot. 11


occurs in the sex ratio of this species in the Santa Fe; of 500 indi-
viduals examined 78.6 per cent were females.

35. Heterandria formosa Agassiz. The least killifish is rather
widely distributed in the Santa Fe; 97 specimens were taken below
and 177 above the natural bridge. It not only occupies the same
type of habitat as Gambusia affinis holbrooki but its breeding season
is also similar. The largest specimen collected was a female 30
mm long.

36. Mollienesia latipinna Lesucur. The sailfin mollie is eury-
haline in Florida and has breeding populations in coastal, estuarine,
and strictly fresh waters far inland. Maximum populations occur in
shallow, marginal situations, especially those with vegetative cover
or other type of protection from predation. In the Santa Fe, it is a
shallow water, marginal form found almost exclusively in small
backwaters with little or no current, some filamentous algae and
small amounts of Lemna on the surface, and a bottom of sand or
rock overlain by silt. We collected 721 specimens below the natural
bridge and none above it. Mollies have a prolonged breeding season.
Ova were present in February specimens and females containing eyed
embryos were found from April through September. No March
collections contained Mollienesia. A disproportionate sex ratio ex-
isted; of 709 specimens examined 72.2 per cent were females. Al-
though these fish were normally found near or at the surface, they
invariably retreated toward the bottom when disturbed.

37. Syngnathus scovelli (Evermann and Kendall). The gulf pipe-
fish is another of the few completely euryhaline fishes in Florida,
maintaining breeding populations in strictly fresh water as well as
in the highly saline waters along the coast. In fresh waters it is
usually found in the densely vegetated littoral zone of clear streams
and rivers. We collected no pipefish above the natural bridge, but
took 108 below it in heavy beds of vegetation, principally Najas, along
the margins of the river and in the spring runs. Gravid females oc-
curred in samples from every month except March, June, October,
and December. Males with embryos in the brood pouch were taken
from April through September and in January and February. A
count of the embryos in the pouches of 19 males gave an average of
12.2 per pouch, which seems abnormally low. McLane (1955) re-
ported an average of 29.5 per pouch in 66 males from the St. Johns
River, and 44.7 per pouch in 48 males from Cedar Key, Florida.



S. scovelli probably breeds throughout the year in the Santa Fe
River as it does in the St. Johns.
38. Aphredoderus sayanus (Cilliams). Pirate perch were well dis-
tributed and associated with aquatic vegetation of all types. In
October 1956 we collected 53 specimens with poison over a clean
sand bottom at Station 8; the fish were in a hole approximately 35
feet deep and relatively free of current. During some seven hours
of underwater observations at all seasons in this hole with scuba
gear no Aphredoderus were seen; possibly they were hidden in small
concavities and crevices along the sheer rock walls of the hole.
Very few pirate perch were collected with seines anywhere, but
poisoning the same areas with rotenone often produced large num-
bers. These fish, together with Noturus gyrinus, Noturus leptacan-
thus, and the gars, were the last species the poison affected. Females
with ripe eggs were found in February, April, and December. The
length-frequency data in Table 2 indicate a late winter breeding sea-
son. Over 95 per cent of the fish examined were heavily parasitized
by Acanthocephala; 14 specimens also had small leeches attached to
the membranes of the caudal and dorsal fins.
39. Centrarchus macropterus (Lacepede). I collected one flier
64 mm in length in September 1956, from the Santa Fe north of
Louise, Alachua County. The river at this point resembles a small
flatwoods stream with a sand bottom and moderate current. An-
other Santa Fe specimen in the University of Florida collections
was taken approximately 2 miles upstream from Louise.
40. Chaenobryttus gulosus (Cuvier). The warmouth was most
abundant above the natural bridge where 50 of the 56 specimens
were collected, ranging in length from 16 mm to 140 mm, both
taken in August. The habitat preference of Chaenobryttus is mark-
edly restricted in the Santa Fe; 92.3 per cent of the specimens were
taken from heavily vegetated still waters over either a mud or silt
bottom. One collection containing 20 specimens was taken from a
small flood plain pond 4 by 5 yards in extent and 2 feet deep next to
the river at Worthington Springs, Union County. This pond was
completely devoid of vegetation, and had a mud bottom. Judging
from the surrounding terrain it is flooded periodically, which might
account for the presence of warmouth.
41. Elassoma evergladei Jordan. In the University of Florida
collections are seven specimens of the Everglades pygmy sunfish

Vol. I


'- Om-i m1 c VD C
- -

e1 t- 0 01

Ci 10 1 0n C~ '" 10 -

SmO l -

- C

1 3 QC 001 01 -

- 01

- 01 -

- CO V -

=0I 0I 0 in co C IC C IC m IC C 10 C=i
0C Ct' CO c? I 1 C t-- m01 0 c

-1101~ 01CCC '''1 1 0 CCC1-1 1


taken in Hatchet Creek, 10 miles north of Gainesville and 9 miles
west of Waldo, and one taken in the Santa Fe 7 miles northwest
of Waldo by Carter R. Gilbert in November 1961. I did not find the
species during my study.
42. Elassoma okefenokee Bohlke. Bohlke (1956) states this spe-
cies lives under overhanging plants in slow-moving, acid streams, in
essentially the same habitat as Leptolucania ommata. Of 66 fish
collected, 32 were from heavily vegetated areas in slow-moving trib-
utary streams and 33 were collected at Station 3. Only one specimen
was taken below the natural bridge. The Okefenokee pygmy sun-
fish was collected only in March, July, September, and November.
The smallest fish (10 mm) was taken in September, the largest (24
mm) in November. Gravid females were found in July.
43. Elassoma zonatum Jordan. Most of the banded pygmy sun-
fish (77.8 per cent) were collected from heavily vegetated areas be-
low the natural bridge. These fish were usually found in beds of
Vallisneria, Najas, and Chara, occasionally in patches of Ludwigia.
The specimens ranged in length from 29 mm in November to 15 mm
in July. Females with fully developed ova were taken in February
and April. Of the 45 fish collected, 25 had approximately 20 Acan-
thocephala in the coelomic cavity per fish.

44. Enneacanthus glorious (Holbrook). All the 172 specimens
collected were from above the natural bridge. The smallest indi-
vidual was 14 mm long, taken in July, and the largest 51 mm, was
caught in September. We collected 57 specimens in a vegetation-
free flood plain pond with mud bottom near Worthington Springs.
Gravid females were found in August and September.

45. Enneacanthus obesus (Girard). The University of Florida
collections have one specimen of the banded sunfish Carter R. Gilbert
collected in Hatchet Creek 10 miles north of Gainesville and 9 miles
west of Waldo in April 1961. I did not find the species during my

46. Lepomis auritus (Linnaeus). The redbreast is by far the
commonest centrarchid in the Santa Fe. Although we found it in
all types of habitats, it was most abundant in sand bottom creeks
(41.5 per cent) and in heavily vegetated sloughs (37.4 per cent). The
young preferred shallow vegetated zones ih still water but were
also found in heavily vegetated tributary streams. Gravid females

Vol. 11


were collected in April, June, and July. From June through Janu-
ary we took 59 specimens under 20 mm, 51 of them in June, July,
and August. Redbreasts were seen guarding redds in April. Though
this fish has a prolonged spawning season in the Santa Fe, a pro-
nounced breeding peak occurs in late spring and early summer.

47. Lepomis macrochirus purpurescens Cope. The young and
adult bluegills show decided differences in habitat preference in the
Santa Fe. Young specimens under 50 mm were taken principally in
heavily vegetated still waters. Large individuals were collected and
observed in more open waters. Seven adult bluegills were taken at
Station 8 in July, and one collection in August made by dynamiting
under a large fallen tree yielded 10 adult specimens. The presence
of fish under 20 mm in collections from July and November indicates
a prolonged breeding season. The largest specimen was taken in
July and measured 163 mm.

48. Lepomis marginatus (Holbrook). Carter R. Gilbert collected
24 dollar sunfish in the Santa Fe 7 miles northwest of Waldo and 2
in Hatchet Creek 9 miles west of Waldo in November 1961. I did
not take the species during my study.

49. Lepomis microlophus (Gunther). The redear sunfish is pri-
marily a fish of the open waters of larger rivers, springs and lakes.
The 244 specimens collected, 211 from above and 33 from below the
natural bridge, ranged in size from 16 mm in June, July and Novem-
ber to 220 mm in December. The relative numbers of fish caught
above and below the natural bridge may be misleading because 163
were taken in five unusually large collections at Station 3. Under-
water observations below the natural bridge showed redear sunfish
much more plentiful than the 33 fish collected there would indicate.
Adult redears are by far the most common sunfish in the deeper,
unvegetated areas at midstream, so numerous in fact that it was im-
possible to keep exact records of them while swimming downstream on
prolonged observations. Short samplings made by swimming across
the river and back gave some definite figures on redear abundance;
in each of three such short trips at Station 7 in July, November, and
April, we counted no less than two and as many as seven large red-
ear sunfish near the middle of the river. These large fish are diffi-
cult to collect. When we tried to spear some with an underwater
gun' they proved so wary of the diver that we took only one speci-
men" in this manner. Young redears were found most frequently in



vegetated shallow waters and were relatively easy to collect with
a seine.
McLane (1955) reports the peak spawning season for the St. Johns
is in June and July. The 16 mm specimens in our collections from
June, July, and November suggest some spawning probably occurs
earlier and later in the Santa Fe.
50. Lepomis punctatus punctatus (Valenciennes). This was the
only member of its genus collected in greater numbers below (233)
than above (123) the natural bridge. We found the spotted sunfish
in all types of habitats in the Santa Fe, but it was most abundant in
some of the deep holes below the natural bridge, where 42 per cent
of the specimens were collected. Fishing with a hook and line with
bread for bait at Blue Springs, Gilchrist County, in January 1956,
I caught both bluegills and spotted sunfish on the sandy bottom of
the boil area, but when I threw my hook over the edge of the rock
crevice from which the spring issues, I caught only spotted sunfish.
The spotted sunfish was also common in and around dense beds of
aquatic vegetation at Stations 3 and 4 (41.4 per cent). No large num-
bers of small L. punctatus were ever taken in a single collection, as
often happened with other sunfish. Carr (1947) found that small
spotted sunfish tend to disperse soon after leaving the redd. Females
with ripe ova were collected in February and from April through
September. Caldwell et al. (1957) report that this species breeds
throughout the year in Silver Springs, Marion County, Florida.
Specimens under 20 mm in length were collected in June, August,
and September, the smallest (14 mm) in August. The largest speci-
men was one measuring 232 mm taken in June.
51. Micropterus notius Bailey and Hubbs. The Suwannee bass,
a relict form in northern Florida, is the most generalized species in
the genus Micropterus (Bailey and Hubbs, 1949). Known only from
swift-flowing rivers in Florida, it is represented by relatively few
specimens in collections. During this study we collected 223 speci-
mens and observed several hundred others during diving operations.
Although we took M. notius and M. salmoides floridanus regularly
in the same seine hauls, the two species occurred most frequently in
distinctly different habitats. Underwater observations and our catch-
es in the Santa Fe showed AM. notius to be essentially a fish of rapid
waters and M. salmoides floridanus one of quieter reaches. In
rapids we caught 21 per cent of the total number of M. notius and
only 1.2 per cent of M. salmoides floridanus. Above the natural

Vol. 11


ro ~ ci

0] cc~c c

- E W -

ci 4N,-I

1 x in NCI C1 r0 1

i-( C I- -

= -

1 0 Ci ~

T i (1c -

C O'C cl C C C C C

Sc c0000 0 0
i- 0] C1i 10 cc -.D lIC CT C



*- ( 10

no M s
-^ MC -


bridge where rapids are not a dominant feature, only 58 M. notius
were present in catches that produced 334 M. salmoides floridanus.
Below the natural bridge where rapids and swift channels occur fre-
quently, the ratio was 164 notius to 107 salmoides. The length-fre-
quency data in Table 3 strongly indicate a spring spawning season.
In April 1956 one individual was observed guarding a redd con-
structed on a clear sand bottom in approximately 2 feet of water.
The redd itself was indistinguishable from those constructed by
M. salmoides floridamns.

52. Micropterus salmoides floridanus (Lesueur). This bass and
M. notius are the most common large predatory fish in the Santa Fe.
Adult largemouth bass occur in every major habitat but are most
abundant where submerged trees and other debris provide cover.
Bass were found in every series of observations we made, at all
depths and in currents of every magnitude in the river. Small in-
dividuals were most often collected and observed in sand bottom
shallows and in sparse patches of Ludwigia, Vallisneria, Najas, and
Chara. Specimens under 25 mm were taken from April through Au-
gust and one gravid female 208 mm long, was collected in April.
Bass were observed guarding redds in April and June at Station 1
and immediately upstream from Station 2. Carr (1942) describes the
construction of redds for this species. The ones observed in this
study were on a clear sand bottom free of vegetation in 2 to 3 feet
of water.

53. Pomoxis nigromaculatus (Lesueur). Black crappie are not
common in the Santa Fc. Six specimens taken above the natural
bridge were the only ones seen.

54. Etheostoma edwini (Hubbs and Cannon). This darter, as its
describers (1935) note, frequents swift, clear, vegetated streams. Of
the 54 brown darters we collected, 42 per cent were from rapids.
We took 68.5 per cent of the total below the natural bridge, where
the most suitable habitat exists. As females with ripe ova were
taken in January, February, April, June, July, September, and De-
cember, these fish evidently breed throughout the year in the
Santa Fe.

55. Etheostoma fusiforme barratti (Holbrook). The 38 swamp
darters we collected, 29 above and 9 below the natural bridge, ranged
in size from 48 mm in July to 26 mm in August. No collections con-

Vol. 11


training this species were made in January, May, or October. Adults
with breeding tubercles and ripe gonads were collected in February
and March. Of the 38 fish examined 9 contained Acanthocephala
in the coelomic cavity.

56. Percina nigrofasciata (Agassiz). Although we collected 82
blackbanded darters above and 9 below the natural bridge, the rel-
ative numbers are not at all indicative of the species actual abun-
dance, as its major habitat below the natural bridge precludes suc-
cessful collecting. Principally bottom dwellers, these fish may be
seen plentifully in the current of the main river bed resting on the
bottom behind rocks and pieces of debris. We counted approximately
50 individuals while swimming once across the river and back at
Station 6 in July. Every longer sampling showed these darters
plentiful in their favored habitat. Above the natural bridge we
found the species principally in the shallow rapids at Station 2 (26.4
per cent) and on the sand bottom in the main current at Station 1
(33.0 per cent). The smallest specimen (24 mm) was collected in
April and the largest (73 mm) in September. Adults in breeding
condition were found in February and April and from July through
October. No collections made in January, March, May, November,
or December contained Percina.

57. Eucinostomus sp. David K. Caldwcll and I saw several
mojarras in April and May at Station 6. As our attempts to collect
these fish were unsuccessful, no positive identification could be made.
McLane (1955), Carr and Goin (1955), and Herald and Strickland
(1949) list Eucinostomus argenteus Baird and Girard as occurring in
Florida fresh water rivers, and most probably the fish we saw were
of this species.

58. Mugil cephalus Linnaeus. Although the striped mullet is prin-
cipally a fish of marine lagoons, estuaries, and other coastal shal-
lows, it frequently invades fresh waters. Many mullet were observed
below the natural bridge in all months except September, October,
and November; no mullet were taken or seen above it. These ma-
rine invaders may be present throughout the year, but they do not
become plentiful until early winter. They were common in the river
from February through August. Local fisherman catch large num-
bers on hook and line using small wads of Spirogyra or earthworms
for bait. No small specimens were taken or seen during this study,
and the adults collected showed little gonadal development.



59. Labidesthes sicculus vanhyningi Bean and Reid. The brook
silversides is distinctly a surface form and often occurs in small
schools. We found it very abundant above the natural bridge at
Stations 1 and 3 (76.9 per cent of the fish collected) and less so in the
small tributary streams (7.9 per cent of the total). It was not very
common below the natural bridge, but could be found there in
springs and a few other places where the current was weak. Over
95 per cent of the specimens were collected from areas of very
slowly flowing or still water. The species has an extended breeding
season in the Santa Fe. Females with ripe gonads were taken from
February through April and from June through September; no
specimens were collected in May. The smallest specimen collected
was one of 12 mm in June 1956. No other individuals under 20 mm
were collected.

60. Trinectes maculatus (Bloch and Schneider). We collected
143 hogchokers on clean sandy bottom below the natural bridge and
found none above it. Our fish ranged in length from 22 mm in May
to 65 mm in July. We collected specimens under 25 mm in April,
May, July, and November. We noted no gonadal development in
any of our specimens, and it is improbable that this fish spawns in
the Santa Fe. Mansueti and Pauly (1956) give 111 mm standard length
as the minimum size of sexually mature individuals in the Patuxent
River, Maryland.


The fish fauna of the Santa Fe is a diversified one characterized
by a large number of species of wide ecological differences. How-
ever, the bulk of this fauna is made up of relatively few species well
adapted to the principal types of habitat available in the river, re-
flecting the fact that a few habitats are quite extensive while many
others are restricted in extent.
Several marine species occur in the Santa Fe, where two of them
maintain breeding populations. In view of the extremely low salini-
ties (10 parts per million maximum) and the distance from the Gulf
of Mexico (approximately 70 miles), this marine invasion of euryha-
line fish is remarkable.
The natural bridge is a more effective barrier to upstream mi-
gration than to downstream. Many fish, fry, and eggs may be car-
ried under it by the current, but any upstream migration through
the underground channel must be active, not passive.

Vol. 11


The fish fauna of the Santa Fe may be divided into three major
groups on the basis of distribution above and below the natural
The first and largest of these three groups is the one including
those fishes that occur both above and below the natural bridge.
The species in this group may be further subdivided into those that
are principally stream forms, those that are found most often in still
water, and those that can adapt to either situation. Generally the
stream forms are found wherever the current is a significant feature.
Those that occur most often in streams are Lepisosteus osseus, Miny-
trema melanops, Noturus leptacanthus, Lepomis auritus, Lepomis
punctatus punctatus, Micropterus notius, Etheostoma edwini, and
Percina nigrofasciata.
Notropis hypselopterus, Leptolucania ommata, Elassoma okefeno-
kee, Elassoma zonatun and Enneacanthus glorious are also princi-
pally stream forms, but they are found most often in small streams
where the current is not so evident.
The still-water forms found both above and below the natural
bridge are Lepisosteus platyrhincus, Amia calva, Opsopoeodus emiliae,
Erimyzon sucetta, Fundulus chrysotus, Fundulus notti, Chaenobryt-
tus gulosus, Etheostoma fusiforme barratti, and Labidesthes sicculus
Those fishes found commonly in both lakes and streams are
Notemigonus crysoleucas, Notropis petersoni, Ictalurus catus, Ictal-
urus nebtdosus, Noturus gyrinus, Lucania goodei, Gambusia affinis
holbrooki, Heterandria formosa, Aphredoderus sayanus, Lepomis mac-
rochirus purpurescens, and Micropterus salmoides floridanus. One
catadromous species, Anguilla rostrata, occurs both above and below
the natural bridge, and is the only euryhaline form found above it.
The second major group of fishes includes the species found only
above the natural bridge. These fishes, all either lake or small
stream forms, are Esox americanus, Esox niger, Notropis chalybeus,
Notropis maculatus, Fundulus cingulatus, Centrarchus macropterus,
and Pomoxis nigromaculatus. All these species occur very infre-
quently even in suitable habitat above the natural bridge, and prob-
ably they also occur infrequently below it. However, during the
period covered by the study, the water level of the river was very
low and suitable habitat for these fishes was almost non-existent.
One other species recorded from above the natural bridge, Umbra


pygmaea, is known in Florida only from a few localities and is prob-
ably very rare in the Santa Fe drainage.
The last group includes the fishes found only below the natural
bridge. These can be subdivided into the primary freshwater forms
Hybopsis harper, Ictalurus sp., Fundulus seminolis, and JordaneUa
floridae; and the euryhaline Strongylura marina, Mollienesia latipinna,
Syngnathus scovelli, Mugil cephalus, and Trinectes maculatus.
Hybopsis harper, one of the few vertebrates known from the sub-
terranean waters of Florida, also occurs in surface waters with con-
siderable subterranean drainage, which is the general situation be-
low but not above the natural bridge. Ictalurus sp. is known in
Florida to inhabit deep, rocky holes in swift, clear water, a habitat
found in the Santa Fe only below the natural bridge. Possibly the
natural bridge acts as a barrier to the upstream migration of Fundu-
lus seminolis, which is not known from subterranean waters. Only
one specimen of Jordanella was taken during the entire study; prob-
ably this species, usually found in roadside ditches, strayed accident-
ally into the Santa Fe.
Of the euryhaline species, Alosa alabanae, the only anadromous
species found in the river, was seen only in October. Although at
times relatively abundant, this fish probably does not play a signifi-
cant role in the overall ecology of the Santa Fe. The sturgeon and
the mojarra are probably only accidental and sporadic visitors. Tri-
nectes maculatus also is probably only a visitor; it was found only
below the natural bridge where it was rather abundant, but no large
or very small specimens were encountered. Present knowledge in-
dicates the sea as the species' place of reproduction as well as the
habitation of most of the population. Though Mugil cephalus forms
a very conspicuous part of the Santa Fe fish fauna, as no evidence of
breeding can be found and the species apparently leaves the river
during the fall months, this mullet must also be called a euryhaline
Strongylura marina was found moderately plentiful, but no evi-
dence was obtained to indicate that it maintains a breeding popula-
tion in the river. As this species spawns in the St. Johns River sys-
tem and maintains heavy populations in some of the larger lakes in
that drainage (McLane, 1955) possibly the needlefish may spawn in
the Santa Fe.
Mollienesia latipinna and Syngnathus scovelli were abundant be-
low the natural bridge. That both species Ihaintain breeding popu-
lations in the river is proved by the presence in the collections of

Vol. 11


adults in breeding condition and of fry. No term is currently avail-
able to distinguish fishes that maintain breeding populations in both
fresh and marine waters from other euryhaline types. For such spe-
cies the writer proposes the term "dihaline".
Of the 60 species found in the Santa Fe, 43 species were taken
above and 42 species below the natural bridge. This practically
even distribution in species numbers suggests that approximately the
same number of ecological niches for fishes occur in each end of
the river.
Above the natural bridge 18 species made up approximately 90
per cent of the population as shown by numbers collected and per
cent of total catch (Table 4). Below the natural bridge 13 species
contributed over 90 per cent of the individuals collected (Table 5).
Four of the species concerned, Micropterus notius, Lepomis m. pur-
purescens, Lepomis microlophus and Percina nigrofasciata, were
actually more abundant than the percentages indicate. Repeated
direct observations confirmed that the catches did not reflect their
abundance accurately.
The difference between the upper and lower portions of the river
apparently is not in the number of different habitats available, but
in the relative extent of these habitats. Thus it is concluded that,
although both segments of the Santa Fe are about evenly endowed
with types of habitats, more of the habitats above the natural bridge
are more extensive than those below it, where a few principal hab-
itats are very extended and the remainder quite restricted.

The ichthyofauna of the Santa Fe may be divided into the
following groups according to their area of derivation as based on
known ranges given chiefly by Briggs (1958), and Carr and Goin
1. A northern element composed of fishes widespread to the
northward and reaching their southern limit in Florida. These spe-
cies are:
Lepisosteus osseus Notropis chalybeus
Amia calva Opsopoeodus emiliae
Umbra pygmaea Erimyzon sucetta
Esox americanus Minytrema melanops
Esox niger Ictalurus catus
Notemigonus crysoleucas Ictalurus natalis



Ictalurus nebulosus marmoratus
Ictalurus punctatus
Noturus gyrinus
Gambusia affinis holbrooki
Aphredoderus sayanus
Centrarchus macropterus
Chaenobryttus gulosus

2. A southeastern element compo
southeastern U.S. These are:
Lepisosteus platyrhincus
Notropis hypselopterus
Notropis maculatus
Notropis petersoni
Noturus leptacanthus
Fundulus chrysotus
Fundulus cingulatus
Fundulus notti
Heterandria formosa
Elassoma evergladei

3. A typically Floridian element
lowing three groups:

Elassoma zonatum
Enneacanthus glorious
Enneacanthus obesus
Lepomis auritus
Pomoxis nigromaculatus
Etheostoma fusiforme barratti

sed of fishes widespread in the

Elassonm okefenokee
Lepomis nacrochirus pur-
Lepomis marginatus
Lepomis microlophus
Lepomis punctatus punctatus
Etheostoma edwini
Percina nigrofasciata
Labidesthes sicculus

subdivided further into the fol-

a. Fishes typically Floridian but not exclusively so.
Hybopsis harper Jordanella floridae
Ictalurus sp.

b. Species endemic to Florida.

Fundulus seminolis
Leptolucania ommata

Lucania goodei
Micropterus salmoides floridanus

c. Species nearly endemic to the Santa Fe Drainage.
Micropterus notius

4. An element derived from marine waters.
Acipenser oxyrhynchus desotoi Mollienesia latipinna
Alosa alabamae Syngnathus scovelli
Anguilla rostrata Mugil cephalus
Strongylura marina Trinectes maculatus

Vol. 11


It is interesting to note that of the dominant species in the Santa
Fe, 9 are from the southeastern fauna, 7 from the northern, 4 from
the typically Floridian and 1 from marine waters.
The following species that were not collected during the present
study might be expected to occur in the Santa Fe on the basis of
their known ranges:

Dorosoma cepedianum Notropis texanus
Dorosoma petenense Notropis welaka
Notropis cummingsae Acantharchus pomotis
Notropis leesdi


Bailey, Reeve M., and Carl L. Hubbs
1949. The black basses (Micropterus) of Florida, with description of a new
species. Occasional papers, No. 516, Univ. Mich., Mus. Zool. 40 pp.,
2 pls.

Bailey, Reeve M., Howard E. Winn, and C. Lavett Smith
1954. Fishes from the Escambia River, Alabama and Florida, with ecological
and taxonomic notes. Proc. Acad. Nat. Sci. Phil., 106: 109-164.

Bailey, Reeve M., et al.
1960. A list of common and scientific names of fishes from the United States
and Canada. Am. Fish. Soc. Spec. Publ. 2: 1-102.

Berry, Frederick H.
1964. Review and emendation of: Family Clupeidae, pp. 257-454. By Sam-
uel F. Hildebrand, with emendations by others, and sections on Haren-
gula by Luis R. Rivas and on Dorosoma by Robert R. Miller. (In)
Fishes of the Western North Atlantic. Sears Foundation for Marine
Research, New Haven, Connecticut, Memoir I, Part 3. Copeia, 1964
(4): 720-730.

Bbhlke, James
1956. A new pygmy sunfish from southern Georgia. Notulae Naturae, Acad.
Nat. Sci. Phil., 294: 1-11.

Briggs, John C.
1958. Florida fishes and their distribution. Bull. Fla. State Mus. Biol. Sci.,
2(8): 223-318.

Caldwell, David K., Howard T. Odum, Thomas R. Ilellier, Jr., and
Frederick H. Berry
1957. Populations of spotted sunfish and Florida largemouth bass in a con-
stant-temperature spring. Trans. Amer. Fish. Soc., 85: 120-134.


Carr, Archie, and Coleman J. Goin
1955. Guide to the reptiles, amphibians and freshwater fishes of Florida.
University of Florida Press, Gainesville, Florida. 341 pp.
Carr, Marjorie H.
1942. The breeding habits, embryology and larval development of the large-
mouth black bass in Florida. Proc. New England Zool. Club, 20: 43-77.
1947. Notes on the breeding habits of the eastern stumpknocker, Lepomis
punctatus punctatus (Cuvier). Proc. Fla. Acad. Sci., 9(2): 101-106.
Chable, Alphonse C.
1947. A study of the food habits and ecological relationships of sunfishes of
northern Florida. (Unpubl. Master's Thesis, Univ. of Fla. Library).
Eyles, Don E., and J. Lynne Robertson, Jr.
1944. A guide and key to the aquatic plants of the southeastern United States.
Public Health Bull. 286.
Ferguson, G. E., C. W. Lingham, S. K. Love, and R. O. Vernon
1947. Springs of Florida. Fla. Geol. Sur. Bull., 31: 1-196.

Gunter, Gordon
1942. A list of the fishes of the mainland of North and Middle America re-
corded from both freshwater and sea water. Amer. Midl. Nat., 28(2):

Herald, Earl S., and Roy R. Strickland
1949. An annotated list of the fishes of Homosassa Springs, Florida. Quart.
Jour. Fla. Acad. Sci., 11(4): 99-109.

Hubbs, Carl S., and Mott D. Cannon
1935. The darters of the genera Holoepis and Villora. Misc. Publ. No. 30,
Univ. Mich., Mus. Zool. 93 pp., 3 pls., 1 fig.

Kilby, John D.
1949. A preliminary report on young striped mullet (Mugil cephalus Linnaeus)
in two Gulf coastal areas of Florida. Quart. Jour. Fla. Acad. Sci.,
11(1): 7-23, pl. 1.

McLane, William M.
1955. The fishes of the St. Johns River System. (Unpubl. Doctoral Disserta-
tion, Univ. of Fla. Library).

Mansueti, Romeo, and Ralph Pauly
1956. Age and growth of the northern hogehoker, Trinectes maculatus macu-
latus, in the Patuxent River, Maryland. Copeia, 1956(1): 60-62.

Marshall, Nelson
1947. Studies on the life history and ecology of Notropis chalybaeus (Cope).
Quart. Jour. Fla. Acad. Sci., 9(3/4): 163-188, pls. 2.

Vol. 11


Odum, Howard T.
1953. Factors controlling marine invasion into Florida waters. Bul. Mar.
Sci. Gulf & Carib., 3: 134-156.

Stubbs, Sidney A.
1941. Solution a dominant factor in the geomorphology of Peninsular Florida.
Fla. Acad. Sci., 5: 148-167.

West, Erdman, and Lillian E. Arnold
1956. The native trees of Florida. University of Florida Press, Gainesville,
Florida. 218 pp.


It q U
t-~ CD -i i

C~ b- -0 C 4 Ci

CCO C O C]] I C0
CDO C) CD it-t I-- Co Cq

C] 0 1-.IC V ClI]

CO to CO N 00 N CD I 1 0
V 10 if O SD c4 I-

I I I 1I I I I I I I I

-- CeCeN-
-N c]-
cqC 0-

c- 00 0C CD C
Ce -~ "~"

O CO 0 CO 1 I CO 0 I -
10 b co C 10i -

-3 C] -

V0) inm~ N IC

j~~in lI Ii -

-C] V~"


- Ce -

8 w8

-~ .B
-C c4.

Vol. 11

O0 in c- 0) in
Ce Co CO 1- CO ] CV

N 01

1n 10

C] -


Io I

S I oI mI I I c Vc 1 c

0 0 I I 0 f CO 00 C i I [1 |

l I c c C I | l 1 001 I 1

bOO CD CQ1 OI PCi II r- hO I I

I 1 C^ I I I I I I I

w0 II o1i I t I cO

S1 Il l I I I i I

I 1 l 1 i i | 1 1 l



o C



r mS 13 C CI I II I I I$


05a 'C

C 1-

se 0 ] I I I I I I1 5

1PC s0 5


- C E


0 10I 0

t- CS to v 0 C
wn ml

Cl CO 0m 10 I^ i0
i-i ^~ u

1 I I I I

ill-lcO C

f c m c ) 0-*

6 C61-H 0C 6CD qO V o ci iC oi Coi 1-4

N .
dM i- i- m

Nc N so co m 00o m

10 I m lI C
N 'tGo co of I v cq cc co 'IT O co N'

cdI o o mc co o

oo m CC9 Co

0- O10 10 q |O tI- CO I
C CO O 10010Co to

co H o m co N I
o 10

c c cq cq | cq co co I co (


0) 00

O I I] n 1

ml Cq I c C Ic

Io- I" I I


eq I I

II l I I

li ~I

10Il112 100j II-a II

I11 1S I 1 III 3
0.C CO

r. r

~ ~ZuCeO ecdO4.JOO

c 0 01 0 ,I! Do COd 0i

c CcOeq

qI c 1 -1c "

I-I It I I

CI jj

*^ a

B a
i -

$ uO

.4aJ -- a lilllli1l

0 co

I 1111111 1

o Il.


I 0 I xE

8 c4 oi do -

I I i 1 I I 1 1 I

O -

- 1-


I I q L I I c
I I0 K
0000 .coC


Sooo om oo

i 2888 8

I 0


II I 25

co 0 -

I1l Is 1l l l 1 I I

- C C 100110 eq
10 C t-

I oI o co

eq -


co0 0N0 C10 Iq
.-4 t-7q

Ic I l I 0

I l I I Il
- G0 00

1 'lI I [
^f '"' 10Ci

I 11

IE- oI
C] ~'


0] C


co eqoe
CD0 eCq-

00 I 10 10 Co
C0 10 10 01 C


'- &

I I I I I I I I I I 1 1 1 I I I 1 q

Sq q o oo q oq qq +o o
ioe m o o 16 co >i o6

Co o0 Ci 0 I I| ( 1 I06

q01 O 00

I l' c Ir 1 i l I : .4 I I
0 c lo m Ito
o l- l c6' l Ic. l I

p. e~


I t I I I I

0 o0 01 00t 00
2 BG 8082

CO [ C O] 00c

I- il0-(
cq oi q q 0o
<-< 1Q 0 C6'

It II 0 o

I 1
t- I

cv In CD m
,- -
5- mCoo

1 II 1 q
I 1 ai I I cs o


S00 0 c

I I "1 t 1
i- IT 1 co
in o i- C

I I 0 I0 1 0I q

- I C II I I I

CaO- 0



^ o o s e-

SI I q I c
0o in CO CI C

U) ~ U





v Co


-00 01


U) p

3 s N 1

sl g. .. i

c1C 00000 tOo 000
i- ---

I I I IO I C I 1 O


SCO I I I Im* I I I C'

Mt- 0 0 0
mom 01


I m
C-m C

I I 0 --I
i 7wm


SqO e oq o00 0 C
4co cc e goC 0i

I | o M c I I I
0l- C10-e eq-
1 1 1 1 1 1 1 1 1 1 1 1

I I I l I IQ R I
1~- CO

C | CO [ 00 q q q

10 il; I Ij %6C6o|CO0.-| fI |1|
com c v v 0140

I~ l I I-Z K liii

000 0 q'i0 In 1) 0 CO 0- 00C 0 C
kO -4eq co co c -i-l co o om o ,V
-, 1~- 'IV -

0 )

-e= 5 -^

8 M.2~
*C ft


z -z
pg |i
-4- 4-
Ot00 l
zzzj C




ot- I I I I I Iml

v, II

Contributions to the BULLETIN OF THE FLORIDA STATE MUSEUM may be in any
field of biology. Manuscripts dealing with natural history or systematic problems
involving the southeastern United States or the Caribbean area are solicited
Manuscripts should be of medium length-50 to 200 pages. Examination for
suitability is made by an Editorial Board.
The BULL.TIN is distributed worldwide through institutional subscriptions and
exchanges only. It is considered the responsibility of the author to distribute his
paper to all interested individuals. To aid in this, fifty copies are furnished the
author without cost.

Highly recommended as a guide is the volume:
Conference of Biological Editors, Committee on Form and Style.
1960. Style manual for biological journals.
Amer. Inst. Biol. Sci., Washington. 92 p.
Manuscripts should be typewritten with double spacing throughout, with ample
margins, and on only one side of the paper. The author should keep a copy; the
original and a carbon must be submitted. Tables and legends of figures should
be typed on sheets separate from the text. Several legends or tables may be
placed on a single sheet.
Illustrations, including maps and photographs, should be referred to as "figures."
All illustrations are reduced to a maximum of 4-1/4 by 7-1/8 inches. Size scales,
wherever they are necessary, should be incorporated into the figure.
References to literature should conform with the bibliographic style used in recent
numbers of the BULLETIN. Spell out in full the titles of non-English serials and
places of publication.
Footnote material should be kept to a minimum. However, provide copy for a
footnote detailing the title, affiliations, and address of the author (see recent
numbers of the BULLETIN).
Manuscripts must be accompanied by a synopsis-a brief and factual summary
(not a mere description) of the contents and conclusions, which points out the
presence of any new information and indicates its relevance. In it list all new
organisms described and give their ranges; indicate all taxonomic changes pro-
posed. The synopsis, written in full sentences, should be concise, but completely
intelligible in itself without references to the paper, thereby enabling the busy
reader to decide more surely than he can from the title alone whether the paper
merits his reading. The synopsis will be published with the paper. It does not
replace the usual conclusions or summary sections. It may also serve as copy
for the abstracting services.
Manuscripts and all editorial matters should be addressed to:
Managing Editor of the BULLETIN
Florida State Museum
Seagle Building
Gainesville, Florida