PROCEEDINGS OF
THE FLORIDA ACADEMY OF SCIENCES
(Issued Quarterly)
VoL. 6 Nos. 3-4
A PRELIMINARY STUDY ON THE DISTRIBU-
TION AND HABITS OF SOUTH
FLORIDA TERMITES
E. MORTON MILLER AND DOROTHY B. MILLER
University of Miami
This paper should be regarded as a progress report on a single
phase of studies being made by the writers on the ecology of termites
in Florida. Acknowledgement is given of the aid received from the
Council of the Florida Academy of Sciences through their annual Re-
search Grant for 1940; but this paper does not embody the final
results of the work made possible by that grant. Further progress
reports will be published from time to time, and, when some measure
of completeness is reached, such conclusions as may be warranted will
be presented in a final summarizing paper.
The only studies made on Florida termites prior to those here dis-
cussed were by Banks and Snyder, published in 1920. Collections
made in recent years by Dr. A. E. Emerson and ourselves have added
new information, noted herein, and have furnished the data necessary
for the construction of a practical key for the identification of the
Florida species, which follows this report.
All of the thirteen species of termites reported from Florida are
represented in the southern half of the state, below a line stretching
from Tampa to Melbourne. Two of these however-Kalotermes ap-
proximatus and Calcaritermes nearcticus-are not known to occur south
of Bartow, and seem to be more characteristic of the central and
northern parts of the state. The distribution of all of these species
is still quite imperfectly known, but examination of the records and
accompanying field notes leads us to formulate certain tentative gen-
eralizations concerning their occurrence.
Kalotermes bequaerti has been found only on the upper east coast
keys-Largo and Elliott-and is apparently rare. Little can be said
of its habits, except that its galleries closely resemble those of the dry
wood termite, Cryptotermes cavifrons, and it is evidently confined to
hardwood hammocks.
102 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
Kalotermes jouteli is largely limited to coastal zones. All of our
records fall within ten miles of the shore line. Jouteli is very abund-
ant within this area, particularly in dead wood in' mangrove swamps.
Galleries may be made in very hard wood and the occupied galleries
are nearly always in fairly substantial material. Well before any
portion decays to the point of fragility the termites of this species
:apparently abandon the zone and work into sounder wood. The spe-
cies is essentially a dry wood form, although not to the same extent as
K. snyderi. Soldiers appear to vary in size in correlation with the
age and population of the colony.
Kalotermes schwarzi is similar in habits and distribution to jouteli,
except that sckwarzi is encountered more frequently in coastal and
lower Everglade hammocks than is jouteli. The colony workings
are difficult to distinguish from those of jouteli in the field; the sol-
diers, however, lack the pigmented eye that is so characteristic of the
latter species.
Kalotermes snyderi has sometimes been recorded under the name
marginipennis. It is the most widely distributed Kalotermes in Flor-
ida, and seems to be the only one that occurs far inland. It is very
characteristic of cypress swamps; in fact, it seems to select cypress
wood in preference to some other kinds. In South Florida, at any rate,
untreated cypress telephone poles are frequently infested by snyderi,
while nearby wood of other types escapes attack. Man-built structures
are also occasionally infested. The moisture requirements appear to
be much less than for jouteli and schwarzi; and although snyderi is
habitually encountered in cypress swamps, it is not found in damp
wood. No explanation can yet be given for these differences in habit.
One hesitates to suppose that the distribution is related only to food
preferences, because in laboratory colonies all species of Kalotermes
are able to utilize cellulose from various sources.
The subgenus Cryptotermes includes dry wood and "furniture"
termites. Two species, cavifrons and brevis, are encountered in Flor-
ida. Cavifrons is apparently a native species, while brevis is very
likely introduced. Cavifrons has been found in natural habitats only,
and brevis exclusively in furniture and buildings. Brevis has been col-
lected only in Key West and the Miami area. Both species occur in
air-dry wood.
Cryptotermes cavifrons is the most abundant species found in South
Florida tropical hammocks. In some localities 75% of the dead,
standing limbs and stumps show signs of cavifrons attack. Dead
stumps or saplings that have begun to decay in the heartwood zones
usually have cavifrons in the outermost, well-cured and non-decayed
zones. Old galleries are frequently filled with excrement in the form
A PRELIMINARY STUDY ON SOUTH FLORIDA TERMITES 103
of minute, regularly-formed pellets or granules. Such droppings are
also accumulated in the galleries of brevis, and when shoved out
through small exit holes constitute the first clue to workings within
furniture or woodwork. Brevis is apparently being spread through-
out South Florida in infested second-hand furniture. Why cavifrons
has not yet become of economic importance is a puzzle, but it is our
opinion that as man encroaches more and more upon the natural habi-
tat of the species, his buildings may become the subjects of attack
by this form.
It is of interest here to note that soldiers in colonies of both brevis
and cavifrons are rare and difficult to locate. The soldiers of both
species have truncate heads without enlarged mandibles, and appear
to be able to function in colony defense chiefly by using the heads as
armored plugs to the invaded galleries.
One other representative of the family Kalotermitidae is found in
south Florida. This is Neotermes castaneus Burmeister. A second
species, angustoculus, has been listed by Snyder, but collections by
Dr. A. E. Emerson and the senior author at the type locality have pro-
vided a graded series of forms whose characters merge into both spe-
cies categories. Angustoculus is, therefore, now considered as synony-
mous with castaneus. Neotermes castaneus is damp-wood inhabiting,
and is frequently found in living trees, where it utilizes the sap as a
source of moisture. Castaneus is known from the east coast keys, the
Miami area, and the Pine Crest area in the lower Everglades. There
are no other records in Florida except a single, yet unverified, report
from Vero Beach region in 1900. In Dade County the species seems
to be confined to well-developed hammocks, although a few instances
of infestation of avocado trees are at hand. In these cases the original
colonizing appears to have occurred via dead, decayed wood at the
base of branches. It is doubtful whether entrance is ever made through
live, uninjured tree bark.
The genus Prorhinotermes is represented in our fauna by one spe-
cies, P. simplex. It is apparently restricted to Dade County, although
no ecological reasons for this have yet been deduced. The genus else-
where, however, is confined to tropical or near-tropical conditions and
has never been found more than a few miles inland from marine coasts.
Simplex, our species, is not known to occur more than five miles in-
land from the bay shore. The suggestion has been made that salt
water is somehow involved in the requirements of these forms, but
experiments by University of Miami students have shown no differ-
ential survival between experimental colonies supplied with fresh and
with sea water moisture. Temperature would seem to be an obvious
limiting factor in distribution northward, but simplex survives chilling
104 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
to immobility without observable injury. We are not yet satisfied that
the known limits of distribution will remain such with further study.
Prorkinotermes simplex is found in two major habitats. It occurs
in damp, but not immersed, wood in mangrove swamps. In such lo-
cations use is made of sapwood and, after partial decay, of the heart-
wood. A large, old colony will frequently have eaten out the inside
wood completely and replaced it with a hard, carton-like material com-
posed of excrement and chewed wood. This is the nearest approach in
this area to the so-called nest-building habits of West Indian and
other tropical species. Pine stumps are also occasionally inhabited by
P. simplex; the heartwood does not seem to be attacked, as a rule. In
either habitat considerable moisture is required. P. simplex is definitely
a damp-wood, not a subterranean type.
Three species of the subterranean genus Reticulitermes are repre-
sented in South Florida: flavipes, hageni, and virginicus. These spe-
cies, together with Kalotermes approximatus, form the temperate ele-
ment in our termite fauna; all of our other species appear to have
been derived from Antillean or Yucatan origins.
All species of Reticulitermes require ground contact and moisture
for the establishment of a thriving colony; but of the three, flavipes has
the highest moisture needs. Hageni and virginicus withstand consid-
erable drying, and, when sealed within galleries, can tolerate "dry
wood" conditions. Distribution in the field follows the moisture fac-
tor. Hageni and virginicus are more typical of the dry sand dunes of
the West Palm Beach area, the pine flatwoods and dry prairies; while
flavipes is encountered in moister situations such as hammocks or
locally moist spots in the foregoing areas. Species of Reticulitermes
are less common in the Miami oolitic pineland than in nearby sandy-
surfaced zones, and are rare on the upper and lower Keys of the east
coast. Flavipes is apparently absent from the Keys. This distribu-
tion, of course, is obviously related to the nature of the substratum.
These three species develop the most populous colonies of any
Florida termites. Furthermore, though classed as subterranean, a
strong colony may extend branches ten to twenty feet upward unto
wooden structures above ground.
Until 1940, Calcaritermes nearcticus, the only representative of
its genus in the United States, was known only from three isolated
winged specimens described by Snyder in 1933. In the fall of 1940
we were fortunate enough to discover colonies of this species near
Winter Park and Bartow. A description of the previously unknown
soldier and nymphal forms was published in this journal (Vol. 6, No. 1,
pp. 5-8, 1943), and their characters are given in the key which follows
this paper.
A PRELIMINARY STUDY ON SOUTH FLORIDA TERMITES 105
Although the genus is essentially tropical, this species has not been
found in the near-tropical tip of south Florida, and its presence there
now seems unlikely in view of the amount of searching that has been
done. This presents a zoogeographic problem to which we have no
answer as yet.
Judging from the colonies found in north and central Florida, this
species has a development somewhat similar to that of Cryptotermes
cavifrons. Calcaritermes, however, occurs in damper wood than
Cryptotermes, and lines its galleries with a brownish plastering sub-
stance.
SEASONS OF ALATE PRODUCTION BY FLORIDA TERMITES
(Eased on collections to 1940 by A. E. Emerson, T. E. Snyder, and E. M. Miller)
|. S S .
V" isti
January x x x x x*
February x* x x
March x* x* x x x* x* x*
April x* x x x x x*
May x* x* x* x* x
June x x x* x* x
July
August x x
September x x
October x x x x* x x x
November x x x x x* x
December x x x* x
x-Months in which maturely pigmented alates have been collected, either flying;
or in colonies.
*-Probable peak or center of swarming period.
One of the most important phases of termite biology concerns the
swarming activities of the winged caste, since flying alates are largely
responsible for establishing new colonies and dispersing the species.
Information about swarming thus far obtained has been summarized
in the table given above. It should be emphasized that this is a
tabulation of alates, not all of which were actually flying. However,
the major swarming period can be estimated and is indicated therein.
Generalizations which seem to be justified are these:
106 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
1) Calcaritermes has a well-defined period of alate production,
January through April.
2) Cryptotermes brevis swarms with regularity in the summer,
May through September. Flights have been observed in
evening, and in early morning up to 10 A. M.
3) Species of Kalotermes appear to be less closely restricted in
alate production to any single season than are the pre-
ceding. This is not unexpected in view of the primitive
nature of the genus and the equable climate of south
Florida.
These Kalotermes species, so far as known, are night or
early morning flyers.
4) Prorhinotermes simplex swarms regularly from October through
January; it habitually flies at dusk between 6 P. M. and
9 P. M.
5) Reticulotermes hageni and R. virginicus, which are very close
taxonomically, appear to be reproductively isolated, i. e.,
swarm at different periods. Virginicus swarms and its
alates have been taken only from March through May,
while hageni alates occur in the fall and winter months.
Virginicus swarms throughout the daytime until 5 P. M.;
hageni flies from noon to 5 P. M.
The nature of termite castes is still largely to be investigated. Are
the castes genetically or environmentally induced? Is caste determined
irrevocably early in the individual termite's life history, or is there plas-
ticity until a later instar? Some evidence showing that in P. simplex
nymphs are capable until late instar of becoming reproductive,
soldiers or workers has been obtained and will be published elsewhere.
Individuals with developing wings may, in the absence of soldiers, be-
come soldiers, and in the absence of reproductive become functional re-
productives within four weeks' time. Hence the superstition, once
common, that destruction of the queen in a colony would result in the
death of a colony must be abandoned for this species.
Evidence on the nature of castes may come also from study of
intercaste individuals. Several such individuals of P. simplex and two
of Calcaritermes nearcticus have been collected. In both instances
these have occurred in laboratory colonies under somewhat artificial
conditions.
The most extreme and consistent damage by termites to human
property in south Florida is done by Reticulitermes hageni and vir-
ginicus. These subterranean forms have practically always gained
access to the structure by faulty construction which permitted ground
A PRELIMINARY STUDY ON SOUTH FLORIDA TERMITES 107
contact with lumber. In Dade County, Prorhinotermes simplex does
considerable damage to the structures which have a good moisture
content. This species is not subterranean, but attacks moist wood,
in contact with the ground or receiving moisture from leaky walls
or roofs.
Kalotermes snyderi seems to be the major dry wood termite doing
damage to electric line poles in the southern half of the state. In
Dade County great numbers of untreated cypress poles have most of
the sapwood destroyed. The electric company periodically strips
off the damaged wood. This is no doubt useful in arresting rot which
follows dry wood termite galleries; but it does not always destroy
all the termites.
In Dade County the "furniture" termite, Cryptotermes brevis,
appears to be spreading-partly through the second-hand furniture
trade. Severe damage to valuable furniture may be done by this
species, although as a rule the colony grows slowly. Damage to door
frames, wood flooring, etc., has also been observed.
Recommendations as to prevention and control have been made
from the construction standpoint by E. S. Frash, of the University
of Florida Engineering School, and from the biological viewpoint by
Snyder (1935), and Kofoid and others (1934). It should be em-
phasized, however, that knowledge of termite species and habits is a
prerequisite for effective control measures, and that each case of in-
festation must be analyzed individually. Furthermore, the examina--
tion and licensing of control agencies by a properly qualified board
would be desirable in this state.
A KEY TO THE TERMITES OF FLORIDA
A. E. EMERSON AND E. M. MILLER
University of Chicago and University of Miami
IMAGOES
1. Three or more parallel chitinized veins near the costal border
of the w ing....................................................................... ........... 2
Two parallel chitinized veins near the costal border of the
w in g ....................................................................... ..................... .... 8
2. Median vein much more weakly colored than radius.................. 3
Median vein as strongly colored as the radius, or nearly so........ 6
3. Median vein ending near the tip of the wing................................ 4
Median vein joining the radius about two-thirds the length
of the wing from the suture..................... ........................... 7
4. Width of head including eyes 1.38-1.65 mm. Length of fore-
wing from costal end of suture 9.40-12.22 mm. Yellow-
brown.........................................................................K alotermes schwarzi Banks
Width of head including eyes 1.23-1.35 mm. Length of fore-
wing from costal end of suture 8.27-9.40 mm. Yellowish
Kalotermes snyderi Light
Width of head including eyes 1.23-1.27 mm. Length of fore-
wing from costal end of suture 6.75-6.96 mm. Dark-brown.
Kalotermes approximatus Snyder
Width of head including eyes .97-1.00 mm. Length of fore-
wing from costal end of suture 7.52-7.71 mm. Yellowish.
Kalotermes bequaerti Snyder
5. Radius with numerous branches and cross veins........................ 6
Radius with few or no branches and cross veins. Wing mem-
brane coarsely punctate........................................Ccaritermes nearcticus Snyder
6. Hairs on pronotum short about .06 mm. or less in length.
Area between ocellus and eye same color as rest of head
Kalotermes jouteli Banks
7. Width of head with eyes .85 -.97 mm.............Cryptotermes cavifrons Banks
Width of head with eyes 1.05-1.15 mm.............Cryptotermes brevis (Walker)
8. Forewing scale barely overlapping the base of the hind wing
scale. Profile of top of head convex.................................... .......... 9
Forewing scale overlapping at least about half the length of the
hindwing scale. Profile of head straight.....Prorhinotermes simplex (Hagen)
9 Color of head and body light brown to yellowish brown
Reticlditermes hageni Banks
Color of head and body dark brown to black........................... 10
10. Ocelli plainly less than their diameter from the eye
Reticdlitermes virginicus Banks
Ocelli either their diameter or more than their diameter from
the eye..................................... .............................R eticditermes flavipes (Kollar)
2(Neotermes angustoculus Snyder is now believed to be synonymous with N. castanets,
since characters formerly used for distinguishing these forms overlap in a graded series of
specimens from Paradise Key, the type locality for angustoculus.)
108
A KEY TO THE TERMITES OF FLORIDA
SOLDIERS
1. Pronotum approximately as wide or wider than the head........... 2
Pronotum distinctly narrower than the head.............................. 9
2. Junction of front and vertex fairly flat; head not sharply
tru n cate ............................................................... ..... ........................... 3
Junction of front and vertex nearly a right angle; head sharply
truncate ........................................................................ ..... .......... 7
3. Third antennal joint distinctly larger and darker than the second
or fourth joint................................... ................ ......... ............... 4
Third antennal joint not conspicuously different from neighbor-
ing joints...................... .............................Neotermes castaneus (Burmeister)
4. Eye pigmented........................... ..............................Kalotermes jouteli Banks
E ye unpigm ented ........................................................ .......... ... .. 5
5. Third joint of antennae usually longer than fourth and fifth
together. Gula proportionately wide in small forms. Labrum
truncate at tip..... ..............................................Kalotermes schwarzi Banks
Third joint of antennae about as long as fourth and fifth
together. Gula proportionately narrow. Labrum bluntly
pointed at the tip................................................... ................... 6
6. Anterior margin of pronotum deeply to mediumly emarginate.
Profile of head with a nearly smooth curve between the front
and vertex.................................. ........ ..........................K aloterm es snyderi Light
Anterior margin of pronotum mediumly emarginate. Profile
of head with obtuse angle between front and vertex. Small
species. Head width about .97-1.06......................Kalotermes bequaerti Snyder
Anterior margin of pronotum slightly emarginate. Profile
of head with obtuse angle between the front and vertex
Kalotermes approximatus Snyder
7. Front slightly oblique, relatively smooth, a deep furrow at
the vertex..............................................................Calcaritermes nearcticus Snyder
Front conspicuously concave, more or less roughened.................. 8
8. Top of head somewhat concave and smooth....Cryptotermes cavifrons Banks
Top of head with distinct depression, rough............Cryptotermes brevis Walker
9. Sides of head fairly parallel.................................... ................... 10
Sides of head distinctly converging toward the front
Prorhinotermes simplex (Hagen)
10. Width of pronotum .81-1.10 mm..................... Reticulitermes flavipes (Kollar)
Width of pronotum .67-.81 mm....................................................... 11
11. Gula narrower with more abruptly demarcated anterior portion;
usually larger and often with more hooked tips on mandibles.
Reticulitermes virginicus Banks
Gula wider with more gradually demarcated anterior portion;
usually smaller and with less hooked tips on mandibles
Reticulitermes hageni Banks
FISHES OF SILVER SPRINGS, FLORIDA
CARL L. HUBBS AND E. Ross ALLEN
Museum of Zoology, University of Michigan
and
Florida Reptile Institute, Ocala, Florida
Silver Springs, located five miles east-northeast of Ocala, Marion
County, comprise one of the largest and most widely known of the
crystal-clear outflows from the limestone cap of northern Florida. The
chief flow emerges at a constant temperature of about 720 F. from a
funnel-shaped depression more than 35 feet deep. The combined dis-
charge is estimated at 22 million gallons per hour. The overflow, Silver
Springs Run, meanders 7 miles before entering Oklawaha River,
which is a tributary of St. Johns River.
The chemical analysis of the water in parts per million is: silica 32,
iron .06, calcium 70, magnesium 11, sodium and potassium 4.4;
radicals-bicarbonate 205, sulphate 43, chloride 7.6, nitrate .86; total
hardness as calcium carbonate 220.
The springs form a pond with a surface of 2 acres. The current
in the pond varies from none to moderate; it is about 4 miles per hour
at the spring source. On account of the extreme clarity of the water,
rank growths of aquatic vegetation (see figures), with a coarse-leaved
Sagittaria predominating, beautify not only the shallows but also
the sides of the deep spring pits. The bottom is of marl and some-
what cavernous limestone. The shores are low and in part marshy.
In 1929, when in a more or less natural state, the Silver Springs
pond was almost wholly surrounded by a marsh, containing much
shallow water rich in plant growths. This marsh has since been filled
in to a large extent, thus eliminating much of the spawning area for the
fish of the spring pond. The edges of the springs have been cleared
out and filled in, and a high stone wall has been built around the north-
east side of the lake. These changes are probably to a large degree
responsible for noticeable decreases in the fish population. Waves
from the speed boats continually passing back and forth on the river
have probably been a contributing factor, by destroying nests and
frightening off the guarding males. Overfishing is not the factor,
for no fishing is allowed in the spring or in the outlet. The fish seem
to be in good condition, and comparatively free of parasites. Silver
Springs fish seldom rub against submerged logs and sticks, as do fish
infested with external parasites. Excepting Signalosa no species
has been observed to die in large numbers. Predators are also few.
Fish-eating birds are not very common and only occasionally feed
FISHES OF SILVER SPRINGS, FLORIDA
around the springs proper. Racoons though plentiful downstream do
not feed in the springs. Softshell turtles occur but are not very
common, and they have been seen to feed very little upon fish (in
fact these turtles feed every day on the bread that is used to attract
the sunfish). Up to seven years ago giant fresh-water shrimp, Macro-
brachium jamaicense (Herbst), ranging in over-all length to 22 inches,
were rather common in Silver Springs, living in the deepest holes and
coming out at night to feed on fish, but since then none has been seen.
For these reasons we assume that neither predators nor parasites nor
overfishing is responsible for the decimation of fish in Silver Springs.
The altered environment seems to be the controlling factor.
Species which have become particularly scarce of late are Mollien-
isia latipinna, Huro salmoides, Esox niger, Amia calva and Elassoma
evergladei. The sunfishes, which are fed daily as a tourist attraction,
are thriving.
Through the windows of glass-bottom boats, a diving bell and the
"Photo-Sub," many thousands of visitors have enjoyed the entrancing
under-water vistas and have observed the aquatic life in a fish's-eye
view. The transparency of the water makes it possible to identify
the fishes at almost unbelievable distance and to observe their intimate
ways of life (see figures). Despite these facts no authoritative account
of the fishes of Silver Springs has been published.
One of the authors (E. Ross Allen), as proprietor of the Florida
Reptile Institute, has had the opportunity of observing the aquatic
life of Silver Springs since 1929. From August 28 to 31, 1935, he
assisted Dr. Samuel F. Hildebrand in making an initial inventory of
the fishes in these waters. The collection preserved on that occasion
has been supplemented from time to time. During the Florida meet-
ing of the American Society of Ichthyologists and Herpetologists, on
April 5, 1941, Carl L. Iubbs, with several colleagues and students,
observed the fishes in the clear waters and examined the preserved
collections in the Florida Reptile Institute. Except for the observa-
tions of this date the natural history notes were made by Allen. In
1942 a series of the specimens which had been collected locally was
sent for further study to the University of Michigan Museum of
Zoology. The latest collection was made on August 4, 1943.
The equipment used in collecting fish in the deep, clear water of
Silver Springs was a face mask (goggles) and a spear. The spear or
dart was shot from a rubber slingshot. This weapon kills fish at a
distance of twenty feet. They can be shot while they are swimming
by leading them a few inches. The face mask enables one to see
clearly under water.
112 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
The systematic remarks are by Hubbs. The paper contributes
to our knowledge of (1) the systematics of southeastern fresh-water
fishes, (2) the local fish fauna, and (3) the life ways of the fishes of
Silver Springs.
Dr. Samuel F. Hildebrand of the Fish and Wildlife Service has
generously placed at our disposal his notes on fishes observed in Silver
Springs in August, 1935, with remarks on specimens sent to him by
Allen in 1936. Dr. Archie F. Carr, Jr., of the University of Florida
has also supplied information on the fishes of the Springs.
Collections of Silver Springs fishes are preserved in the United
States National Museum, the University of Michigan Museum of
Zoology, the University of Florida Department of Biology, and the
Florida Reptile Institute.
LEPISOSTEIDAE
I4episosteus platyrkincus De Kay
FLORIDA GAR
The common gar of Florida has been confused by most authors, for ex-
ample by Jordan and Evermann (1896: 110-11) with the shortnose gar, L.
platostomus Rafinesque. Recent studies, however, show that platostomus is re-
stricted largely to the silty rivers of the Mississippi Valley. The Florida species
are well described and figured as Lepisosteus platyrhincus by De Kay (1842:
273, pl. 43, fig. 137), and as Cylindrosteus castelnaudii by Dumeril (1870: 347,
355-56, pl. 21, figs. 2-2b) and by Fowler (1911: 605, 609, pl. 38. figs. 13-14).
C. megalops Fowler (1911: 605, 609-11, pl. 38, figs. 15-16) was apparently based
on a large-eyed example of the same species. Data on proportions in L.
platyrhincus were given by Hammett and Hammett (1939: 197-209, charts 1-5).
These authors suggested that a striped and spotted variety may occur near Engle-
wood, Florida, but large series from the same region, made by the same col-
lector (Stewart Springer), indicate that the differences in coloration are due to
age and individual variation.
To judge from the collections examined and from local testimony, this
species seems to be the only gar, other than L. osseus, in at least the major part of
peninsular Florida. On the Atlantic slope it ranges north at least to the
Okefenokee Swamp in southeastern Georgia. It is the Florida representative of
the spotted gar of the Mississippi Valley, Lepisostemu products Cope, which
ranges into western Florida. A specimen from Thomas's Mill Pond, 8 miles south
of Marianna, Jackson County, is in the University of Michigan Museum of
Zoology. On the basis of this specimen L. oculatus was listed for Florida by
Carr (1936: 78), before it'was learned that oculatus is a synonym of pfbductus.
That species was described as Cylindrosteus products by Cope (1865: 86-87),
Dumeril (1870: 347, 357), and Fowler (1910: 605, 609, pl. 38, figs. 10-11).
Hubbs has recently contrasted L. products with L. platostomus, with which it
has long been confounded. The full data have not been published, but have
been given in summary in papers by Kuhne (1939: 19-21) and by Hubbs and
Lagler (1943: 76). In the paper last cited the ratios for the proportionate length
of the snout were inadvertently transposed.
L. platyrhincus agrees much better with L. products than with L. platostomus
in all the characters in which those species disagree, with one exception: the
Fig. 1. Main spring at head of Silver Springs, showing a school of stripedmullets (Mugil cephalus) passing over a bed of Sagittaria.
PhotoeraDh by E. Ross Allen.
0
,' ,j I
^^ ^^^^^^ ^- cn*
Fig. 2. Largemouth bass (Huro salmoides), striped mullets (Mugil cephalus) and an eastern bluegill (Lepontis macrochirus purpurescens)
in ''Silvpr Rivpr"A hptd nf ttnria -- r, -f rf +, Tb i.1 kI 4-, DL *. 1. r -- ..- .. .
FISHES OF SILVER SPRINGS, FLORIDA
snout is quite as short and broad as in platostomus. The ecological conditions gen-
erally prevailing through the range of platyrhincus are like those selected by
products.
This species is fairly common in Silver Springs, and specimens of several
sizes are in the collections. On April 5, 1941, a number were seen, some sol-
itary, others in a group at the shallow edge of the deep spring. Here they
lined up to await the schools of Signalosa, on which they were feeding. Some
broke the surface, obviously to breathe. The largest were about two feet long.
Lepisosteus osseus osseus (Linnaeus)
EASTERN LONGNOSE GAR
The single Silver Springs specimen at hand, although only 195 mm. long to the
end of the lateral line scales, already shows the roughening of the anterior trunk
scales (apart from the denticles) that is characteristic of the Atlantic Coastal
form of L. osseus. This character was pointed out by Cope (1865: 86-87),
Dumbril (1870: 326) and Fowler (1911: 606, pl. 38, figs. 3-4), but otherwise
has generally been overlooked. Whether it distinguishes a full species, as
thought by the authors quoted, or only a geographical subspecies, remains to be
determined. An adult was collected in Silver Springs on August 31, 1935, but it
has not been studied by us.
The local form of the longnose gar attains a larger size than does the Florida
gar: the largest ones are about four feet long and robust in build.
Longnose gars are much less plentiful in the Silver Springs region than they
are in many streams of central Florida. Some, however, may be seen at any time
of the year except on cold days, when they probably go into hiding. They seem
to become more abundant in warm weather and probably migrate upstream in
the spring. They seldom frequent the main spring "boil" at the stream head
(none were seen there on April 5, 1941): they usually stay downstream in deep
holes and eddies. In the deep water they may be seen during the day swim-
ming slowly back and forth, occasionally rising to break the surface in a smooth
roll. Others may be observed in shallow water in dark, shady places, under
bushes and logs, resting on the bottom as though asleep. Once in a while one
may be seen with a four or five inch "brim" (Lepomis, sp.) in its mouth. They
seem to do much of their feeding nocturnally. They have a habit of lying near
the surface at night, and persons paddling along in a canoe in the dark are
often startled by a sudden splash as the gars ruh off. On such occasions small
gars sometimes jump into the boat.
The courting habits of the longnose gar have been observed locally. During
the day two males have often been seen following one female. The fish then swim
very close together, with the males just above or just below the female.
AMIIDAE
Amia calva Linnaeus
BowrIN
Locally known as mudfishh," this species is not very common in the springs.
One specimen was collected on August 28, 1935, and two others on August 4, 1943.
A few were seen in the head pond, in the daylight on April 5, 1941. They show
no marked seasonal changes in abundance. Usually about a dozen can be seen by
looking for them at night with a light. Shallow pools just off the main stream
seem to be a preferred habitat. While swimming with a face mask in the spring
holes I (Allen) have seen them hiding on the bottom in the deep-water "grasses."
116 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
Occasionally they swim in a vertical position to the surface, tadpole-fashion, to
gulp air and return to the bottom.
At times the bowfin are seen nosing along the bottom, presumably in search
of crayfish. It is commonly thought that this fish is a destructive predator.
The Negroes consider the flesh good eating, but the meat is unattractive to the
white people because it is so soft. The largest local specimen on which there is a
definite record weighed 10.5 pounds.
A compact swarm of young mudfishh," resembling a school of young catfish,
was once found next to the bank. The vividly-marked fish were about two inches
long. They were packed so closely that they could be scooped up by the handful.
Obviously the bowfin in Florida has habits similar to those observed in the
North (Reighard, 1903).
CLUPEIDAE
Signalosa petenensis vanhyningi Weed
FLORIDA THREADFIN SHAD
The genus Signalosa was unknown as far east as Florida until 1925, when
Weed (pp. 142, 145-146) described the Florida form as S. atchafalayae varnhyn-
ingi. A study of an abundance of material, coming from the coastal low-
lands from Florida to El Peten, Guatemala, seems to indicate that neither S.
mexicana nor S. atchafalayae should be specifically separated from S. petenensis
(Giinther).
This small relative of the gizzard shad is not regularly present in the pool
of Silver Springs but occasionally invades these waters in immense droves. The
invaders probably come from St. Johns River, whence Fowler (1941: 229, figs.
1-2) reported the species (as S. vanhyningi) to be common. So far as known,
schools first arrived in 1933 but these soon disappeared. The next occurrence
was in 1941, when countless numbers suddenly appeared, and many died, to pro-
duce a smelly nuisance. They deserted the springs on March 15 but returned about
a week later. During the latter half of March the schools were too dense to be
seen through. The fish even got into one's bathing suit. On April 5, these shad
were somewhat scarcer but still excessively abundant. Later they disappeared.
Their return will be eagerly watched for. The possibility of cyclic behavior will
be kept in mind.
These fish swam in dense, compact schools, acting in surprising unison (as
indicated by widely shown motion pictures as well as direct observation). Whole
areas flashed aside on attack by a predator, but the gap thus made was soon
filled in. Others jumped to escape the enemy.
The individuals of a school maintained remarkably parallel courses, as they
rushed about ceaselessly, day and night. Most of the schooling was near the
surface, or in large sheets extending from near the surface to the mid-depths of
the deepest spring hole. Some schools penetrated into the deeper waters. Much
of this behavior agreed with the descriptions and theories of schooling published
by Parr (1927).
It is something of a mystery how so many thousands of plankton-feeding
fish found subsistence in the crystal water of Silver Springs. They did not appear
to be feeding.
The threadfin shad observed on April 5 varied much in size but all were
small fish. They were very silvery, but one could see the dark streaks of which
two stood out sharply in lateral view. The shoulder spot was not very con-
spicuous.
FISHES OF SILVER SPRINGS, FLORIDA
Locally, these fish were identified as "menhaden," and were fancifully supposed
by some to have entered the springs from the sea by the way of underground
waters!
Dorosoma cepedianum (LeSueur)
GIZZARD SHAD
Some specimens of gizzard shad from Silver Springs are in the preserved col-
lection. A few were seen on April 5, 1941, schooling with Signalosa. They were
recognizable in the water by reason of their much deeper body, larger size, and
more prominent shoulder spot. The gizzard shad travel in small schools, usually
in the deeper spring holes, and have been seen feeding like mullets off the vegetation.
CATOSTOMIDAE
The paucity of suckers in peninsular Florida is exemplified by the finding
of only one species in Silver Springs.
Erimyzon sucetta sucetta (Lac6epde)
EASTERN LAKE CHUBSUCKER
On April 5, 1941, a few large adults of this sucker, carp-like in size and build,
were seen in the springs, at depths of 10 to 25 feet. They were cruising over the
Sagittaria beds. The species is very common locally and is frequently seen
travelling during the day in small schools or in pairs. A specimen 274 mm. in
total length was collected on August 28, 1935.
CYPRINIDAE
Only two species of the minnow family were observed in Silver Springs, and
neither of these was abundant. In few parts of the Holarctic Realm, apart from
the peninsula of Florida, could one find such a rich fresh-water fish fauna with
so poor a contingent of Cyprinidae.
Notemigonus crysoleucas boscii (Valenciennes)
SOUTHEASTERN GOLDEN SHINER
Silver Springs is inhabited by a type of golden shiner so unlike the ordinary
representatives of the species as to be scarcely recognizable on first sight. After
making allowances for magnification in the clear water it seems that the golden
shiners here grow to a length of almost or quite one foot. Even more surprising
is the robustness of the body. A specimen 222 mm. in standard length is 85 mm.
in greatest body depth, and is therefore nearly two-fifths as deep as long; the
greatest width is 40 mm. The very deep scales are set off with unexpected con-
spicuousness by dark basal bars. In this example there are 16 principal anal rays:
a typical number for subspecies boscii. The caudal fin was seen to be red in life.
Most of the golden shiners seen in Silver Springs are large adults. Smaller,
less bulky individuals are probably quickly consumed by the hordes of predacious
fishes. Small ones were collected in 1935 and 1936.
Notropis, species
FLORIDA SHINER
A small, dark-striped minnow was observed on April 5, 1941, and on other
occasions, near the weeds in the shallow waters of the springs. It is probably
the Florida shiner, a species which has been confounded with N. xaenocephalus
and N. roses.
118 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
AMEIURIDAE
Ictalurus lacustris punctatus (Rafinesque)
SOUTHERN CHANNEL CATFISH
Among the myriads of catfish in Silver Springs one observes (as we did on
April 5, 1941) a number of very large fish with caudal fins more deeply emargi-
nate than in the common I. catus. Since in ameiurids this fin becomes more
rounded and less forked with age, these large fish surely represent a second spe-
ces. They are unlike the usual run of channel catfish in being blackish, without
evident dark spots, but mature adults of I. lacustris exhibit this color phase, and
all fish in Silver Springs tend to be dark. A few moderately light-colored half-
grown individuals were seen.
In the depths of the spring holes, in limestone recesses from which the water
boiled out, the largest catfish were seen on April 5 to be in territory-holding
pairs. One fish in each couple, certainly the male, was broad-headed and thick-
cheeked, whereas the mate was narrow-headed. Similar observations have been
made in the Great Lakes region and in the Mississippi Valley. In the North
such adult males correspond with the descriptions of Ameiurus lacustris (or
nigricans), a supposed species often referred to Haustor and to Villarius. The
females and all young agree with Ictalurus punctatus. In the Mississippi
Valley similar breeding males (not always large) have passed as Ictalurus
anguilla. This sexual dimorphism is general throughout the family (Hubbs,
1940: 209-10). For these reasons we have been treating Ictalurus punctatus
as specifically identical with "Ameiurus" lacustris (and with I. anguilla). The
problem, however, is complicated, because age, individual and geographical varia-
tions involve the very characters which distinguish the sexes in mature adults,
and many fishermen insist that there are two species of this type.
The largest catfish in Silver Springs when seen at close range on April 5
were judged to weigh 40 to 50 pounds, but the property of water to magnify
objects may have deceived the observer. The record weight of locally caught
fish is 42 pounds.
Observations made over several years show that these catfish hide during the
day in spring caves, heading against the current, and under rocks and logs, and that
they come out at night, apparently to feed, and can then be seen with a light
in all parts of the spring pond. They are very wary and fast.
In late spring many of these fish migrate into the stream, when they can be
seen during the day hiding in groups under low-hanging trees, along with other
fish near the bottom.
Some of the larger catfish have odd white spots, like gray hair, on the head,
and other marks by which the boat operators can recognize the individuals. The
same fish may be seen every day in the same retreat.
Most of these large catfish occur in spring holes downstream from the head
spring. A place of particular abundance is called "Catfish Hotel, with running
water in every room."
Ictalurus cats (Linnaeus)
WHITE CATFISH
The elimination of Villarius (Haustor) as a distinct genus, or subgenus of
Ameiurus, and the synonymizing of this group with Ictalurus, throws open the
problem of generic separations in the Ameiuridae. The species catus (obviously
distinct) has been associated systematically with the nominal species based on
males of Ictalurus punctatus. In all known characters it is obviously intermedi-
C
S *-
-
4'
Fig. 3. Largemouth bass (Huro salmoides) and sunfish (Lepomis macro chirus purpurescens and-one fish to right-Lepomis
microlophus microlophus) feeding on shrimp freed from floating vegetation in Silver Springs.
Photographby W. Nase, reproduced by permission.
a)~ra~
Photographlby W. Nase, reproduced by permission.
Fig. 4. "Christmas Tree Spring," with striped mullets (Mugil cephalus) and eastern bluegills (Lepomis macrochirus purpurescens)
in plain view. Photograph by E. Ross Allen.
FISHES OF SILVER SPRINGS, FLORIDA
ate between Ictalurus and Ameiurus. We may either counter convention and con-
venience by uniting the genera, or we may, rather arbitrarily, put catus (and some
similar Mexican forms) either in Ictalurus or in Almeiurus. To preserve a group
distinction, "caudal fin more or less deeply forked" as opposed to "caudal fin
weakly emarginate, truncate or rounded," it seems best to refer catus to Ictalurus.
The taxonomic and nomenclatorial conclusions regarding the larger catfishes were
first indicated by Carr (1936: 80), on our advice. Preserved specimens, collected
in 1935 and 1943, seem typical of catus.
The white catfish rather belies its name in Silver Springs, where it, along
with the other fishes, takes on a dark hue. The larger ones are blackish, like
the local adults of I. punctatus. Some pairs of catfish, definitely identifiable as
I. catus, were seen on April 5, 1941, in rock recesses. They were smaller than
the breeding adults of I. punctatus, and had less deeply emarginate caudal fins.
There was agreement, however, in that each pair consisted of a broad-headed
male ("Ameiurus lophius") and a narrow-headed female (ordinary catus).
The catfish which are chummed in swarms into the view of visitors are almost
entirely of this species. Along with the "bream" (Lepomis, spp.) they feed on
bread which is regularly thrown out from the glass-bottom boats. The point of
greatest abundance is the already-mentioned Catfish Hotel.
Although they are often quite active by day, the white catfish are also seen
at night in shallow water. They have the habit of lying very quietly on the
bottom in the mud, or just under a log. In the daytime I (Allen) can swim
down and touch them before they seem to be aware of my presence.
Ameiurus natalis erebennus Jordan
SOUTHEASTERN YELLOW BULLHEAD
The bullhead erebennus is obviously the southern form of Ameiurus natalis,
for its differentiae parallel the geographical trends exhibited throughout the
family (Hubbs, 1940: 209-10). It remains to be determined whether this varia-
tional tendency is sufficiently sharp to warrant retaining erebennus as a subspecies.
No bullheads were seen in Silver Springs on April 5, 1941 (they are secretive
by day), but one yearling of A. n. erebennus is included in the locally obtained
collection and 3 adults were collected on August 28, 1935.
This very inactive fish lies on the bottom and therefore little of interest has
been noted concerning its habits. Masses of algae have been found in the stom-
ach of this bullhead. At times it can be heard very distinctly grunting under
water, for instance when speared.
Mottled catfish are rarely seen in Silver Springs. They may be Ameiurus
nebulosus marmoratus (Holbrook), but we prefer not to list the species from this
locality until specimens have been collected.
ESOCIDAE
Esox americanus Gmelin
BULLDOG PICKEREL
Adults of this species are in the local collection. A small, red-finned Esox,
probably americanus, ranges south to the east coast of Florida. In the western
part of the state the little pickerel (referred, unsatisfactorily, to Esox vermiculatus)
do not have red fins, and other differences exist.
No bulldog pickerel were seen on April 5, 1941, or on any other date, in the
spring pond. Nor have they been seen in the main outlet stream. They occur
in shallow waters leading off from the outlet, and in adjacent marshes. Those
122 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES vol. 6,
Nos. 3-4, 1943
that were collected were picked or scooped up by hand at night, in water less
than six inches deep.
Esox niger LeSueur
CHAIN PICKEREL
Larger pickerel, up to 2 feet in length, occur solitarily and rather rarely in
Silver Springs, chiefly in the spring of the year. On April 5, 1941, one was
seen lying stiff and motionless near the surface in shallow weedy water, close
to shore, following the local custom of the species. Its yellowish mid-dorsal stripe
was vividly distinct. The color is always bright in Silver Springs. A specimen
was collected on August 28, 1935; another on August 4, 1943.
In Silver Springs, as elsewhere, the chain pickerel is ordinarily quiescent, but
swims very fast, in short darts, when it does move. One is occasionally seen
striking at a smaller fish. When a chain pickerel is disturbed it dives down in
the weeds to hide. The flesh is very good eating.
BELONIDAE
Strongylura marina (Walbaum)
BILLnFsH
Fish known as "needlefish" or "needle gars" often occur in the spring of
the year. None were observed on April 5, 1941, and they usually arrive some-
what later. Obviously this fish is a Strongylura, and presumably is S. marina,
which is the only species along the United States coast that is known to penetrate
commonly into fresh waters.
Day and night the billfish swim about on the surface in schools of 2 to 8
fish, often breaking water. They are seen more often downstream than in the
head pond. The only specimen that was collected was shot.
ANGUILLIDAE
Anguilla bostoniensis (LeSueur)
AMERICAN EEL
Eels occur in side streams and pools as well as in the head pond and outlet
stream.
On account of their secretive, nocturnal habits, the eels are not often seen
by day. They occur then in deep water. I (Allen) have scared them out of mud
as I walked on the bottom in 30 feet of water. At night they are seen feeding
in shallow water on a mud bottom, in which they often disappear when they are
disturbed. They are easily caught with a dipnet at night. Some of the speci-
mens have been preserved. The eels have a good flavor, like catfish.
A number of eels have been taken from rainbow snakes (Abastor erythrogram-
mus Latreille) which had caught these fish and taken them onto the bank to
swallow.
CYPRINODONTIDAE
Species of this family, known as killifishes, top-minnows, etc., are fairly com-
mon along the weedy edges of Silver Springs. Few specimens have been collected,
however, and the list given is probably incomplete.
Bangham (1942: 297) listed Fundulus similis (Baird and Girard) as from
Silver Springs but it is virtually certain that this essentially marine species does
not occur here. Dr. Bangham indicates by letter that he was confused in the iden-
FISHES OF SILVER SPRINGS, FLORIDA
tification of species of Fundulus, that his identifications were not checked and
that he preserved no specimens.
Fundulus chrysotus Giinther
GOLDEN TOPMINNOW
This is probably the commonest topminnow in Florida. Specimens were
collected in August, 1935, in Silver Springs and in nearby lakes.
Funduls dispar lineolatus (Agassiz)
SOUTHEASTERN STARHEAD
This is the topminnow generally known as Fundidus nottii. That name, how-
ever, along with Zygonectes guttatus and Z. hieroglyphics, was originally based
on material from near Mobile, Alabama. A study of fresh and preserved material
from various points along the Gulf coast indicates that the form ("nottii") of the
southeastern states and the peninsula of Florida does not occur about Mobile or
elsewhere to the westward, and that only one form, exhibiting striking differences
with age and sex, occurs about Mobile. This is the type commonly called
F. guttatus, but in view of the synonymy as drawn up by Garman (1895: 120-21),
under Zygonectes nottil, we must treat Z. guttatus (with Z. hieroglyphics) as a
synonym of Fundulus dispar nottii. The form commonly called nottii will there-
fore need another name, and may be called Fundulus dispar lineolatus (Agassiz).
This application of the names nottii and lineolatus, on our advice, was given first
notice in the key to Florida fishes by Carr (1936: 82). Both nottii and lineo-
latus are now provisionally referred to Fundulus dispar, because it appears that
they are the terminal elements in a graded chain of forms ending with F. dispar
dispar in the North.
Specimens of F. d. lineolatus were obtained in Silver Springs on August
23, 1935.
Chriopeops goodei (Jordan)
REDTAIL KILLIFISII
Specimens of this attractive fish were collected in Silver Springs and in a
nearby lake on August 28, 1935.
Leptolucania ommata (Jordan)
OCELLATED KILLIFISH
This minute species, with eye-like spots, is not yet reported from Silver
Springs proper, but almost surely occurs in marginal vegetation. Seven specimens
were seined in a nearby lake on August 28, 1935.
Jordanella floridae Goode and Bean
FLAGFISH
This beautiful cyprinodont occurs in the shallow water of Silver Springs,
among "grasses." One preserved specimen is at hand.
Lucania parva (Baird and Girard)
RAINWATERFISH
Several examples of this plain-colored killifish were collected at Silver Springs
on August 28, 1935. Another was obtained on August 4, 1943.
124 PROCtEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6
Nos. 3-4, 1943
POECILIIDAE
Gambusia affinis holbrookii (Girard)
EASTERN MOSQUITOFISH
This little fish, effective in mosquito control, is rather common along the
weedy shores of Silver Springs, but it is less abundant than it was formerly. Black-
spotted individuals occasionally occur. Series were collected on August 28, 1935,
and August 4, 1943.
Heterandria formosa Agassiz
LEAST KILImsH
This tiny live-bearer, once regarded as the smallest of all vertebrates, is well
represented in the local collection of August 30, 1935.
Mollienisia latipinna LeSueur
SAILFIN MOLLY
The local form of Mollienisia is typical of M. latipinna. The number of dorsal
rays (Table I) is average for the species, partaking neither of the increase char-
acteristic of the salt-water types of Pensacola Bay and Key West nor of the de-
crease that marks the races of Georgia and the Carolinas.
TABLE I
DORSAL RAYS IN MOLLIENISIA LATIPINNA FROM SILVER SPRINGS
Number of rays
13 14 15 Average
Number of Specimens......................... ............... 5 19 1 13.84
This popular aquarium fish is still rather common about the remaining marshy
edges of Silver Springs, but it is not as abundant as it was formerly. The active,
sail-finned males, with the blue reflections resplendent in the sunlight, always attract
attention as they strut about and circle the females in courtship. They keep up
this activity all day, but become quiet at night, when they stay in very shallow
water. One male in the preserved series (of aquarium stock?) is almost solid
black, and spotted ones are seen.
APHREDODERIDAE
Aphredoderus sayanus (Gilliams)
PIRATEPERCH
The occurrence of this weird, pugnacious little fish at Silver Springs is
attested by the collections. It is found only in side streams and pools, under
water lettuce.
FISHES OF SILVER SPRINGS, FLORIDA
PERCIDAE
Hadropterus nigrofasciatus Agassiz
CRAWLABOTTOM
The confused nomenclature'of the etheostomatines of the Hadropterus nigro-
fasciatus-scierus group is under much needed review. This is one of the few
types of darter that penetrates the peninsula part of Florida.
Archie F. Carr, Jr., informed us that he knew from observation that Silver
Springs harbored a species of darter, though no specimens had been obtained. Con-
sequently we watched particularly for darters during the trip of April 5, 1941,
and on shell-littered shoal bottom saw a few which were almost surely referable
to Hadropterus nigrofasciatus.
What is probably the same darter is very common on rock ledges and other
rock and clay bottom, in the rather swift deep current of the springs. One
rock in only 4 feet of water, downstream from South Beach, is a favorite
habitat.
CENTRARCHIDAE
Microplerus dolomieu dolomieu Lac6pede
NORTHERN SMALuLOUTH BASS
There is yet no proof that the smallmouth bass inhabits Silver Springs, but
there is some evidence that it has penetrated into these waters, which would seem
to be suitable to it as to the largemouth. A few years ago some small-
mouths were planted in Lake Weir which has been connected by canal with
Oklawaha River about 20 miles away. Since the planting and the connection were
made, several persons have reported seeing smallmouths in the Spring. On April
5, 1941, Hubbs saw a bass with darkened scales that may have been a smallmouth
bass, or an unknown Florida species, and Boyd W. Walker saw two or three
fish which he took to be M. d. dolomieu. The identifications, however, were not
all certain.
Huro salmoides (Lacepede)
LARGEMOUTH BASS
The classification of the largemouth bass has not received the revisionary
study which has been accorded the species and subspecies of Micropterus (Hubbs
and Bailey, 1940). It is therefore not possible at this time to name more precisely
the form of largemouth bass that occurs in Silver Springs. It is an intensely
marked fish, with the black band on the caudal fin rather conspicuous. The dark
band on the body tends to be broken up into blotches or short vertical bars.
Largemouth bass (Fig. 2), of small to large size, are common at various depths
in Silver Springs, and prove to be among the most interesting of the fish to the
visitors. The largest ones, some of which would seem to weigh as much as 12
pounds or more, hide under floating plants, water lettuce, logs, tree branches, and
similar shelter.
The bass in Silver Springs feed on fish and crayfish, and seem to be par-
ticularly fond of Palaemonetes paludosa (Gibbs), local known as "fairy
shrimp" (see Fig. 3). Some, becoming very tame, follow me (Allen) as I swim
along with bunches of Ludwigia pulled up from the bottom. They follow as
close as 2 feet, to snap up the shrimp as they are washed out of the plants.
The bass readily take frogs that are thrown into the middle of the pool.
126 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
Chaenobryttus coronarius (Bartram)
WARaiUTHr
As shown by Harper (1942: 50) the proper specific name of the warmouth
is coronarius rather than gulosus.
Few warmouths were observed among the thousands of centrarchid fish seen
on April 5, 1941, presumably for the reason that this species is common only
around the margin, in weeds and around logs, stumps and like cover. It is more
commonly seen at night than by day. Specimens have been collected.
Lepomis punctatus punctatus (Cuvier)
EASTERN STU3MKNOCKER
This form is accorded trinomial nomenclature because punctatus and miniatus
have been found to intergrade completely between the Mississippi River and the
Florida peninsula. This nomenclatorial conclusion has been adopted by Carr
(1936: 85). We follow Bailey (MS) in referring all the common sunfishes to a
single genus, Lepomis.
The interesting and true vernacular, stumpknockerr," refers particularly to
this species, and we urge its adoption as the "book name."
The eastern stumpknocker is characterized by the numerous small blackish
spots on its light-colored sides. In the water, however, these spots are less
conspicuous and diagnostic than are the bright silvery, creamy, or rosy margins
along the soft dorsal and caudal fins. Most of the stumpknockers stay during
the day in only moderate depths, swimming leisurely over the weeds near the
bottom. As the diving bell nears the surface stumpknockers swim against the
glass windows, as though curious. At night they come inshore, where they may
be caught easily. They also come inshore to spawn in slight excavations near the
edge of the pool, as observed on April 5, 1941. Stumpknockers were also observed
nesting in Silver Springs during June, 1943. Most of the redds were located in
groups on hard bottom, of white sand or snail shells. The guarding parents are
faithful, deserting their nests only when considerably disturbed.
Lepomis auritus (Linnaeus)
REDBREAST SUNFSH
No valid grounds have been found for the separation of solis as a distinct
southern subspecies, or specific representative of auritus.
This is one of the scarcer species of sunfish in Silver Springs. Only a few
adults were seen on April 5, 1941. However, there is no doubt as to its occur-
rence here, for the observations were clear-cut and 3 preserved specimens are
at hand.
Lepomis macrochirus mpurpescens Cope
EASTERN BLUEGILL
The modern use of the name macrochirus for the bluegill dates from Hubbs
(1935: 2-5). The form indigenous along the Atlantic coast from the Carolinas
to the Florida peninsula is quite distinct from the common bluegill, L. m.
macrochirus (which ranges into western Florida). In young and half-grown
stages it differs from the western form in having reddish fins (fading with age),
and in the broader, more even bars and higher fin spines (as shown in the figure
by Fowler, 1940: 14, fig. 19). These characters persist in the adult but become
less obvious with age. The adults are more purplish (this is particularly true of
the breeding males), and frequently have a cream-colored bar across the nape,
connecting with a large blotch or wide bar of the same color across the opercular
FISHES OF SILVER SPRINGS, FLORIDA
region (Archie F. Carr, Jr., tells us that the "crackers" insist that the bluegills
so marked are a different kind, and "eat better.") The anal soft rays, particu-
larly in Florida and southern Georgia, average more numerous than in typical
macrochirus (more often 12 than 11, rather than typically 11, rarely 10 or 12).
The "Georgia bluegill" or "Georgia brim," as it is known to fish culturists, retains
these characters where is has been introduced, as in Texas, and in the Yaqui
River system of Sonora. It proves to be a larger, faster-growing and more
resistant fish than the common bluegill. It is no doubt a genetically distinct and
recognizable form. It may even prove specifically different.
The characters here ascribed to L. m. purpurescens were evident in the many
bluegills seen in Silver Springs on April 5, 1941. It is the dominant sunfish in
these waters. The bluegills (Figs. 2-4) occur more often in the open water and
in the deeper spring holes than do the other species. They have been trained to
come to food (bread) so as to be easily seen in large numbers by the visitors.
Next to the catfish, they are probably the most conspicuous fish in the Springs.
Lepomis nicrolophus micolophus (Ginther)
EASTERN SHELLCRACKER
The application of the specific name microlophus to this sunfish is based on
the discussion by Hubbs (1935: 5-9). The vernacular, "shellcracker," is par-
ticularly appropriate and pleasing. This is the largest sunfish in Florida, and
the most highly regarded by anglers. Although it feeds largely on snails, it may
be caught on worms.
The shellcracker, recognizable by the red "ear-spot," is a common species
in Silver Springs, in both deep and shallow water. On April 5, 1941, most of
those seen were large adults. One is shown in Fig. 3. Some of the males, in
water of considerable depth, were guarding nests which were conspicuous, bowl-
shaped depressions in dense stands of the coarse Sagittaria. They are faithful
guardians. Several adult specimens from the Springs have been preserved.
Enneacanthus, species
Specimens of Enneacanthus have been collected but none have been ex-
amined by the senior author to make sure of the specific identification. E. obesus
and E. glorious both occur in Florida.
Pomoxis nigro-maculatus (LeSueur)
BLACK CRAPPIE
The use of the name P. nigra-maculatus in place of P. sparoides was validated
by Bailey (1941).
No crappies were seen in Silver Springs during the observations of April 5,
1941, but an adult specimen speared by Allen is in the University of Michigan
collection, and another taken May, 1936, is in the National Museum.
ELASSOMIDAE
Elassoma evergladei Jordan
EVERGLADES PYGMY SUNTISH
This species, one of the smallest of all the spiney-rayed vertebrates, is rep-
resented in the preserved collection at hand from Silver Springs. Another speci-
men, of 13 mm., was taken on August 29, 1935.
It is one of the species which frequent dense weeds in shallow water.
128 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
Mugil curema Valenciennes
WHITE MULLET
A mullet labelled as collected in Silver Springs by Hildebrand and Allen on
August 30, 1935, is an example of this species. In general the white mullet is less
common in fresh waters than is the striped mullet, but it is known to penetrate at
times far inland (as in Texas). This white mullet may have been a stray, or may
have been mislabelled. The record needs confirmation.
Mugil cephalus Linnaeus
STRIPED MULLET
Most if not all of the mullets that now occur in Silver Springs, almost cer-
tainly all of the larger adults, are of this species (Figs. 1, 2 and 4). The dark
stripes along the scale rows are evident in life. The depth of the body was seen
to vary greatly. Many of those observed on April 5, 1941, were feeding. They
shook their heads from side to side as they browsed over the rock bottom and
particularly up the broad blades of the Sagittaria. As they continually feed in
this fashion they expel clouds of dirt from their mouths.
These very active fish travel in schools of about a dozen, most commonly in
the deep pools. They frequently jump out of water. They are also active at
night but apparently do not feed then. They are not bothered by other fish.
Extremely large mullet are never seen in Silver Springs.
ATHERINIDAE
Labidesthes sicculus vanhyningi Bean and Reid
SOUTHEASTERN BROOK SILVERSIDE
This is a rather poorly defined geographical subspecies. One character is the
large eye.
No brook silverside were observed on April 5, .1941, but there are specimens
in the local collection. It is to be watched for at the surface.
ACHIRIDAE
Trinectes maculatus fasciatus (Lacepede)
SOUTHERN HOGCHOKER
The generic and the specific nomenclature of this common American sole
were treated by Hubbs (1932). The name for the species should stand as
Trinectes maculatus (Bloch) rather than as Achirus fasciatus Lac6pede. The
northern form, T. n. maculatus, is distinguished from the Gulf of Mexico sub-
species (commonly called brown) chiefly in having the blind side of the adult
conspicuously spotted. The proper subspecies name for the southern form could
not be determined until a good series of topotypes of Achirus fasciatus, from
Charleston, South Carolina, became available. This desideratum was adequately
supplied by E. Milby Burton of the Charleston Museum. Some Charleston
specimens are more or less weakly spotted on the eyeless side, but the majority
are unpigmented on this side. Achirus fasciatus was therefore based on inter-
grades, but on a population approaching the southern form much the more
closely. For this reason the name fasciatus is applied to the southern subspecies.
Another feature of T. m. fasciatus is its frequent occurrence in lakes and
streams far inland from the saline or even tidal influence of the sea. It is
therefore not at all surprising to find that this sole occurs in Silver Springs. Sev-
eral were seen here on April 5, 1941, by members of the American Society of
Ichthyologists and Herpetologists, and there is one specimen in the preserved
collection. It was caught by hand at a depth of 25 feet, in Big Spring.
FISHES OF SILVER SPRINGS, FLORIDA
LITERATURE CITED
BAILEY, REEVE M.
1941 The scientific name of the black crappie. Copeia, 1941: 21-23.
BANGOCAM, RALPH V.
1942 Parasites of fresh-water fish of southern Florida. Proc. Fla. Acad.
Sci., 5, "1940": 289-307.
CARR, A. F., JR.
1936 A key to the fresh-water fishes of Florida. Ibidem: 1: 72-86.
COPE, E. D.
1865 Partial catalogue of the cold-blooded Vertebrata of Michigan. Part
II. Proc. Acad. Nat. Sci. Phila., 1865: 78-88.
DE KAY, JAMES E.
1942 Zoology of New-York or the New-York Fauna; comprising detailed
descriptions of all the animals hitherto observed within the State of New
York, with brief notices of those occasionally found near its borders,
and accompanied by appropriate illustrations. Part IV. Fishes. In:
Natural History of New York. Albany: i-xv, 1-415, pls. 1-79.
DUMERt., AUG.
1870 Histoire naturelle des Poissons ou ichthyologie g6enrale. Tome Second
Ganoides, Dipn6s, Lophobranches. Paris: 1-624, and Atlas: 9-12, pls.
15-26.
FowLE, HENRY W.
1911 Notes on chimaeroid and ganoid fishes. Proc. Acad. Nat. Sci. Phila.,
1910: 603-12, pl. 38.
1940 A collection of fishes obtained on the West Coast of Florida by Mr. and
Mrs. C. S. Chaplin. Ibidem, 92: 1-23, pl. 1, figs. 1-37.
1941 A collection of fresh-water fishes obtained in Florida, 1939-1940, by
Francis Harper. Ibidem: 227-44, figs. 1-13.
GARMAN, S.
1895 The cyprinodonts. Mem. Mus. Comp. Zool., 19: 1-179, pls. 1-12.
HAhLMETT, F. S. AND D. W.
1939 Proportionate length growth of gar (Lepisosteus platyrhincus De Kay).
Growth, 3: 197-209, charts 1-5.
HARPER, FRANCIS
1942 The name of the warmouth. Copeia, 1942: 50.
HuBBs, CARL L.
1932 The scientific name of the common sole of the Atlantic Coast of the
United States. Proc. Biol. Soc. Wash., 45: 19-22.
1935 The scientific name of two sunfishes, Helioperca macrochira (Rafin-
esque) and Eupomotis microlophus (Giinther). Occ. Pap. Mus. Zool.
Univ. Mich., 305: 1-12.
1940 Speciation on fishes, Am. Nat., 74: 198-311.
HBBss, CARL L., AND BAILEY, REEVE M.
1940 A revision of the black basses (Micropterus and Huro), with de-
scriptions of four new forms. Misc. Publ. Mus. Zool. Univ. Mich., 48:
1-51, pls. 1-6, maps 1-2.
130 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
HUBBS, CARL L., AND LAGLER, KARL F.
1943 Annotated list of the fishes of Foots Pond, Gibson County, Indiana.
Inv. Indiana Lakes and Streams, 2, "1942": 73-83.
JORDAN, DAVID STARR, AND EVERANN, BARTON WARREN
1896 The fishes of North and Middle America: a descriptive catalogue of
tht species of fish-like vertebrates found in the waters of North Amer-
ica, north of the Isthmus of Panama. Bull. U. S. Nat. Mus., 47 (1):
i-lviii, 1-1240.
KUTNE, EUGENE R.
1939 A guide to the fishes of Tennessee and the mid-South. Tenn. Dept.
Cons.: 1-124, figs. 1-81.
PARR, ALBERT EIDE
1927 A contribution to the theoretical analysis of the schooling behavior of
fishes. Occ. Pap. Bingham Oceanogr. Coll., 1: 1-32, figs. 1-2.
REIGHARD, JACOB
1903 The natural history of Amia calva Linnaeus. Mark Anniversary Volume,
Art. 4: 57-109, pl. 7, fig. A.
NEEDLE RUSTS OF PINE TREES IN FLORIDA
CAUSED BY COLEOSPORIUM SPECIES
GEORGE F. WEBER
Agricultural Experiment Station, University of Florida
Needle rusts are commonly found and observed in Florida on the
different species of pines so abundant in the State. The brilliant orange
color of the maturing aecia on the green pine foliage frequently attracts
attention to the rusts caused by Coleosporium spp. No one has
ventured a theory as to the origin of these needle rusts, but it is
thought that they are indigenous to Florida. Davis (1913) has shown
how a certain species was probably introduced into Wisconsin.
Published information concerning these rusts is not plentiful, and
as indicated by Dearness (1939) and Hedgecock (1939a) it is not
entirely usable either by the amateur or the professional, unless the
latter is an expert. Definite identification in the field of all the fungi
causing these needle diseases is almost impossible. There are a number
of species of Coleosporium occurring in Florida. Being similar in their
manifestations, and as a rule only identifiable by microscopic examina-
tion, in this paper they will be treated as a group in relation to the
diseases which they cause, but separately with respect to their tax-
onomic characters.
The information here presented has been obtained from observa-
tions and collections made by the author and associated co-workers
over almost a score of years in Florida, and from the literature, to
which Hedgecock and his collaborators (1913-1939) have been the
principal contributors. Dr. Hedgcock has accompanied the author
and his colleagues on some of their collecting trips.
THE DISEASE
Needle rusts have been collected in Florida in widely separated
localities. From the available information one suspects their occur-
rence to be more or less general over the entire state wherever pine
trees are growing. There are certain large areas in south Florida,
possibly 30 miles across, where no species of pine occurs. South of a
line drawn from Ft. Myers to Palm Beach, Pinus clausa and P. caribaea
are the only native pines present. In general, needle rust disease seems
to be scarcer toward the tropics and more abundant northward, though
it has been reported by Stevens (1917) from Puerto Rico. In southern
Florida, judging from scattered collections, it is rather uncommon, but
throughout the central and northern parts of the state needle rust is
very plentiful. Certain species of Coleosporium have been collected
131
132 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES ol. 6,
Nos. 3-4, 1943
on a half dozen species of Pinus, and one or more species of the fungus
can be found infecting pines wherever these trees occur in Florida.
At present the following eight species of pine are considered in-
digenous to Florida: Pinus australis (longleaf pine), P. caribaea
(southern slash pine), P. clausa (sand pine), P. echinata (shortleaf
pine), P. glabra (spruce-pine), P. palustris (northern slash pine), P.
serotina (pond pine or black pine), and P. taeda (loblolly or oldfield
pine). Reports of collections of diseased needles by Hedgcock (1933,
1939b) show that each of the pines has been found infected by one
or more Coleosporium species. In all of the collections made by
Hedgcock and by the author and his associates, only a single species
of Coleosporium has been found infecting Pinus clausa; but this same
fungus has also been found in Florida on P. australis, P. caribaea, P.
glabra, P.palustris, P. serotina, and P. taeda... In contrast, another
of the needle rusts, which was found on Pinus glabra near Gaines-
ville, Florida, and described by Hedgcock and Hunt (1917g), has
never been found anywhere else in the world, although it still occurs
annually at the type locality, and has there since been found on Pinus
taeda.
The pines are more or less infected by one or more of these fungi
wherever they grow. The disease may be scarce, as intimated by
Graves (1914), or very prevalent even on commercial plantings as
reported by Guterman (1935).
All of the needle rusts occurring in Florida are heteroecious. The
pycnia and aecia are found on Pinus spp., and the uredinia and telia on
alternate hosts which are usually broad-leaved plants, the majority of
them belonging to the Carduaceae.
Boyce (1938) stated that he considered needle rusts most active
(or severe) on young pine trees in the seedling and sapling stages.
Hubert (1931) stated that young seedlings are sometimes severely in-
fected and suffer some stunting but that death of the plant due di-
rectly to attacks of the disease has not been noted. Rankin (1918)
noted that the needle rusts occasionally caused defoliation. In what
follows nursery stock is taken as including plants 1 year old or less;
seedling ranges to 3 feet high, a sapling to 4 inch diameter, and
a tree is more than 4 inches in diameter. Observations by the author
in Florida have shown that seedlings are certainly more heavily in-
fected than saplings or trees of the same species. The disease has
been found occasionally in nursery stock but not in sufficient quantity
to be recorded as more than a trace. This is probably accounted for
by the fact that the few nurseries visited were located in extensively
cultivated areas where the fungus did not reproduce plentifully, pos-
sibly because of the lack of alternate hosts. Also, artificial protection
NEEDLE RUSTS OF PINE TREES IN FLORIDA
(such as cloth covering and fungicides) may have been a contributing
factor. The young plants in nature, less than 1 year old, are seldom
more than a third as tall as nursery stock and have seldom been found
infected. In contrast, however, seedlings between 1 and 2 feet tall
are frequently very heavily infected in the vicinity of Gainesville.
Seedlings of Pinus taeda growing in a location very favorable for the
development of the disease have been killed over a two year period.
Others were very badly stunted by the disease. Seedlings of Pinus
australis between 1 and 2 feet tall in this vicinity have been found in
the spring showing the rust infection on 95% of the needles which
resulted in no serious effects during the summer, other than a possible
temporary stunting. Continued observations during the following
spring of these identical seedlings showed no infection. This area
was burned in the interim and all infections on the pine and alternate
host were apparently destroyed. Seedling infection is widespread,
commonly found and occasionally severe enough to be considered of
economic importance. The disease on saplings is seldom of import-
ance although almost every plant under ten feet in height that possesses
low spreading branches shows some infection. The high-headed sap-
lings and pine trees show little needle rust, and under the most fa-
vorable circumstances almost none.
Symptoms.-There are several species of Coleosporium that cause
infection of the needles of pine trees. The symptoms produced are
generally very similar; slight exceptions to this statement will be men-
tioned only when pertinent.
The principal symptom that characterizes the disease is the ap-
pearance of yellow- to orange-colored areas, several millimeters to
several centimeters in length, located anywhere on the needle from
fascicle sheath to the tip. The infection is less frequent at the apex
than elsewhere. A single infection apparently causes one colored band
across the otherwise healthy green leaf. Several such etiolated areas
may occur on a single needle, caused by infections by more than one
species of the fungus. On a particular host the color of the infected
tissue is consistently the same for each of the Coleosporium spp.
There are slight differences in this respect between hosts, and also
between different fungus species infecting the same host. These
colored areas on the needles are conspicuous and easily seen but
cannot be considered proof of rust infections unless upon closer ex-
amination pycnia or aecia of the fungus are found. These structures
not only indicate that the infection has been caused by a Coleosporium
sp., but are safe diagnostic characters for determining the exact
Coleosporium sp. involved. The pycnia may appear on pine needles
in central Florida anytime after early January, being most common
134 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
during the last half of February. They are small, circular to oval,
slightly raised, rounded, smooth surfaced, reddish-orange to black
areas, 1-4 mm. in diameter. The areas may be in single, continuous
or intermittent, irregular lines, in several parallel short lines, or in
somewhat irregular clusters on the yellowish areas, all usually on one
side of the needle. The pycnia are followed in appearance by the
aecia, which become evident three to six weeks later in central
Florida. The aecia break through the epidermis of the leaf opposite
the pycnia. This stage of the fungus is very conspicuous. The
aeciospores contained within the peridial walls are bright orange in
color. The peridial walls are several millimeters in height. The
aecia vary considerably among the Coleosporium spp. in size, color
and shape, but each species is rather uniform on the various Pinus spp.
The symptoms found on the pine needles, including the color of the
spot and the size, color and location of the pycnia and aecia, are
diagnostic characteristics of the needle rust diseases.
The symptoms of the disease on the alternate hosts are much less
conspicuous. The uredinia and telia are similar and appear as in-
conspicuous yellow, cushion-like areas on the leaf surface of herbs
and weeds. These infections cause yellow areas to appear on the
foliage and frequently result in the death of these plants.
THE FUNGI
Taxonomy.-Common names of the diseases found in literature are
leaf rust, needle rust or pine needle rust.
The genus name Coleosporium was first used by Leveille in 1847
in reference to the fungi causing needle rust. Since that time new
species have been discovered and named. Saccardo (1887-1924) lists
54 described species. Hubert (1931) suggests about 80 and Ludwig
(1915) reports 25 species. Arthur (1934) published descriptions of
20 Coleosporium spp. on the genus Piits in the United States and
Canada. Hedgcock after examining all available collections verified
23 species for the United States; his earlier published key (1928) con-
tains 18 for this area. Boyce (1938) reports 16 species, and Rhoads
(1918) presents notes on 14 species. After carefully checking all au-
thentic reports of Florida collections supplemented by our staff collec-
tions it is found that there are 10 Coleosporium spp. in the state. They
are as follows:
NEEDLE RUSTS OF PINE TREES IN FLORIDA 135
TABLE I
List of Pine Needle Rusts Found in Florida, Including Name of Fungus, Pine
and Alternate Hosts
Hosts
Fungus O, I | II, III
Coleosporium apocynaceum Cke. Pinus australis Amsonia ciliata
Pinus palustris
Pinus taeda
Coleosporium delicatulum (A. & K.) Pinus australis Euthamia caroliniana
H. & L. Pinus palustris Euthamia leptocephala
Pinus serotina Euthamia minor
Pinus taeda
Coleosporium elephantopodis (S.) Pinus australis Elephantopus carolinianus
Thum. Pinus glabra Elephantopus elatus
Pinus palustris Elephantopus nudatus
Pinus serotina Elephantopus tomentosus
Pinus taeda
Coleosporium heterotheca Hedge. Pinus australis Ileterotheca subaxillaris
Coleosporium ipomoeae (Schw.) Burr. Pinus australis Colonyction aculeatum
Pinus palustris Ipomoea carolina
Pinus serotina Ipomoea pandurata
Pinus taeda Ipomoea pes-capreae
Ipomoea speciosa
Ipomoea triloba
Pharbitis barbegina
Pharbitis cathartica
Pharbitis hederacea
Coleosporium laciniariae Arth. Pinus australis Laciniaria chapmanni
Laciniaria elegans
Laciniaria graberi
Laciniaria gracilis
Laciniaria laxa
Laciniaria pilosa
Laciniaria scariosa
Laciniaria tenuifolia
Coleosporium minutum H. & H. Pinus glabra Adelia ligustrina
Pinus taeda
Coleosporium. solidaginis (Schw.) None in Florida Aster puniceus
Thum. Chrysopsis scabrella
Solidago brachyphylla
Solidago chapmannii
Solidago fistulosa
Solidago puberula
Solidago rugosa
Solidago sempervirens
Solidago stricta
Coleosporium vernoniae B. & C. Pinus australis Vernonia augustifolia
Pinus clausa Veronia blodgettii
Pinus echinata Vernonia flassidifolia
Pinus glabra Vernonia oligantha
Pinus palustris Vernonla ovalifolia
Pinus serotina
Pinus taeda
Coleosporium viguierae D. & H. None in Florida Verbesina lacinata
136 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
It would be very helpful to include here a description of each of
the 10 Coleosporium species occurring in Florida, but space does not
permit. Detailed descriptions of the fungi with synonyms, host and
geographical range are published by Arthur (1934); Hedgcock (1913,
1916, 1939a); Hedgcock and Long (1913, 1917); Hedgcock, Bethel
& Hunt (1919); Hedgcock, Hunt & Hahn (1920); Rhoads, et al
(1918); and Weir and Hubert (1916).
As indicated in the table above no authentic specimen of either
Coleosporium solidaginis or C. viguierae has been found on any pine
in Florida. The 10 species found in the state are reported from the
alternate hosts. The data accumulated through artificial inoculations
by Hedgcock and Hunt (1917a-1933) show that the range of host
plants is limited to certain families and genera. Consequently the iden-
tification of rusts when either the uredinia (II) or telia (III), or both
are present is reliable.
Arthur's key (1934) to rusted alternate hosts, abridged to include
only Florida species, and slightly revised, is as follows:
Alternate hosts belonging to family Oleaceae-Coleosporium minutum H. & H.
Alternate hosts belonging to family Apocynaceae-C. apocynaceum Cke.
Alternate hosts belonging to family Convolvulaceae-C. ipomoeae (Schw.) Burr.
Alternate hosts belonging to family Carduaceae
Tribe Vernonieae
Genus-Vernonia-C. vernoniae B. & C.
Genus-Elephantopus-C. elephantopodis (S.) Thum.
Tribe Eupatorieae-C. laciniariae Arth.
Tribe Astereae
Genus-Euthamia-C. delicatulum (A. & H.) H. & L.
Other genera-C. solidaginis (S.) Thum. (including C. heterothecae H.
& H.)
Tribe Heliantheae-C. verbesinae D. & H.
The above key is based entirely on the infected alternate host and
urediniospores produced thereon. It contains Coleosporium solidaginis,
C. verbesinae and C. heterotheca none of which have been found to
produce the peridermium stage, or pycnia and aecia, on pines in
Florida. Arthur (1934) includes the latter under C. solidaginis.
On the other hand only 7 Coleosporium spp. have been definitely
identified as parasitizing Pinus spp. in Florida. The various diseases
on pines show the pycnia (0) and aecia (I) stages of the different
fungi. These stages are very consistent and uniform for each fungus
on each host and, except to the expert, are likewise quite similar to
each other. Hedgcock (1928) has worked out a key by which he is
able to distinguish the species in the United States. Herewith is a
revised and abridged version of the part of this key containing Coleo-
sporium spp. on pine needles, applicable only to Florida.
NEEDLE RUSTS OF PINE TREES IN FLORIDA
I. Dehiscence of the peridium irregularly apical and longitudinal.
A. Aecia very conspicuous usually in single extended rows.
1. Aecia irregularly flattened rhomboidal, pycnia orange-rufous, auburn
or chestnut colored.................................... Coeosporium vernoniae B. & C.
2. Pycnia olivaceous black to brownish black.
Coleosporium elephantopodis (Schw.) Thum.
B. Aecia small and inconspicuous, usually clustered or in short rows, pycnia
olive to olivaceous black.
1. Pycnia usually in single extended line.
Coleosporium iponoeae (Schw.) Burr.
2. Pycnia usually clustered.............................Coleosporium laciniariae Arth.
II. Dehiscence of the peridium circumscissile.
A. Aecia large and conspicuous on yellow chlorotic areas. Pycnia hazel to
chestnut brown.............................................. Coleosporium apocynaceum Cke.
B. Aecia small and inconspicuous.
1. On reddened chlorotic areas, pycnia orange chrome to English red.
Coleosporium delicatulum (A. & K.) H. & L.
2. On yellowish chlorotic areas, pycnia tawny to buckthorn brown.
Coleosporium minutum H. & H.
This key is workable for Florida needle rusts after a little experi-
ence in the field with color variations and other factors.
Morphology.-These rusts are all of the Eu-type in which the 0,
I, II, III stages are present during each season. They are heteroecious
in host relationship, with the pycnial and aecial stages appearing on
the pine needles, and the uredinial and telial stages appearing on
various herbaceous dictoyledonous plants.
The pycnia are subepidermal, mound-shaped to flat, oval to linear,
paraphysate, dehiscing by a slit. They may be located in lines con-
tinuous or intermittent, or in several short parallel rows, or in groups.
In color they vary considerably, from yellow and orange to olive or
blackish; and they range from 0.5 to 2.0 mm. in length.
The aecia are foliicolous, amphigenous, hypophyllous or epiphyllous,
erumpent, with colorless peridium prominent and of various shapes and
sizes; dehiscing slit is longitudinal at crest in smooth or rough line or
circumscissile. The aeciospores are ellipsoidal or globoid, with ver-
rucose colorless walls. The contents are usually orange, and en masse
cause the aecia to be conspicuous because of their bright color.
The uredinia are erumpent, pulverulent, without peridia, round or
oval, raised, usually small, orange or yellow when fresh. On either
surface of the leaves, usually more numerous on lower surface. Ure-
diniospores ellipsoid to globoid or obovate, catenulate, of various
sizes, with colorless thin walls, finely or strongly verrucose.
The telia are indehiscent except through weathering, waxy, or gel-
atinous according to Cummins (1938), becoming gelatinous upon ger-
138 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol 6,
Nos. 3-4, 1943
mination, usually on lower surface of leaves, scattered or clustered,
and of variable size and color. The teliospores are sessile or obscurely
catenulate, one-celled, cylindric, thin, smooth, the colorless walls more
or less thickened at round or acute ends.
These rusts are all obligate parasites, requiring both of the hosts to
complete their lie cycle in any single season. They cannot be grown on
sterile nutrient decoctions, but can be cultured on either host in
controlled environments such as greenhouses. The aeciospores and
urediniospores germinate in water when mature, especially in the
presence of host materials, and are capable of causing infection within
24 hours under conditions favorable for their growth. The teliospores
germinate in place, freeing the basidiospores in abundance. These
spores are small and easily carried by air currents to the pine.
PATHOGENICITY: It is primarily through the extensive inoculation
experiments conducted by Hedgcock (1933) and Hedgcock and Hunt
(1917a, c, d, f, g), that we know of the extensive host range of these
various species of rust. The data definitely show that these fungi
are highly pathogenic on one or more of the native pines of Florida.
Likewise they have shown that the uredinial and telial stages of a single
species are found on as many as nine herbaceous hosts in the state.
These experiments were usually verifications of observational data,
substantiated by collections in the field, although the inoculations
have extended the host range beyond those found in nature.
THE HOSTS AND FUNGI IN RELATION TO THE DISEASE
The seasonal development of the disease is dependent on the prox-
imity of the hosts. It should also be borne in mind that the advance
of spring in the various regions of the United States from South to
North influences markedly the dates of appearance of the disease.
Hedgcock (1939b) states that the pycnia of Coleosporium species
appear in central Florida in December and mature in January to
February and that they precede the aecia by at least a month. The
writer has not determined the limits of early pycnia appearance, but
on February 2, 1941, aecia were collected on Pinus taeda a few miles
from Gainesville that already had completely shed their aeciospores.
During the previous year their first appearance was about 5 to 6 weeks
later. Cool weather retards their development. In relation to each
other, however, certain ones are always the first to appear and others
are consistently later. The dates of appearance of the first and the
last may cover a period of 6 to 8 weeks from February 1 to April 1.
The aeciospores are shed during this period and disseminated by the
wind and rain to plants in the immediate vicinity. The herbaceous
hosts of several species usually show new growth during later Jan-
NEEDLE RUSTS OF PINE TREES IN FLORIDA
uary. Many of these alternate host plants continue growth in the
rosette stage throughout the winter and have been found showing ma-
ture uredinia spores every month of the year. Thus they may continue
to develop during the entire season without utilizing the stages on pine.
This condition was found by Maines (1916) to exist north of the
Ohio River. The uredinial stage continues during the summer and the
telial stage develops on the older foliage in summer and fall early
enough to infect the needles of the young pine seedlings. This infec-
tion develops slowly during the fall and winter. The pycnia appear
in January and February on these infections followed by the aecia in
February and March. The fungus overwinters in several ways, namely
(a) on the pine needles infected in the summer and fall, (b) in the
telia on the alternate host producing basidiospores in the spring, and
(c) in the uredinia on the alternate host where spores are produced
throughout the year. Eriksson (1921), in addition to these, sus-
pects a mycoplasm symbiosis.
The infected herbaceous alternate hosts are the source of the
inoculum in the summer and fall with which the pines are infected.
The sources of the inoculum in the spring by which the herbaceous
hosts are inoculated are the aecia on the pine needles. The aeciospores
are not capable of reinfecting the needles of pines, but must infect the
herbaceous plants. The teliospores, upon germination produce basidio-
spores. Basidiospores cannot reinfect the herbaceous host upon which
the teliospores were produced but can infect only the pine. The
urediniospores very readily reinfect the herbaceous host upon which
they were produced. The spores produced on each of the hosts are
disseminated by the wind. It is known that the aeciospores may
withstand adverse conditions and over extensive periods travel long
distances before losing their viability, whereas the basidiospores are
delicate and short-lived and are disseminated usually less than a mile
before dying.
The time of infection of both hosts is very shortly after the first
mature spores are produced. The aeciospores may infect the herba-
ceous hosts any time during February, March and April and these
plants may suffer additional infection by urediniospores which are
produced on these plants after a 2 to 3 weeks incubation period
throughout the summer. The basidiospores as a source of inoculation
cause infection primarily through late summer and fall. Infection
of the herbaceous hosts in all instances is direct; that is the germina-
tion tubes of the spores, after a short period of growth, grow directly
through the epidermis, although stomatal entrance may be possible.
Johnson (1934) observed no tropic responses. Penetration of the pine
needle is generally considered to take place through the stomata.
140 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
The production of aecia and aeciospores may occur several months
after infection, appearing usually scattered, but found by Spaulding
(1917) to be as many as 20 per needle. They appear only in the
spring following infection of the previous year. The production of
uredinia occurs less than a month after infection of the alternate host
by aeciospores. Usually the fungus does not extensively invade the
host tissue and only small areas of needles show any yellowing, result-
ing apparently from invasion and not from toxic effects as produced
by many parasites.
CONTROL
The primary factor in control of the disease on pines would be
the elimination of the herbaceous alternate host. This can most easily
be done by planting pine seed beds in cultivated fields as far as possible
from woodlands, roadsides and fence rows where alternate hosts usually
grow. Ordinarily the protection given seed beds for control of other
diseases would be sufficient for this trouble. No control is suggested
for transplanted seedlings, except in plantings used as ornamentals,
where the greater value of the plants warrants spraying with a copper
fungicide.
NEEDLE RUSTS OF PINE TREES IN FLORIDA
LITERATURE CITED
ARTHUR, J. C.
1934. Manual of the Rusts in the United States and Canada. Perdue Res.
Fund, Lafayette, Indiana. 438 pp.
BOYCE, J. S.
1938. Forest Pathology. McGraw Hill Book Co., N. Y. 600 pp.
CUMaMNS, GEORGE B.
1938. A new microcyclic Coleosporium on timber and pinion trees. Phy-
topath., 28: 522-23.
DAVIS, J. J.
1913. The introduction of a European pine rust into Wisconsin. Phy-
topath., 3: 306-8.
DEARNESS, JAKOB
1939. Notes on Coleosporium crowellii. Plant Disease Reporter, US.D.A.,
23: 134.
ERIKSSON, JAKOB
1921 The mycoplasm theory-is it dependable or not? Phytopath., 11:
385-88.
GRAVES, ARTHUR H.
1914. Notes on diseases of trees in the Southern Appalachians, II. Phy-
topath., 4: 5-10.
GUTERMAN, C. E. F.
1935. Control of Aster leaf rust. (Abstract). Phytopath., 25: 17-18.
HEDGCOCK, GEORGE G.
1913. Notes on some western Uredineae which attack forest trees, II.
Phytopath., 3: 15-17.
1916. Identity of Peridermium montanum with P. acicolum. Phytopath.,
6: 64-67.
1928. A key to the known aecial forms of Coleosporium occurring in the
United States and a list of the host species. Mycologia, 20: 97-100.
1933. Notes on the distribution of some species of CoSeosporium in the
United States and adjacent regions. Plant Disease Reporter, U.S.D.A.,
17: 20-27.
)939a. Notes on the occurrence of Coleosporium crowelii Cum. in the
United States. Plant Disease Reporter, US.DA., 23: 133-134.
1939b. Notes on the occurrence of Coleosporium in the southeastern United
States during 1938 and 1939. Plant Disease Reporter, U.S.D.A., 23:
268-277.
HEDGCOCK, GEORGE C., AND N. R. HUNT.
1917a. A Peridermium belonging to Coleosporium ipomoeae. (Abstract).
Phytopath., 7: 67.
1917b. A Peridermium belonging to Coleosporium terebinthinaceae. (Ab-
stract). Phytopath., 7: 67.
1917c. An alternate form of Coleosporium helianthi. (Abstract). Phy-
topath., 7: 67-68.
1917d. Notes on some species of Coleosporium. (Abstract). Phytopath.,
7: 68.
1917e. Some new hosts for Coleosporium solidaginis (Schw.) Thum. (Ab-
tract). Phytopath., 7: 68.
142 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
1917f. Some new hosts for Coleosporium inconspicuum. (Abstract). Phy-
topath., 7: 67-68.
1917g. New species of Peridermium. Mycologia, 9: 239:342.
1922a. Notes on some species of Coleosporium, I. Mycologia, 14: 244-257.
1922b. Notes on some species of Coleosporium, II. Mycologia, 14: 297-310.
1933. Notes on some species of doleosporium, III. Mycologia, 25: 392-396.
HEDGCOCK, GEORGE G., AND W H. LONG.
1913. Note on cultures of three species of Peridermium. Phytopath., 3:
250-251.
1917. The aecial stage of Coleosporium elephantopodis (Sch.) Thum. (Ab-
stract). Phytopath., 7: 66-67.
HEDoGocx, GEORGE G., E. BETHEL, AND N. R. HUNT.
1919. Notes on the rusts of pinion pines. (Abstract). Phytopath., 9: 53.
HEDGcocx, GEORGE G., N. R. HUNT, AND G. G. HAuN.
1920. New species and relationships in the genus Coleosporium. Mycologia,
12: 182-98.
HUBERT, E. E.
1931. Outlines of Forest Pathology. John Wiley & Co., N. Y., 543 pp.
(pp. 253-254).
JOHNsoN, THORVALDUR.
1934. A tropic response in germ tubes of urediniospores of Puccinia graminis
tritici. Phytopath., 24: 80-82.
LUDWIG, C. A.
1915. Notes on some North American rusts with Caeoma-like sor. Phy-
topath., 5: 273-281.
MARINES, E. B.
1916. The wintering of Coleosporium solidaginis. Phytopath., 6: 371-372.
RANKIN, W. H.
1918. Manual of Plant Diseases. Macmillan & Co., N. Y., 398 pp.
RHOADs, A. S., GEORGE G. HEDGCOCX, E. BETHEL, AND CARL HARTLEY.
1918. Host relationships of the North American rusts other than Gymno-
sporangium which attack conifers. Phytopath., 8: 309-352.
SACCARDO, P. A.
1887-1924. Sylloge Fungorum. Vols. 7, 9, 11, 22.
SPAULDING, PERLEY.
1917. Needle rusts on Pinus resinosa. Phytopath., 7: 225.
STEVENS, F. L.
1917. Noteworthy Porto Rican plant diseases. Phytopath., 7: 130-134.
WEIR, JAMES R. AND E. E. HUBERT.
1916. Inoculation experiments with Periderminum montanum. Phytopath.,
6: 68-70.
THE LOWER VERTEBRATE FAUNA OF THE
WATER HYACINTH COMMUNITY IN
NORTHERN FLORIDA
COLEMAN J. GOING
University of Florida
Although the water hyacinth, Piaropus crassipes (Mart.) Britton,
is extremely abundant and widespread in the southeastern states, very
little attention has been given to the fauna that is now associated
with its luxuriant growth. Many bodies of water in this region support
large masses of the hyacinth, and some are entirely covered by this
floating plant. In the lakes and larger sluggish rivers it drifts to and
fro with the wind, and it often completely blankets the small ponds,
canals, and ditches.
The roots of Piaropus develop directly in proportion to the depth
of the water. In the shallow water the roots are short, but where
the water is deeper they may extend downward to a depth of three
feet. These large masses of filamentous roots, shaded as they are by
the floating portions of the plants, provide an extensive cover for
numerous animals. Moreover, there is a huge and continuous ac-
cumulation of hyacinth detritus on the bottom of all hyacinth-
supporting bodies of water, and this detritus not only modifies the
physical conditions on the bottom but, where the plants are thick, also
supplies an enormous mass of organic food. At Newman's Lake, for
example, although the greater part of the lake is open water and only
the margins are typically covered with hyacinths, a bottom sampling
made with an Eckman dredge at nearly any point in the lake will be
found to contain many fragments of dead hyacinths.
Professor M. D. Cody, of the University of Florida, informs me
that this plant was first brought into the United States in 1835 by
the Venezuelan delegates to the Centennial Exposition at New Orleans.
Visitors to the Exposition received these plants as souvenirs and, by
carrying them home, introduced them to many parts of the Southeast.
It was not until about 1840 that the plant became established in
Florida. After its first establishment it was carried to different parts
of the state by cattlemen who believed the hyacinth was of value as
cow feed.
The abundant and luxuriant growth attained by the hyacinths has
attracted a great number of aquatic and semiaquatic animals which
have become associated with it. Among the invertebrates characteris-
tic of the hyacinth community may be mentioned oligochaete worms
143
144 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
(Dero), jawed leeches (Gnathobdellida), amphipods (Hyalella), a
crawfish (Procambarus fallax), water spiders (Dolomedcs), and midges
(Chironomus).
In the course of other studies, I have kept records of the fishes,
amphibians, and reptiles found in this community. Most of these
records are based on about two years' active and intensive collecting,
while random notes have accumulated over a period of some six
years. In addition, records from the University of Florida Department
of Biology collections have been included in cases where the speci-
mens are known to have been collected among water hyacinths. My
absence from Florida during the summers has prevented me from
collecting during the months of June, July, and August. All of the
records given here are for the vicinity of Gainesville, Alachua County,
Florida. Representatives of all species have been deposited in the
Carnegie Museum or in the Department of Biology, University of
Florida, but it did not seem advisable to collect large series of the
abundant species. However, records of all of the rarer and less well
known species are based upon laboratory-identified material. Un-
doubtedly some of the species less commonly associated with the
hyacinths have been overlooked, but I believe that I have records for
all of the conspicuous elements of the lower vertebrate fauna in this
region.
In collecting animals from this community, I used a dredge which
was built specifically for the purpose, and which has been recently
described (Goin, 1942: 183). This dredge can be inserted under a
mat of hyacinths, then raised and carried ashore, where the mass of
plants is taken apart and the animals removed.
Many friends have at various times assisted me with the collect-
ing, but I wish in particular to thank Mr. J. C. Dickinson for his help
on many occasions. I also wish to thank Professor M. D. Cody for
information regarding the water hyacinths, and Dr. A. F. Carr for
his continued aid and assistance and especially for verifying the
identifications of the fishes. Finally I wish to thank Professor J.
Speed Rogers for his aid and advice throughout the study and in the
preparation of this manuscript.
Except where proposed changes have been published, the names
of the fishes in the following list are as given by Carr in his key to the
fresh-water fishes of Florida (1937), and the nomenclature of the
amphibians and reptiles follows Stejneger and Barbour's recent check
list of North American amphibians and reptiles (1943).
LOWER VERTEBRATE FAUNA OF THE WATER HYACINTH 145
FISHES
Lepisosteus platyrhincus De Kay
FLORIDA GAR
This form is common in the hyacinths around the edge of Newman's Lake.
I have records for April, May, June, September, October, and Novmber.
Amia calva Linnaeus
BowriN
The few spring records I have for this species (locally called the mudfish)
do not give a true picture of its abundance. I have seen four or five grown in-
dividuals on ithe edge of Payne's Prairie in a single evening. Records for
February, April, May, and October.
Signalosa petenensis vanhyningi Weed
FLORmA THREAnUD SHAD
While I have never collected this form among hyacinths, Dr. A. F. Carr in-
forms me that in Newman's Lake the eggs, which are laid in open water, frequently
come in contact with the roots of the hyacinths and adhere to them.
Erimyzon sucetta sueetta (Lacepede)
EASTERN LAKE CHUB-SUCKER
Only once have I collected this species among the water hyacinths, but there
are records in the Biology Department collection for January, February, March,
April, October, and November.
Notemigonus crysoleucas boscii (Valenciennes)
SOUTHEASTERN GOLDEN SHINER
I have records for January, March, April, October, and November.
Notropis maculatus (Hay)
SPOTTED SHINER
My only records are for February, April, and October.
Ictalurus cats (Linnaeus)
WHITE CATTISH
Although rare in the smaller bodies of water, this catfish is often caught by
fishermen in many of the larger lakes and streams. The only locality in which
I have found it definitely among the hyacinths is where a small stream enters
Newman's Lake. Commercial fishermen quite often take it in the same region.
Records for February, March, and October.
Ameiurus nebulosus marmoratus (Holbrook)
MARBLED BROWN BULLHEAD
This fish is commonly found among the water hyacinths. Records for
February, March, April, May, October, and December.
Ameiurus natalis erebennus Jordan
SOUTHEASTERN YELLOW BULLHEAD
The yellow bullhead is not so common as the preceding. Records for February,
March, and November.
146 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
Schilbeodes gyrinus (Mitchill)
TADPOLE MADTOM
This form is abundant around the roots of the hyacinths. Records for
February, March, April, October, and December.
Esox niger LeSueur
CHAIN PICKEREL
The chain pickerel, or jackfish, is seldom abundant in this habitat as an adult,
but in the spring of the year the young are sometimes plentiful. During March,
1941, young individuals from 5 to 25 mm. in length were very numerous
among the hyacinths on the edge of Newman's Lake.
Chriopeops goodei (Jordan)
RED-FINNED KILLIISH
This fish is rarely found among hyacinths. My only records are for the
month of February.
Fundulus chrysotus (Ginther)
GOLDEN TOPMINNOW
Fundulus is fairly common. Records for January, February, March, April,
May, September, October, November, and December.
Jordanella floridae Goode and Bean
FLAGFISH
This is one of the most abundant fish in muddy, hyacinth-covered water.
It can be collected with ease even after the water has been stirred up and is exceed-
ingly murky. Records for January, February, March, April, October, November,
and December.
Heterandria formosa (Agassiz)
LEAST KILLIFISH
This species is very rare among the water hyacinths. I have one record for
March.
Gambusia affinis holbrookii (Girard)
EASTERN MOSQUITO-FISH
This fish is commonly found, but never in the great numbers in which some
of the other small fish occur. Records for January, February, March, April, May,
October, November, and December.
Hololepis barratti (Holbrook)
FLORIDA SWAMP DARTER
I have records of the darter for February, April, October, and November.
Pomoxis nigro-maculatus (LeSueur)
BLACK CRAPPIE
The black crappie, or speckled perch, is abundant in Newman's Lake but is
seldom taken except during the months of February and March, when the fish
enter the hyacinth-clogged streams that empty into the lake. At this time as
many as thirty or forty may be caught on a hook and linein a single afternoon,
when topfish (Fundulus and Gambusia) are used as bait. During these spring
LOWER VERTEBRATE FAUNA OF THE WATER HYACINTH 147
months they may also be caught all around the shore, but never in as great
numbers as in the mouths of the small creeks.
Huro salmoides (Lac6pide)
LARGEMOUTH BASS
Although I have never collected this bass among the hyacinths with my dredge,
it certainly occurs there. Fishermen often catch it among the hyacinths both with
hook and line and with artificial bait.
Enneacanthus glorious (Holbrook)
BLUE-SPOTTED SUNFISH
I do not believe that my few spring records give a true picture of the abund-
ance of this species. Records for January, February, and March.
Chaeniobryttus coronarius (Bartram)
WARMOUTH BASS
This form, locally called chub, is very abundant in the shallow waters near
the edges of lakes and streams. A voracious feeder, it can be taken by any
moving bait. I have seen as many as eighty individuals caught by one man in
a single afternoon while fishing with a hook and line on Payne's Prairie, a large,
hyacinth-covered marsh about four miles south of Gainesville. I have records
for January, February, March, April, October, November, and December.
Lepomis macrochirus purpurescens Cope
EASTERN BLUEGILL
This locally important panfish is often caught among the hyacinths. Records
for January, March, April, October, and November.
Lepomis marginatus (Holbrook)
FLORIDA LONG-EARED SUNFISH
I have records for February and March.
Elassoma evergladei Jordan
EVERGLADES PIGMY SUNFISH
The pigmy sunfish is not common. Records for January and February.
Labidesthes sicculus vanhyningi Bean and Reid
SOUTHEASTERN BROOK SILVERSIDES
In central Florida Labidesthes is characteristically found in clear bodies of
water. I have two records for its occurence among the hyacinths, once in Feb-
ruary and once in March.
AMPHIBIANS
Amphiuma means means Garden
TWO-TOED AMPHIUMA
I have never found this form common in the water hyacinth community.
I have records for January, February, March, October, November, and December.
148 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
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Triturus perstriatus Bishop
FLORIDA STRIPED NEWT
This newt is relatively rare in the Gainesville region. The only time I have
taken it definitely in the water hyacinths was on February 18, 1937, when I
collected four adult specimens in a small, hyacinth-covered, woods pond about
three miles east of Gainesville.
Triturus viridescens loidsianensis (Wolterstorff)
LOUISIANA NEWT
The Louisiana newt is abundant in shallow water among the roots of the
water hyacinths during the winter months. A gravid female was collected and
placed in an aquarium on February 8, 1940. She deposited three eggs on the
bottom that night, and five additional eggs on February 12. The first three
eggs were stuck together in a string, but the five eggs laid on the twelfth were
deposited singly. An adult male collected on the same date regurgitated an adult
Gambusia. Records for January, February, March, and October.
Manculus quadridigitatus (Holbrook)
DWARF SALAMANDER
Van Hyning (1933: 3) lists this form as common "among leaves in dried-up
ponds in woods, and among roots of the water hyacinths (Piaropus crassipes)."
It is often found in small woods ponds that are covered with hyacinths, but I
have seldom taken it among the hyacinths in large bodies of water. Records
for January, February, March, October, and November.
Pseudobranchus striatus axanthus Netting and Goin
NARROW-STRIPED SIREN
No other aquatic vertebrate in Florida seems to be so restricted to the water
hyacinth habitat as is P. s. axanthus. I have collected extensively in Alachua
County for this species, and while it is usually abundant in all of the hyacinth-
covered lakes, prairies, and ditches that I have worked, I have never had any
success seeking for it in other situations. Records for every month of the year
except June, July, and August.
Siren lacertina Linnaeus
GREAT SIREN
This species is quite common in and among the roots of the water hyacinths
in the Gainesville region. I have seen several dozen in a single evening while
wading near the shore in Payne's Prairie. Records for February, March, April,
October, and December.
Acris gryllus (Le Conte)
CRICKET FROG
Acris gryllus is extremely common at all seasons of the year around the edges
of bodies of water covered with Piaropus. I once saw a large water spider
(Dolomedes) catch and kill a full grown cricket frog. Records for January,
February, March, April, May, October, November, and December.
Hyla cinerea cinerea (Schneider)
GREEN TREE FROG
The green tree frog has been found climbing about on the emergent por-
tions of water hyacinths on several occasions. Concerning this species, Kilby
LOWER VERTEBRATE FAUNA OF THE WATER HYACINTH 149
(MS 1936) says, "By far the greatest concentration of individuals is found
among lake or pond shore vegetation and on the floating rafts of water hyacinths
that are common in Florida Lakes." My own records are for January, February,
and October.
Hyla crucifer bartramiana Harper
SOUTHERN SPRING PEEPER
The six specimens from Alachua County, Florida, listed by Harper (1939: 2)
in the original description of this form, were taken in a small woods pond which
is nearly completely covered with hyacinths. The greatest number of individuals
I have seen at any one time in northern Florida was in the above mentioned pond,
where I collected eighteen on the evening of December 31, 1940. In the
Gainesville region the peepers usually enter the ponds and marshes in late De-
cember, and continue to call spasmodically until early spring. My only col-
lection records are for February and December.
Rana catesbeiana Shaw
BULLFROG
In the Gainesville region adults of this species are seldom found in the
hyacinth-covered ponds and streams, but immatures are quite common in this
habitat in the spring. Records for January, February, March, and December.
Rana clamitans Latreille
BRONZE FROG
The bronze frog is very seldom found among the hyacinths. In Florida this
form seems to prefer springs and seepage areas in heavily forested areas to any
other habitat. My only records of its occurrence with the water hyacinths are for
January and December.
Rana pipiens sphenocephala Cope
SOUTHERN LEOPARD-FROG
This frog is quite common near the edges of the larger bodies of water.
Records for January, February, October, November, and December.
Microhyle carolinensis (Holbrook)
NARROW-MIAOUTHED TOAD
I have never taken this toad among the hyacinths proper, but it can often be
found under mats of dead hyacinths that have been thrown up on shore. Records
for April, October, and November.
REPTILES
Alligator mississippiensis (Daudin)
ALLIGATOR
Alligators are, unfortunately, becoming quite rare in many parts of the state;
but about fifteen or twenty years ago they were regularly seen in Newman's
Lake, a large lake that has long supported a dense growth of hyacinths. It was
then not at all unusual to be able at a single time to count six or eight large
"gators" floating at the surface of the water.
150 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
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Sternotherus odoratus (Latreille)
STINK-JIM
The few records of this and the following form do not give a true picture of
their relative abundance in the water hyacinth community. They occur com-
monly in many aquatic situations, particularly those that are choked with trash
or vegetation. Records for February and October.
Kinosternon baurii baurii Garman
STRIPED MUSK-TURTLE
Records for January, February, October, and December.
Kinosternon subrubrum steindachneri Siebenrock
FLORIDA MUSK-TURTLE
The Florida musk-turtle can often be seen sunning itself in hyacinth-filled
road ditches, canals, and ponds. Records for January, March, May, and October.
Pseudemys nelsoni Carr
NELSON'S TERRAPIN
This "cooter" is quite common on Payne's Prairie, a large, hyacinth-covered
marsh about four miles south of Gainesville. Records for January, February,
April, May, August, October, and December.
Deirochelys qeticularia (Latreille)
CHICKEN-TURTLE
Unquestionably this is one of the most ubiquitous of all Florida turtles,
occurring in large clear lakes, rivers, marshes, prairies, ponds, and ditches.
The only records I have in which it was definitely among the water hyacinths
are for March and December.
Farancia abacura abacura (Holbrook)
HORN-SNAKE
The horn-snake is abundant in densely vegetated muddy marshes; it is very
common on Payne's Prairie. Records for January, February, March, April,
September, October, and December.
Diadophis punctatus punctatus (Linnaeus)
SOUTHERN RING-NECKED SNAKE
I have never collected this snake among the water hyacinths, but Carr
(1940: 79) states "I have found several in the water among water-hyacinth
roots with Farancia and Pseudobranchus."
Lampropeltis getulus floridanus Blanchard
FLORIDA KING-SNAKE
The Florida king-snake is wide spread but not particularly abundant.
Records for February, March, April, May, September, and October.
LOWER VERTEBRATE FAUNA OF THE WATER HYACINTH 151
Natrix sipedon pictiventris Cope
FLORIDA BANDED WATER-SNAKE
This snake is extremely common and wide spread. Records for March,
April, May, October, November, and December.
Seninatrix pygaea (Cope)
RED-BELLIED MUD-SNAKE
With the possible exceptions of Pseudobranchus s. axanthus and Liodytes
alleni, this snake seems to be more closely correlated with water hyacinths than
any vertebrate known to me. I have records for February, March, May,
October, November, and December.
On the afternoon of October 18, 1939, I collected a large female which that
night gave birth to eight young. The following table gives the measurements
and scale counts of this series of young. The catalogue numbers are those of
the Carnegie Museum.
CM No. Sex Scale Rows Abdominals Sub-caudals Total length Tail length
18753 d 17-17-15 121 54 129 31
18754 d 17-17-15 123 54 127 29
18755 d 17-17-15 126 53 134 30
18756 ? 17-17-15 127 44 123 24
18757 9 17-17-15 125 45 124 25
18758 d 17-17-15 126 55 128 30
18759 9 17-17-15 122 42 124 24
18760 ? 17-17-15 125 45 124 25
The mother (CM18752) had a maximum of 17 scale rows, 122 abdominals,
43 sub-caudals, and a total length of 379 mm. and tail length of 67 mm.
Liodytes alleni (Garman)
ALLEN'S MUD-SNAKE
As stated by Carr (1940: 92), this is probably the most aquatic snake in
Florida, and is fairly common among the water hyacinth roots in shallow water.
Records for February, October, and December.
Agkistrodon piscivorus piscivorus (Lac6pide)
COTTON-1OUTH MOCCASIN
I have never found this snake common among the hyacinths. It seems to me
to be more commonly found along the borders of lakes and streams in heavily
wooded regions. I have records of its occurrence among the hyacinths for
'February, April, May, and December.
152 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
LITERATURE CITED
CARR, ARCHIE F., JR.
1937. A key to the fresh-water fishes of Florida. Proc. Fla. Acad. Sci. for
1936, 1: 72-86, 1 fig.
1940. A contribution to the herpetology of Florida. Univ. Fla. Biol. Ser.,
3 (1): 1-118.
GoLr, COLEMAN J.
1942. A method for collecting the vertebrates associated with water hya-
cinths. Cofkia, 1942, No. 3: 183-184, 1 fig.
HARPER, FRANCIS.
1939. A southern subspecies of the spring peeper (Hyla crucifera). Notulae
Naturae, No. 27: 1-4.
KILBY, Jon~ D.
1936. A biological analysis of the food and feeding habits of Rana spheno-
cephala (Cope) and Hyla cinerea (Schneider). Unpublished Master's
thesis, University of Florida.
STEJNEGER, LEONHARD, and THOMAS BARBOUR.
1943. A check list of North American amphibians and reptiles. 5th ed.,
xix 4 260 p. Bull. Mus. Comp. Zool., 93, No. 1.
VAN HYNING, OTHER C.
1933. Batrachia and Reptilia of Alachua County, Florida. Copeia, 1933,
No. 1: 3-7.
LOWER VERTEBRATE FAUNA OF THE WATER HYACINTH 153
PLATE I
Water hyacinths (Piaropus crassipes)
Upper.-A mass of water hyacinths completely covering the mouth of a creek
entering the St. Johns River.
Lower.-Two individual plants in an aquarium showing the submerged root
system.
-Photographs by Albert M. Laessle.
A CONTRIBUTION TO THE NATURAL HISTORY
OF THE FLORIDA SHORT-TAILED SHREW
JOSEPH C. MOORE
Department of Biology, University of Florida
The Florida short-tailed shrew, Cryptotis floridana Merriam, is one
of the smallest animals native to Florida. It belongs to the order of
insectivores and comes of a family famous for diminutive bodies, ra-
pacious appetites, secretive habits, unresting activity, irritable nature
and none-too-pleasant odor. Other members of this family, the
Soricidae, known to occur in the state are the Carolina short-tailed
shrew (Blarina brevicauda carolinensis), the Everglade short-tailed
shrew (Blarina brevicauda peninsulae, and the Bachman shrew (Sorex
longirostris longirostris).
The Florida short-tailed shrew is a mammal about the size of a
lady's little finger, salt-and-pepper gray above and slightly lighter be-
neath. Its feet appear proportionately small; its rather short tail is
not remarkably hairy. It has a long pointed nose, very small eyes, and
no external ears. The lack of external ear undoubtedly suggested the
generic name Cryptotis or "hidden ear."
Before proceeding with the discussion on this animal, I wish to
express my gratitude to Professor H. B. Sherman whose help, guidance
and encouragement made this paper possible; to Professor J. Speed
Rogers who arranged for me to live on the University of Florida Con-
servation Reserve at Welaka where this study was carried on, and to
the many others who helped, encouraged, or simply endured.
The known distribution records of the Florida short-tailed shrew
were published by Professor Sherman (1937: 105) in his check list
of the land mammals of Florida. More recent records have not ap-
preciably extended the known range.
Recorded information concerning the natural history of this shrew
consists almost wholly of casual references to the types of habitat in
which the species has been taken. Chapman (1894) tells of finding
the remains of one whole individual and the skull of another in the
stomach of a barn owl taken in Gainesville. Both Elliot (1901: 55)
and Smith (1938: 372) found this animal inhabiting palmetto ham-
mocks in the vicinity of Sanford. Merriam (1895: 19), acquainted
with it only through four specimens, laconically gives the habitat as
"palmetto scrub." Bangs, who took many specimens and was of the
opinion that this shrew occurs in all sorts of situations, wrote (1898:
209): "I have taken it on barren sand hills, in old fields, in hummocks,
swamps, piny woods, under brush piles, and at the edge of the salt
155
156 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES vol. 6,
Nos. 3-4, 1943
marsh." He considered its favorite abode to be among the rushes that
fringe the borders of small fresh-water ponds, especially near the edge
of the salt marsh. Bangs also notes that these shrews like others
"have a great fondness for eating each other in the trap; usually they
begin with the head." Blair (1935: 274) was the first to publish
critical data on the relative numbers of individuals trapped in dif-
ferent types of habitats.
HABITAT PREFERENCES: The results of Blair's trapping of
Cryptotis and Blarina on the Biology Station grounds at Newman's
Lake, near Gainesville, appear to be in agreement with my own results
obtained on the University of Florida Conservation Reserve at Welaka,
Putnam County, Florida. Cryptotis appears to have a greater range
of habitat tolerance, having been taken in most of the moist, shady
situations where Blarina was trapped, and also in relatively dry, open
places where Blarina apparently did not occur. Table No. 1 presents
the figures upon which this statement is based.
TABLE No. 1-HABITAT PREFERENCES OF CRYPTOTIS AND BLARINA
1. Number of shrews trapped by W. F. Blair near Gainesville, Florida.'
Dry Pineland Briar Patch Grassy Swamp Border
Cryptotis ........................ 4 3 3
Blarina ............... ........ 0 2 4
2. Number of shrews trapped by J. C. Moore and J. J. Friauf at Welaka, Florida.
Flatwoods Pond Border Bayhead
Cryptotis ........................ 5 1 0
B larina ............................ 0 1 3
1Data from Blair, op. cit., p. 274.
SYNONYMY: Characters distinguishing the Florida short-tailed
shrew, Cryptotis floridana, from its nearest relative to the north,
Cryptotis parva, were first noted by Professor Baird in 1857; he
nevertheless did not describe the Florida form, but referred it to
Blarina cinerea (a synonym of Cryptotis parva). In 1894 Chapman
listed it under two different names, both synonyms of C. parva. The
following year Merriam revised the genus Blarina, including Cryptotis
as a subgenus; he described the Florida form as Blarina floridana, and
indicated its differences from the related parva. Cryptotis was given
generic status by Miller in his 1912 check list. Harper in 1927
placed floridana as a race of parva, but it was given specific rank by
Blair (1935) and. Sherman (1936). This nomenclatorial history is
covered by the following list of citations:
THE FLORIDA SHORT-TAILED SHREW
Cryptotis floridana (Merriam)
1857. Blarina cinerea Baird, Mammals of North America.
1894. Blarina cinerea and Blarina exilipes Chapman, op. cit.
1895. Blarina (Cryptotis) floridana Merriam, op. cit., p. 19.
1901. Blarina floridana Elliott, op. cit., p. 55.
1912. Cryptotis floridana Miller, op. cit., p. 25.
1927. Cryptotis parva floridana Harper, op. cit., p. 270.
1935. Cryptotis floridana Blair, op. cit., p. 274.
1937. Cryptotis floridana Sherman, op. cit., p. 105.
METHODS OF CAPTURE, AND RECORDS: On the Univer-
sity of Florida Conservation Reserve, 3,708 trap-nights with live traps
caught only three specimens of Cryptotis. Mr. James J. Friauf, a
fellow-student who was working on the ecology of the Orthoptera of
the Reserve, took two Cryptotis in from 300 to 500 trap-nights with
"molasses traps" set for crickets and cockroaches. This suggests that
some kind of an open pit trap, properly baited, may be effectively
used for capturing this tiny animal. Dr. T. H. Hubbell informs me
that he has had similar success in capturing shrews with molasses
traps in Michigan, Tennessee and other regions; it is a question
whether the shrews simply blunder into the pits, or are attracted
thereto by the entrapped insects. One Cryptotis was found dead and in
good condition beside the hard road which runs through the Reserve.
In the following table "No." refers to the catalog number of the
specimen in the writer's collection, "Date" to the date of capture,
"Skin" to stuffed skins preserved, "Skl." to body skeletons preserved,
"Alc." to whole specimens preserved in alcohol. Measurements are re-
corded in this order: total length, tail-length, length of hind foot, and
weight. All linear measurements are in millimeters, and weight is
given in grams.
RECORD OF SPECIMENS OF CRYPTOTIS TAKEN ON THE RESERVE
No. Date Skin Ski. Ale. Sex Measurements
M-32 12-13-39 x F not taken
M-40 2-25-40 x x M 79x23x11x5.0
M-48 3-15-40 x F not taken
M-50 3-31-40 x x F 74x24x11x5.0
M-58 5-1-40 x ? 78x22x11x3.6
M-107 10-11-40 x F not taken
The following notes are made from observations of one individual,
a female, which was kept in captivity for more than three months,
Oct. 11, 1940 to Jan. 28, 1941. The name "Shivernose" was given
to this animal by a small boy, who thought it was cold because, as he
said, "It shivered its nose all the time."
158 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
NESTS: The shrew was kept in a glass vivarium with the floor
covered with sand. It was moved to a clean vivarium every ten or
eleven days, and each time this happened it was obliged to make a
new den or nest of whatever material was made available. Scrub oak
leaves scooped off the ground in the scrubby flatwoods where the
shrew had been trapped provided cover in which it hid itself but made
no real nest. During cold weather it was given tufts of absorbent
cotton which it readily made into a nest. These cotton nests were
never so neat and globular as those made under similar circumstances
by harvest mice (Reithrodontomys humulis), cotton mice (Peromys-
cus g. gossypinus), and golden mice (Peromyscus nuttalli aureolus).
On various occasions Shivernose made nests of sphagnum moss or
dried grass. At one time the shrew had only the shelter of a flimsy
camouflage of bits of dry crab grass which it had dragged together in
one corner of its cage. It added to this skimpy cover whenever more
material was made available, but abandoned it when a more suitable
den was offered in the form of an egg shell. This dried shell of an
ordinary hen's egg, into the side of which a hole 18 mm. in diameter
had been broken to remove the contents, was actually large enough
to have held four shrews the size of Shivernose. It had hardly been
placed in the vivarium when the shrew entered it and adopted it as a
rather permanent den. A bit of cotton was placed in it for nesting
material, but the shrew dragged in more and also pieces of dry grass
until there was barely enough room for the shrew.
Smith (1938: 372-373) describes a nest of this species found under
a log in the vicinity of Sanford:
"The nest was loosely constructed of leaves of panic grass (Panicum
mutabile) and was about 5 inches in diameter. There was no evident at-
tempt at weaving or any other means of fastening the leaves in place;
they were loosely piled in a globular mass. Small pieces of shed snake
skin, seemingly from a Florida king snake (Lampropeltis getulus floridanus),
were mixed with the leaves, and were also scattered about the sides of the
log away from the nest."
BURROWS AND TUNNELS: Every time that the dried-out sand
in Shivernose's vivarium was replaced with fresh, moist sand, the
shrew responded by riddling it with tunnels during the night. On two
occasions fresh sphagnum moss was spread over the floor of the cage.
The sphagnum screened Shivernose's activities too well, however, and
in consequence the moss was removed each time after several days.
It was noted that whenever the shrew burrowed in the moist sand or
sphagnum after having been denied the privilege for several days, its
fur become fluffy and more healthy-looking.
THE FLORIDA SHORT-TAILED SHREW
On one occasion two ridges about five inches high were moulded
by hand in the moist sand, so as to divide the vivarium into three
parts. Placed in one of the compartments formed by these ridges, the
shrew ran about until it found places where it could climb over the
sand walls, and for some time scrambled back and forth from one
compartment to another. Presently, I poked a hole with my finger
through one wall at the bottom. When the shrew had used this hole
for a tunnel several times, it proceeded to dig a tunnel through the
other wall. The diameter of the tunnel that it dug was, 13 mm.
vertically and 18 mm. horizontally.
One time when the sand of the vivarium had become too dry and
loose for burrowing, and Shivernose had become accustomed to denning
in a particular corner of the arena, which was only thinly protected
by a few stems of dried grass, dampening the sand stimulated the
shrew to spend much of the night digging tunnels and burrows; al-
though it still spent the days in its grass-covered lair instead of under-
ground. Here the apparently conditioned habit of denning in the
corner above ground appeared temporarily stronger than the more
common habit of denning in a burrow.
On one occasion when the shrew was placed in a vivarium with
fresh sand, water was poured on the sand at one end of the cage with
the intention of making it more suitable for burrowing. This must
i -- mm. ------- I
Fig. 1.-"Shivernose" (Cryptotis floridana) asleep in its burrow. The shrew's
forehead is practically flat on the ground in this sleeping position, and it is
lying more on its belly and the top of its head than on its side.
160 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
have made the sand too wet, however, for after the shrew had been
left in this for a while, it was found asleep in a burrow in the relatively
dry end. The whole burrow was against the glass end of the vivarium,
so that one saw it practically in longitudinal section. The mouth of
the tunnel was nearly closed, through accident or design, and the shrew
apparently felt unusually safe, for this was the only time that it was
found curled up as shown in figure 1.
SANITATION: At first Shivernose defecated only in one corner
of the vivarium, but after a while any corner except that of the
den was used. Faeces and urine were extruded simultaneously, the
faeces usually in one lump. It took but an instant and was accom-
plished with a contortion which looked like peristalsis of the whole
body. In the latter days of imprisonment, the shrew began the prac-
tice of defecating against the glass of its cage, and the faeces stuck
to it about 4 to 8 mm. above the surface upon which the shrew stood.
This behavior has some similarity to that reported by Carl O. Mohr
(1940: 93) of a house mouse (Mus musculus Linn.) which, when
kept in a tall glass cylinder, took its faeces and stuck them up on the
walls of its cage as high as it could. Mohr's remark that "this be-
havior seemed to be an act of sanitation" might well apply in the case
of the shrew, whose behavior at least eliminated the hazard of
walking in its own excrement.
The frequency with which such a voracious creature defecates may
be of some interest. Once when Shivernose had been in a clean cage
nine hours, evidence indicated that it had defecated four times in that
period. On another occasion the shrew was observed to defecate twice
within 23 minutes.
MOULTING: A line of molt was noticed around the shrew's
thorax December 14 to 19.
SENSES: It seemed apparent at times that Shivernose had little
use of its eyes. On many occasions it,ran within two inches of a large
insect without becoming aware of it. Observing the insectivore hunt its
food in the vivarium, occasions the belief that it depends a great deal
on its ears and even more on its nose in locating its prey. Shivernose
often appeared to detect an insect's presence by smell and, while ap-
proaching it, its exact location as well. But the shrew appeared seldom
able to determine the exact location of an insect further away than an
inch or two. By hearing a large insect walk, the shrew apparently
could locate it accurately at a distance of three inches or more, and
would close upon it from that distance with confidence. On one oc-
cision a cornered cricket, which Shivernose approached cautiously,
attempted to rush past about three-quarters of an inch from the shrew's
THE FLORIDA SHORT-TAILED SHREW
head. Instantly aware of this the shrew flirted its head to the side
and caught the cricket in full career. The movement was almost in-
credibly swift.
ALERTNESS: Shivernose was never observed to be relaxed and
still. Regardless of where it was, the shrew raised its head and
"shivered its nose" to test the air, turned its head this way and that,
and whirled about quickly as if to make sure it was not about to be
attacked from the rear. However, these moves were made so swiftly
and punctually that they gave an impression more of wariness than of
timorousness. When the situation was such that the shrew was
obliged to eat in the open, it ran from its food every now and then
to a corner and crouched there sniffing the air for a few moments.
Then it returned to eat some more for a few seconds before repeating
this performance. Another caution behaviorism observed was the
tendency, when suspicious or alarmed, to run along very close to the
walls of the vivarium.
Even on an occasion when the shrew was observed while sleeping
in a burrow, it appeared to be sleeping "on the alert." Every now and
then it jerked up its head, sniffed the air this way and that, and then
curled up again; and sometimes it did an about-face. Also, strange,
periodical fits of trembling seized Shivernose as if the little beast
were having "bad dreams."
HUNTING: The hunting tactics displayed by Shivernose in at-
tacking a grasshopper in its vivarium consisted of creeping up to with-
in an inch or so of it, and reaching out and snatching it back with its
teeth. Then, if the shrew were under the cover of its nest or a tunnel
in the sphagnum, it usually only turned about with its prey and pro-
ceeded to kill it by biting it in the head. If, on the other hand, it were
out in the open, it rushed its victim back to a burrow to kill it there.
And, when no shelter at all was at hand, it carried its prey habitually
to a certain corner or part of the arena each time.
The shrew's method of attack on crickets differed sharply from
that used on grasshoppers. When a cricket was definitely located,
instead of creeping up and snatching it, Shivernose charged and closed
upon it like a football player upon a fumbled ball. If the cricket fled
an instant before the shrew rushed, an exciting but short race ensued.
The only place for the shrew to get a good hold on a cricket appeared
to be the head, and sometimes it chased big crickets several lengths
of the vivarium before getting the right hold. And when it did capture
such a one, the shrew snatched it up and scurried back with it holding
it high and seemingly bursting with excitement and pride, the hair
on the back of its neck standing up high.
162 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6,
Nos. 3-4, 1943
The only animal besides the cricket with which the shrew was ob-
served to throw itself into bodily contact while fighting it was a pray-
ing mantis. The shrew's fighting in this case was all done in a brief
whirl of wrestling and biting from which it emerged chewing the van-
quished mantis's head. In all other cases its fighting consisted of dash-
ing in to nip and jumping back.
Once a small (12.3-gram) soft-shelled turtle, Amyda ferox, was
put into the vivarium with the shrew. Shivernose attacked it at once,
nipping it sharply and jumping back again and again. Sometimes
the shrew stood on its hind feet shooting out a nip or recoiling after one
like a light-weight boxer. After every few such nips, Shivernose ran
off to a far corner, tested the air carefully for a few moments, and
presently returned to the fray. The turtle made a spirited attempt to
escape through the glass but did not offer to do battle. It was taken
out before it suffered any real damage.
FEEDING HABITS: When eating from a large insect or chunk
of meat, the shrew crouched over the food, holding it down with its
forefeet and pulling and tearing at it with its teeth in the manner of
a small lion. The food given to the captive shrew most of the time
was grasshoppers. These were usually Melanoplus femur-rubrum pro-
pinquus, Chortophaga australior, and another common species about
20 mm. long. Smaller and softer individuals were eaten entirely, as
were crickets and other soft-bodied arthropods. With larger grasshop-
pers the procedure was generally as follows. The shrew began by either
eating the grasshopper's head or by biting it loose from the thorax
and licking and scraping out the contents of the head through the
neck hole. Then, turning to the thorax, the Cryptotis tried to clean
it out through the neck hole. Having gleaned as much as possible that
way, it bit and tore the abdomen loose from the thorax and cleaned
out the remainder of the thorax from the posterior end. Next, it ate
the femora of the jumping legs in the manner of a boy with a lolly-
pop, chewing the fleshy part in one side of its mouth while the distal
end protruded as would the lolly-pop handle. Finally, the shrew
laboriously ingested the abdomen-starting at one end and chewing
until the whole was devoured. This last part was sometimes modi-
fied to ripping the abdomen open and stripping it clean, and sometimes
was omitted altogether.
On a big, heavily armored beetle, Passalus cornutus, the shrew's
attack was completely ineffective. When the elytra and pronotum
were removed, however, Shivernose readily ate the beetle.
Several times top minnows, (Fundulus sp.), varying in length be-
tween 20 mm. and 40 mm., were placed in a watch glass which held
THE FLORIDA SHORT-TAILED SHREW
a disc of water 9 mm. deep and 50 mm. in diameter. In attempting to
capture them the shrew stood with both front feet in the water and
moved its head alertly this way and that trying to locate a fish. Find-
ing one, the shrew would snap it up and rush down into the under-
ground runways. Sometimes the insectivore put its nose and mouth
under water and searched about, apparently trying to locate a
minnow by touch. It frequently had considerable difficulty in catching
the minnows, for they were very active and apparently difficult for
the shrew to perceive even in so shallow a disc of water. To eat a
minnow it always started at the head end.
For 48 hours at a stretch Shivernose was sometimes fed nothing
but milk. It once lived on cheese longer than that, and also, on an-
other occasion, on corned beef. After taking a bite or two of peanut
butter, the shrew often "scooted" its chin along in the sand in an
amusing manner and drank water every few seconds for a while. After
eating some of the fresh, sticky turtle's egg, the shrew wiped
the viscid stuff off its lips with its hands, and then was obliged to rake
its hands clean on its teeth. This awkward procedure was very
amusing to watch.
FOOD PREFERENCES: During Shivernose's first two months
in captivity the plentiful grasshoppers, as previously stated, provided
most of its meals, but when cold weather brought a scarcity of grass-
hoppers, the shrew was offered a rather heterogeneous assortment of
other foods, as listed in Table No. 2.
TABLE No. 2-FOOD PREFERENCES OF CRYPTOTIS FLORIDANA
Eaten ravenously:
Crickets
Butterfly
Frogs
Eaten readily:
Grasshoppers
Termites
Beetles (Passalus cornutus)
Minnows (Fundulus sp.)
Hamburger
Ovarian turtle's egg
Beef tenderloin
Soft-shelled turtle flesh
Pocket-gopher flesh
Cotton rat flesh
Flying squirrel flesh
Eaten unenthusiastically:
Sardines in brine
Raw fresh pork
Raw smoked ham
Roast pork
Corned beef
Milk
Cream
Hen's egg
Cheese
Peanut butter
Eaten with reluctance:
Fresh bream and bass heads
Opossum flesh
Beetle larvae (Passalus cornutus)
Walking stick
Rabbit liver
Refused:
Scorpion
Mantis
Smoke-cured breakfast bacon
164 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES Vol. 6.
Nos. 3-4, 1943
The captive Cryptotis showed a definite preference for crickets
over grasshoppers. When several of both were dropped into its cage,
the shrew ran about among them biting the grasshoppers in the head
to kill or cripple them. But usually when it caught a cricket the in-
sectivore stopped to eat it. If a crippled grasshopper stirred more
than suited the shrew, it stopped eating the cricket to bite the grass-
hopper savagely in the head, and then returned to finish eating the
cricket. Only after the crickets were all gone did the shrew start eating
grasshoppers.
Shivernose apparently fasted twenty-four hours rather than eat
smoke-cured breakfast bacon. That is an exceedingly long fast for
a shrew. In regard to cheese, which several times supported Shivernose
for as long as 48 hours, it should be pointed out that with cheese
present in its cage, another shrew of this species died on the second day
of captivity, apparently of starvation.
HOARDING FOOD: Shivernose dug a burrow at one end of the
vivarium so that the chamber was against the glass. I could see the
shrew in this chamber when I began dropping grasshoppers and
crickets into its glass cage two or three at a time. The first several
were crickets. Shivernose came forth and captured and ate the first
three in three minutes, and carried the fourth down into the den. After
remaining below with it for a minute or so, the shrew emerged and
began to carry insect after insect down, until I suspected it of not
eating them and peered into the den through its accidental glass window.
Sure enough, the grasshoppers and crickets were piled three deep.
Each time Shivernose brought an insect in, shoved it over the top of the
others, bit its head vigorously, and dropped it. Then, for a minute
or so the shrew crouched there, darting an additional nip at the head
of whichever victim moved unduly. Presently it went to one of the
two openings to the surface and, after testing the air with its nose,
emerged and darted swiftly about the arena until it located a victim.
Seizing it, the shrew sped back across the surface and down into its
burrow as fast as the eye could follow. Sometimes it carried a big
grasshopper in such a way that the narrow entrance of the burrow
knocked the insect from the shrew's jaws. This never slowed up the
shrew's entrance. It turned about down below, reached up again, and
jerked the grasshopper down. In thirty minutes Shivernose had eaten
four or five (at the most) and hidden away twenty grasshoppers and
crickets in this way.
When the rain of grasshoppers had ceased and the shrew had taken
a last scout around, it rested crouching in its crowded store-house for
about three minutes. Then it began to dig into the sand wall of one
THE FLORIDA SHORT-TAILED SHREW
side of the chamber, kicking the sand over the recumbent grass-
hoppers. In two minutes it had dug a new chamber and completely
covered up its cache. The shrew undoubtedly used some of this store
of food during the 12-14 hours before it was fed again.
DRINKING: During the hunting of the last five or six grass-
hoppers of the hoarding episode, the shrew stopped to drink water
four or five times. It always drank copiously after eating peanut
butter or brine sardines. Its method of drinking, quite different from
that of its relative, the mole, was to take a sip and raise its head to
let the water run down its throat, the way a chicken does. This was
done so quickly that the shrew must have taken two or even three
sips a second. Although it seldom drank more than three or four sips
at a time, it was once observed to drink eleven sips within a minute
or so, when it had been without water for some hours.
BREATHING: Shivernose's breathing rate at rest is about 170 times
per minute. This figure was arrived at by averaging eight counts made
when the shrew was napping. The room temperature was between 70
and 800 F. During the waves of trembling which passed over its body
every fifteen or twenty seconds, the shrew's breathing became extreme-
ly rapid for a moment. This made exact counting very difficult.
VOICE: The captive shrew often emitted tiny, bird-like chirpings.
Just what occasioned them is hard to say. When it was in its nest
under the sphagnum, I tried for a while to teach it to come up for food
when called in a certain way. This experiment was not successful;
but when I called for it to come out and tapped on the sphagnum
above its nest, it nearly always responded with that feeble, cheerful
chirping. Later on, when Shivernose was sitting undisturbed in the
egg-shell nest, we often heard the small beast twittering in that high-
pitched, tiny voice.
166 PROCEEDINGS FLORIDA ACADEMY OF SCIENCES ol. 6,
Nos. 3-4, 1943
LITERATURE CITED
BAIRD, SPENCER F.
1857. Mammals of North America.
BANGS, OUTRAM.
1898. The land mammals of peninsular Florida and the coast region of
Georgia. Proc. Boston Soc. Nat. Hist., 28: 209.
BLAIR, W. FRANK.
1935. The mammals of a Florida hammock. Journal of Mammalogy, 16
(4): 274.
CHAPMAN, FRANK M.
1894. Remarks on certain land mammals from Florida, with a list of the
species known to occur in the state. Bull. Amer. Mus. Nat. Hist., 6:
333-346.
ELLIOT, D. G.
1901. A list of mammals obtained by Thaddeus Surber, in North and South
Carolina, Georgia, and Florida. Field Columbian Museum, Publ. 58:
Zool. Ser., 3 (4): 35.
HARPER, FRANCIS.
1927. The mammals of the Okefinokee Swamp region of Georgia. Proc.
Boston Soc. Nat. Hist., 38 (7): 270.
MERRIAM, C. HART.
1895. Revision of the shrews of the American genera Blarina and Notio-
sorex. North American Fauna, 10: 19.
MILLER, GERRIT S.
1912. List of North American land mammals in the U. S. National Museum.
Bull. U. S. National Museum, 79: 270.
MOHR, CARL O.
1940. Strange action of a house mouse. Journal of Mammalogy, 21 (1):
93.
SHERMAN, HARLEY B.
1937. List of the recent wild land mammals of Florida. Proc. Florida Acad.
Sci. for 1936, 1: 102-128.
SMITH, CLARENCE F.
1938. Nest of Cryptotis floridana. Journal of Mammalogy, 19 (3): 372.
INDEX TO VOLUME 6
[New names are printed in bold-face type]
ALGAE1
algal flora of some Florida soils, 59-65
Allen, E. Ross (see Hubbs, Carl L.)
Ambystoma tigrinum, neoteny in, 37-40
AMPHIBIA (see also Salamanders)
survey of in Florida, 67-68
of water hyacinth community, 143-152.
Bahaman fresh-water turtles, (2.)7--.3
Biological effects of neutron radiation, 2.5-
l8
Bless, Arthur A., Biological effects of neutron
radiation, 2.5-1.8
BRYOPHYTES -
Check list of Florida mosses, (21)-16
Campbell, Robert B., Earthquakes in Florida,
'-4
Carr, A. F., Jr., and Coleman J. Goin, Ne-
oteny in Florida salamanders, 37-40
CHEMISTRY
separation of terpenes, (2)17-32.
Coleosporium species, 131-142.
CORALS, 41-48
key to Madreporaria in Miami area, 43-
45
CRUSTACEA
new species of crayfish, 49-58
Procambarus leonensis, 49
-pycnogonopodus, 53
DeVall, Wilbur B., The correlation of soil pH
with distribution of woody plants in the
Gainesville area, 9-1.4
Earthquakes in Florida, 1-4
ECOLOGY
lower vertebrate fauna of water hyacinth
community, 143-152.
ECONOMICS
theory of normal price, 2.9-36
Ellis, H. R. (see Smith, F. B.)
Emerson, A. E., and E. M. Miller, A key to
the termites of Florida, 1o8-1o9
FISH
of Silver Springs, 11o-130
of water hyacinth community, 143-152
FLORIDA
algal flora of soils, 59-65
amphibia, 67-68
earthquakes in, 1-4
fishes of Silver Springs, 110-130
littoral fauna of Miami area, 41-48
lower vertebrate fauna, 143-152.
mosses of, (2.)1-16
needle rusts of pines, 131-142.
salamanders in 37-40
termites, distribution and habits of,
101-107
termites, key, 108-109
vegetation types in Gainesville area, 9-
2.4
water hyacinth community, 143-15s
Florida short-tailed shrew, 155-166
Fuguitt, Robert E. (see Stallcup, W. David)
Gainesville area, vegetation types, 9-.4
Goin, Coleman J., The lower vertebrate fauna
of the water hyacinth community in North-
ern Florida, 143-152. (see also Carr,
A. F., Jr., and Netting, M. Graham)
Hawkins, J. Erskine (see Stallcup, W.
David)
Hobbs, Horton H., Jr., Two new crayfishes
from the Panhandle of Florida, 49-58,
Hodsdon, L. A., and Jay F. W. Pearson,
Notes on the discovery and biology of two Ba-
haman fresh-water turtles of the genus Pseu-
demys, (2)7-2.3
Hubbs, Carl L., and E. Ross Allen, Fishes
of Silver Springs, Florida, 110-130
INSECTS (see Termites)
Kalotermes (Calcaritermes) nearcticus, 5-8
Littoral fauna of Miami area, 41-48
Madreporaria (see CORALS)
MAMMALS
Florida short-tailed shrew, I55-166
Marine aquaria, (2.)24-16
Miller, Dorothy B. (see Miller, E. Morton)
Miller, E. Morton, The soldier and nymphal
forms of Kalotermes (Calcaritermes) ne-
arcticus Snyder, 5-8
Miller, E. Morton, and Dorothy B. Miller,
A preliminary study of the distribution and
habits of South Florida termites, o01-107
Miller, E. M. (see Emerson, A. E.)
Moore, Joseph C., A contribution to the na-
tural history of the Florida short-tailed
shrew, I55-166
Mosses, check list, (2z)-16
Neoteny, in salamanders, 37-40
Netting, M. Graham, and ColemanJ. Goin,
Progress report on a survey of the amphibia of
Florida, 67-68
JOURNAL OF FLORIDA ACADEMY OF SCIENCES
Neutron radiation, biological effects, 25-z8
Pearson, Jay F. W. (see Hodsdon, L. A.)
Pines, needle rusts, 131-141
PLANT DISEASES
needle rusts of pines, 131-141
Procambarus leonensis, 49
-pycnogonopodus, 53
REPTILES
Bahaman turtles, (1)17-13
of water hyacinth community, 143-152
Salamanders
neoteny in, 37-40
Florida species, 67-68
species to be expected in Florida, 67
Salinity, control in marine aquaria (2)
14-216
Schornherst, Ruth Olive, Check list of Florida
mosses, (2)1-16
Sherman, John H., An attempt at complete
statement of the theory of normal price, 2.9-
36
Shrew, natural history of, 155-166
Smith, F. B., and H. R. Ellis, Preliminary
report on the algal flora of some Florida soils
59-65
Smith, F. G. Walton, Littoral fauna of Mi-
ami area, 41-48
The provision of controlled salinity
variations in experimental marine aquaria,
(1)14-16
SOILS
algal flora of in Florida, 59-65
pH in relation to vegetation, 9-14
Spiral screen fractionating columns, (1)17-
31
Stallcup, W. David, Robert E. Fuguitt, and
J. Erskine Hawkins. Spiral screen fraction-
ating columns for the separation of terpenes,
(1)17-31
Termites
distribution and habits, 101-107
Kalotermes (Calcaritermes) nearcticus, 5-8
key to Florida termites, 108-109
soldier and nymphal forms of, 5-8
Terpenes, separation of, ()2.7-32
Theory of normal price, 29-36
Turtles, Bahaman fresh-water, (1)17-13
Vegetation
types in relation to soil pH, 9-24
VERTEBRATES
lower vertebrate fauna of water hyacinth
community, 143-152
Water hyacinth community, 143-152.
Weber, George F., Needle rusts of pine trees in
Florida caused by Coleosporium species,
131-142.
168
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