hy."

he
ft

Ny take! A
‘ : o 4%. ek Re
A oY wr MOAR te tet Geta
a ‘ a ‘ 4 1 GR Rye
t v) PR let ww 4K nt
; \ : om UTA hot |
% Rta)
‘ ov 4
\ hi my oe
t4
+ a . AS Ya taes,
, on
‘ eit
he ty
Hh Sart sth |
he hn
forthy Actes Pa
Re tenth Ge
Hy es
i SBM thy ra Syl
' B49
5 y st sa
: 5 , ;
a A : ‘ ‘
: . ‘ " ;
, a ‘ aay
' ea
‘ ct 4 :
hy { AAS A E
4 i :
a lbs at
, 1 7
ait :
'
‘a ,
; e ; ' j
,
; , i Sat, Sea Ne 4 F} rye
fe)
: pias i Ae ee
hy i Pace ce Lies 4
eae jenn ! }
r ; ; é - Uetrd any nas i
3 M CUE Ay det) ‘ k
, j P ' ‘ i wd
ed at me. ’ s es
P ‘ #BEIy ws PFE bey
: bth hee
° a j
t 4
"iy 1 ; i
’ 5
’ nt
’ ; " ‘
, :
bo OE ee y
my , ey
‘ rik ae vay} ; }
) rad ; i hs aah Fae
: ; “psig
i hos 5 he i eehs
, , e ”
, i; ei , 7 '
i Pg ae PLR CT ea D Hty Sd 4 ‘ 3 ‘ +8, dpa! ; oy f in a2 9 5 BG ER MAT Te Ah AT Sa Wik ecep cis
’ : ; ; P é { ¢ ; 4 Wparpette ate Gt
f t 4 J d wee 4 Sa:
: : 4 are ,
‘ , i,
‘ ib Sats A Oi ge
’ rot , i ie 7 Neen
eo rae 5k % hse He prAna
; UNG ee ‘ 4 bey eas Seas
Lag (48 Mei
te ‘ 4 cas x ak Aig ha 'niip>
' a ' Hl ie
5 ' Pog 6 1 f ' i 2D. estes
aa Rai hee pa car ara “oth taale ete
: i ga ; wary
Ta oe 2 wy tat Uae: Relat ep Pu ;
. 7 * , ¢ ‘ i ne We
Bd 1%} AC i Tae 2 ‘ ' ” nite Hh re
bey ay Rae Me G4.
; 5 We ile 20 eee at

SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLIWS
Fen DE ae eh cs -
e) see Ss) ai Sie ‘e)
5 “2 5 a> NX §
Es 3 = > Way =
ne - : * a
w
i z e we i - =
NOILNLILSNI SStYVYaIT LIBRARIES. SMITHSONIAN
z= w Zz: eae wn
< = Bed SS =
zZ ~ ma =
© Sis 2. [eye 2
Ww Qe 9) Ww
Ae Oy 5) Oo
ae x os =
e = 2 Ye

Ss

SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS

LIBRARIES SMITHSONIAN

Sa1uve

a WY ——,
on ee tar: a i
WL ME A 2 =
ao Vet Sc oc e
a “Wg = B 2
ay : a a, ra ~
NOILALILSNI NVINOSHLINS S3SIY¥VYdIT LIBRARIES SMITHSONIAN:
ae = 1m 4 2 va
_ E S = 50
= = p ;
> _ > i 5,
= 2 : 3
eS oe m 2 m shy
ep) — ” aa 2) We
SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3S14Va
” = * w = ” eae
| = < = Wy < = UY
\ “A Pi cot ith fy 2 = Yo
. es Oo ar Lp, fo) ma tp
RCREQR. 6) anh 22 JOE 2) w Gd
, \ : a Vp, Ko Wee)
ae a lee = ner = at
VOILALILSNI NVINOSHIIWS SJiuVug Il. LIBRARIES SMITHSONIAN INSTITU
= ie ae ih c 5 > py
n at ” “E a7)
: : 2Q™= $o%
- LG,
Ss We S S
TSE, ee mM —a m 2
S ere Ss = =.
MPERARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31 uve
; = z E 2 a.
= fom EE a ve 5
Ey 2 F = We
Ai : ¥ = E
ag on z a = "aa
eo tuYvuad riot BRARI ES _ INSTITU
, < = i < et = < S ‘ \
= = 4 = = z
D ee Di. wn D
: Si 2 : =z WA
i 2 Bo ee a)
SMITHSONIAN INSTITUTION NOILNLILSNI SS buvy
mi 2 Sot in 2 oe
” ae ” wae oo
ie: iia x a reat
< 3; ps << = <x |
ae Sy _— © ;
. — Oty oneee =p

ES SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLINS SJZIUY:

N

ad Cr pa =
ie ie cS) a:
ke of ah > 0 XE ria
2 | ‘ Lay Yh: SD eS ON ‘ane |
a Gye cad ne VE iE
Det “Ufy ma - Ss, is
” 4 ih <

a Cl z ee

SNI NVINOSHLINS S3IYVYEIT LIBRARIES

NVINOSHLINS S3iuVvudit

NYINOSHLIWS
SMITHSONIAN

S SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS Sa1uy

SMITHSONIA
SMITHSONIAN

#9 z rg 2 6 we

ul 2 _ a x “

Na. can a NG

- = : ZMK =

mo = £0 = SQ co

Mas Zz ng 2 a TG
ISNI NVINOSHLINS S3SIYVUdIT LIBRARIES SMITHSONIAN _INSTITL

| he = i S eh a

or = oo = tg OD

70 Xs = Pe) 2 Ly 70

F VSS E z - YG f/ =

aS SS = “ = OL se

m WW es a ei m

w rf = oO = wn

M1ES SMITHSONIAN INSTITUTION NOILMLILSNI_ NVINOSHLINS SAIUY:

WO Fad on = ”
x z = y Lp Zz of Y
ee oe oO or . Gee ea Cay re i Y
me P & Uy; ® wy
2 - 4) YG fe? i Ae a allege
> Bo. =
z a 2 - 2
ma ne LIBRARIES INSTITU
J os wo prea
iG ul a ws a
eed ox na oc a
c o cs pe cE
o — --{
= ee) or mm. ae.
Oo ~ Oo a oO
Die, ae me ay =
eS Ee eeONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYV
oe z |
S ha S S
- on to i
is ea =) =)
E: a E E
= wn = z

NI_NVINOSHLINS S31UVYSIT_ LIBRARIES

SMITHSONIAN INSTITI

NVINOSHLINS SaIuVvugit

(op) Pa

z = ae <

fiZ =f ne as

oO fa Ne EM oO

(ep) ” yy (op) YW)

a (@ ye ae i

= Z iE e

i % = > ‘ , = i =
RIES SMITHSONIAN INSTITUTION NOILMLILSNI NVINOSHLINS S3IYV

NOILNLILSNI

LIBRARIES

NOLLALILSNI
4
Yip
pf
LIBRARIES

_IBRARIES

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

Vol. 29 December, 1966

VOLUME 29

Editor
PIERCE BRODKORB

= or e ee,

— “~~ Ep =
A 1 bj *
GNU SUN, 4

Published by the

FLORIDA ACADEMY OF SCIENCES
Gainesville, Florida
1966

PUBLICATION DATES OF VOLUME 29

Number 1: June 17, 1966
Number 2: September 8, 1966
Number 3: October 20, 1967
Number 4: February 12, 1968

New Taxa PROPOSED IN VOLUME 29

Actinocleidus georgiensis Price (Trematoda: Dactylogyridae )
Heteronchocleidinae Price (Trematoda: Dactylogyridae )
Anacanthorninae Price (Trematoda: Dactylogyridae )

Kochlorine floridana Wells and Tomlinson (Cirrepedia: Alcippidae )
tNotophthalmus slaughteri Holman (Amphibia: Salamandridae )
Hyla grandisonae Goin (Amphibia: Hylidae )

+Hyla miocenica Holman (Amphibia: Hylidae )

+ Leiocephalus jamaicensis Etheridge (Reptilia: Iguanidae )
Diadophis punctatus acricus Paulson (Reptilia: Colubridae )

+Floridatragulinae Patton (Mammalia: Camelidae )

+ Fossil

ii

76
200
200

27
269

39
270

49
296
182

CONTENTS OF VOLUME 29

NUMBER 1

Photoelectric photometry of 44i Bootis
K-Y. Chen and D. A. Rekenthaler

The Mannich reaction of 2-naphthol with secondary amines
J. E. Fernandez and W. I. Ferree, Jr.

Verification of Monotropa hypopithys in Florida Daniel B. Ward
A Pacific polychaete in southeastern United States John L. Taylor

A new burrowing barnacle from the western Atlantic
Harry W. Wells and Jack T. Tomlinson

Acetes shrimp on the Florida East Coast Edwin A. Joyce, Jr.
A new frog of the genus Hyla from British Guiana Coleman J. Goin
A second specimen of Ophisaurus ceroni J. Alan Holman

An extinct lizard of the genus Leiocephalus from Jamaica
Richard Etheridge

Growth and longevity of rats fed different diets
L. R. Arrington and C. B. Ammerman

Effect of vitamins A and FE on copper in heart of cattle
R. L. Shirley, H. L. Chapman, Jr., J. F. Easley, and T. J. Cunha

NUMBER 2
Organotin esters McDonald Moore and Francis C. Lanning
A new monogenetic trematode from Georgia Charles E. Price
Barnacles of the northeastern Gulf of Mexico Harry W. Wells
Minima of eclipsing binaries VV Orionis and Beta Persei K-Y. Chen
New records of Bahamian Odonata Dennis R. Paulson

Hermaphroditism in mullet, Mugil cephalus Linnaeus
Martin A. Moe, Jr.

Review of the Lutjanus campechanus complex of red snappers
Luis R. Rivas

Centrarchid spawing in the Florida Everglades James P. Clugston

The exotic herpetofauna of southeast Florida /
Wayne King and Thomas Krakauer

The vertebral musculature of Chersydrus (Serpentes) Walter Auffenberg

iil

43

73
OTs
81
96
97

Ii1

ils
137

144
155

NUMBER 3

Variscite from the Hawthorn Formation
Frank N. Blanchard and Stephen A. Denahan

An investigation of ostracode preservation Mervin Kontrovitz
Florida Academy of Sciences award for 1967

Revision of the selenodont artiodactyls from Thomas Farm
Thomas H. Patton

Distribution of Euglenida in North Florida John J. McCoy
Two new subfamilies of monogenetic trematodes CE} Price
Mayfly nymphs from northwestern Florida Robert F. Schneider

Caribbean recruitment of Florida’s spiny lobster population
Harold W. Sims, Jr., and Robert M. Ingle

NUMBER 4

Organotin esters and their reaction with Grignard reagents
McDonald Moore and F. C. Lanning

Wavellite-cemented sandstones from northern Florida
Frank N. Blanchard and Stephen A. Denahan

Notes on spiny lobster larvae in the North Atlantic
Harold W. Sims, Jr.

Account of an octopus bite Arthur C. Wittich
A small Miocene herpetofauna from Texas J. Alan Holman
Diet of the bowfin in central Florida Michael C. Diana

First Gulf of Mexico record for Lutjanus cyanopterus
Martin A. Moe, Jr.

The molid fish Ranzania laevis in the western Atlantic
C. Richard Robins

Iodine toxicity in relation to hormones L. R. Arrington,
R. N. Taylor, C. B. Ammerman, and A. C. Warnick

Variations in some snakes from the Florida Keys Dennis R. Paulson

Officers and members of the Academy

163
Teyal
178

179
191
199
202

207

243

248

257
265
267
276

285

287

290
295
309

>
\\

rGrEs

Quarterly Journal
of the

=! Florida Academy of Sciences

Vol. 29 March, 1966 No.

CONTENTS

Photoelectric photometry of 44i Bootis
K-Y. Chen and D. A. Rekenthaler

The Mannich reaction of 2-naphthol with secondary amines
J. E. Fernandez and W. I. Ferree, Jr.

Verification of Monotropa hypopithys in Florida Daniel B. Ward
A Pacific polychaete in southeastern United States John L. Taylor

A new burrowing barnacle from the western Atlantic
Harry W. Wells and Jack T. Tomlinson

Acetes shrimp on the Florida East Coast Edwin A. Joyce, Jr.
A new frog of the genus Hyla from British Guiana Coleman J. Goin
A second specimen of Ophisaurus ceroni J. Alan Holman

An extinct lizard of the genus Leiocephalus from Jamaica
Richard Etheridge

Growth and longevity of rats fed different diets
L. R. Arrington and C. B. Ammerman

Effect of vitamins A and E on copper in heart of cattle
R. L. Shirley, H. L. Chapman, Jr., J. F. Easley, and T. J. Cunha

Mailed June 17, 1966

1

AT

60

67

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Cargon Copy will facilitate review by referees.

Marcins should be 1% inches all around.

Tires should not exceed 42 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and
ADDRESS in paragraph form following Literature Cited.

LrreRATURE CrTeD follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLES are charged to authors at $16.70 per page or fraction. Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

LecENps for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($15.80 per page, $14.00 per half
page for zinc etchings; $14.50 per page, $13.25 per half page for halftones).
Drawincs should be in India ink, on good board or drafting paper, and let-
tered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. PHoTocRAPHS should be of good contrast, on glossy paper. Do not
write heavily on the backs of photographs.

ProoF must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FLORIDAVAGADEMY (OF SGIENGES

Vol. 29 March, 1966 No. l

Photoelectric Photometry of 441 Bootis
K-Y. CHEN AND D. A. REKENTHALER

PHOTOELECTRIC photometry of eclipsing variable stars at the
University of Florida was begun in January of 1965. The Univer-
sity Observatory, shown in Fig. 1, is located on the north shore of
Bevin’s Arm lake, Gainesville, Florida. The first star under sys-
tematic observation is 441 Bootis. This stellar system was first re-
ported as a binary by Herschel in 1781. It was later found to
be a triple system: a visual binary with an eclipsing variable as
one component of the visual pair. Numerous investigations of
this variable’s light curve have been made by photographic and
photoelectric means, the most recent being of Binnendijk (1955).
Observation of this star on 16-17 April 1965 is given here.

2 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

EQUIPMENT

The equipment used for observation is described in this sec-
tion. Figure 2 illustrates the amplifier power supplies and the re-
corder. The telescope with the photometer and DC amplifier
mounted for observing is shown in Figure 3.

Fig. 2. Amplifier power supplies and recorder.

CHEN AND REKENTHALER: Photoelectric Photometry 8

Telescope. The telescope, built by the Cave Optical Com-
pany, is housed in a sliding roof observatory. It is an f/7.6 New-
tonian reflector with a focal length of 96 inches, supported by an
equatorial mount. A 3-inch refractor is mounted alongside the
reflector for use as a finder scope. The telescope is electrically
driven and has an accessory control box which allows the ob-
server to change the tracking rate in right ascension, and also to
change the direction in declination. Two setting circles, a right
ascension circle and a declination circle, are installed on the axes
of the mount.

Photometer. As the light rays pass from the telescope into the
photometer, they travel first through a small aperture, then through
a filter, and finally through a Fabry lens before impinging on the
phototube. Four diaphragms are at the observers disposal. These
are small circular openings of 0.500, 0.250, 0.079, and 0.039 inch
diameters placed on a disc which can be rotated to select the prop-
er size aperture. The smallest opening (0.039) was used in this
work.

The viewing eyepiece for observing the star visually follows
the aperture in the light path. The eyepiece allows visual obser-
vation of the star only if it is “pushed in” to the photometer. De-
flection of the light rays into the eyepiece is accomplished by use
of a prism. This also blocks the light from reaching the phototube.
With the eyepiece withdrawn, the light path is unobstructed and
a signal can be generated. The outline of the particular aperture
in use can also be seen when the eyepiece is used, as the aperture
is illuminated by a battery-powered bulb. When the eyepiece is
withdrawn to take a photoelectric reading, the bulb is switched off
by a micro-switch. The starlight then passes through a filter.

The filters are mounted on a slide so that the observer can
select one of the ultraviolet (Corning 9863), blue (Corning 5030 and
Schott GG13), and yellow (Corning 3389) filters, which correspond
to the Johnson-Morgan standard U-B-V system.

The Fabry lens serves to converge the bundle of light rays
onto a small area on the surface of the photocathode. It is made
of fused quartz with an anti-reflecting coating on the surfaces, a
physical diameter of 13+1 mm, and a focal length of 1941 mm.

The filtered converging rays impinge on the cathode of the
RCA 1P21 multiplier phototube at the end of the light path.

4 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

CHEN AND REKENTHALER: Photoelectric Photometry 5

One of the three external electrical connections to the photom-
eter leads from the high voltage power supply. The second is
the 12-volt line from the battery to the bulb lighting the aperture.
The third is the short shielded line carrying the output of the 1P21
to the DC amplifier.

High Voltage Power Supply. The power for the phototube
comes from a commercial unit, the Victoreen model HV-25CT.
This provides a stable 1000 volt DC potential which is reduced to
approximately 75 to 150 volts between any two dynodes by means
of a series resistance voltage divided. This voltage divider is
essentially a series of resistors connected directly to the 1P21 tube
socket pins.

DC Amplifier. The DC amplifier is used to amplify a cur-
rent, in the order of a nano-ampere, from the phototube to a level
detectable by the recorder. The properties most important to an
amplifier in a photoelectric system are its linearity and gain sta-
bility. In the amplifier used here these qualities are met by using
large negative feedback. The first stage of the amplifier consists
of an ‘electrometer’ tube of high input impedance. This is put
in parallel with a large load resistor.

The size of the load resistors is determined so as to change the
amplification in increments of 2.5 magnitudes, from 0 to 12.5. The
feedback resistors are in increments of 0.25 magnitude from 0 to
2.50. Magnitude here refers to stellar magnitude. Different com-
binations of these resistors are used to produce the suitable signal
on the recorder chart. The combination of resistors desired is se-
lected with two rotary switches mounted on the amplifier. The
time constant of the amplifier is controlled by a third rotary switch
which gives a choice of four time constants.

The electric circuit of the amplifier is similar to the one de-
scribed by Whitford (1962).

Amplifier Power Supplies. The amplifier requires DC volt-
ages of 1250, —105, +67.5, +4.5, and also filament power. The
+250 and —105 DC voltages, and the 6.3 AC voltage are supplied
by two regulated power supplies of conventional design.

A battery box supplies the +67.5, +4.5, and 1.5 voltages for
the electrometer tube. The filament of the electrometer tube is
wired with a potentiometer in the circuit. By varying the filament
current with this potentiometer, an accurate zero point balancing
of the recorder is possible.

6 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

JD Hel
2430000-+

8867.5771
.o795
0825
0858
.0896
.O916
0943
.6032
.6091
.6146
.6214
.6260
.6289
G32
.6348
.6397
.6416
.6491
.6591
.6612
.6678
6757
.6809
.6832
6867
.6893
6923
.6957
.6983
7101
VOX
.7283
SS)
£7347
£7429
.7493
£7509
.7596
.7618

Yellow Observation of 44i Bootis

Phase

0.4155
0.4243
0.4358
0.4479
0.4623
0.4697
0.4796
0.5130
0.5348
0.5555
0.5809
0.5981
0.6087
0.6229
0.6307
0.6492
0.6561
0.6844
O25
0.7295
Ora 2i
0.7837
0.8032
0.8117
0.8247
0.83438
0.8455
0.8584
0.8680
0.9121

‘0.9717

0.9800
0.9956
0.0039
0.0345
0.0586
0.0832
0.0970
0.1053

TABLE 1

Am

—0.821
0.780
0.884
0.814
0.830
OMOM
0.750
0.788
0.765
0.809
0.891
0.796
0.845
0.918
0.880
0.933
0.875
0.869
0.957
0.940
O:07
0.893
0.950
0.942
0.910
0.902
0.910
0.925
0.887
0.889
0.816
0.745
0.787
0.769
0.766
0.856
0.869
0.838

ae ORO

JD Hel
2430000-+-

8867.7659
.7684
.7835
.7895
.7916
MOON
.7980
.8087
8139
8165
8197
8219
8271
8308
8334
8374
8399
844]
8466
8507
8532
8567
8594
8632
8657
8705
8732
.8766
8789
8823
8846
8879
8900
8933
8953
8986
9008
9039

Phase

0.1203
0.1299
0.1862
0.2087
0.2165
0.2318
0.2401
0.2803
0.2997
0.3093
0.3213
0.3296
0.3490
0.3628
0.3724
0.3874
0.3967
0.4125
0.4216
0.4372
().4462
0.4595
().4696
0.4839
0.4929
0.5108
0.5209
0.5336
0.5422
0.5549
0.5637
0.5759
0.5837
0.5961
0.6036
0.6161
0.6241
0.6355

Am

—0.844
0.858
1.048
1.037
1.028
0.949
1.033
0.913
0.881
0.925
0.972
0.927
0.877
0.854
0.860
0.906
0.872
0.878
0.819
0.827
0.791
0.857
0.828
0.796
0.813
0.823
0.814
0.843
0.818
0.805
0.803
0.837
0.865
0.899
0.876
0.903
0.891
Soe ORC E

CHEN AND REKENTHALER: Photoelectric 'Photometry 7

Recorder. The recorder is a “Speedomax G” built by the Leeds
and Northrup Company. It has a single pen which marks the
continuous strip-chart to correspond with the signal current from
the amplifier. The chart is set to run at a rate of 1 inch per minute.

The recorder responds to signals in the 0 to 10 millivolt range,
a 10 millivolt signal giving a full deflection of the pen (9-1/2
inches).

Miscellaneous Accessories. The observatory is equipped with
a high frequency receiver for receiving WWV time signals on
either 5, 10, or 15 megacycles.

Two electric clocks, one for sidereal time and the other on
Eastern Standard Time were used, as well as a barometer, ther-
mometer, and hygrometer. Readings of all these instruments were
made periodically and the final reading recorded nightly in the
observatory log.

OBSERVATION AND Data REDUCTION

The stars observed on 16-17 April 1965 were 44i Bootis, whose
right ascension (1900) was 15°0"30*, declination (1900) +48° 3’, vis-
ual magnitude 4.76 and spectral type Gl, and the comparison
HR 5581, whose right ascension (1900) was 14°53"4°, declination
(1900) +50° 2’, visual magnitude 5.62 and spectral type F7.

A definite sequence of observations was followed. First the
zero point was set by balancing the equipment with the amplifier
gain at the minimum value. The dark current of the 1P21 was
measured by turning the amplifier gain to a pre-selected setting.
The dark current reading was followed by a measurement of the
sky background. This was accomplished by withdrawing the eye-
piece of the photometer while the telescope was pointed to an
area of sky totally devoid of stars (or apparently so) and close to
the variable. After the sky deflection had been made, the com-
parison star was centered in the aperture and readings made of
its light. The variable reading was made next and a continual
sequence of these readings followed in a general order: sky-com-
parison-variable-sky-comparison-variable-etc. Each object was ob-
served through both the blue and yellow filters with the individual
readings lasting about 45 seconds. For the stars observed, the re-
sponse of the phototube when used with the ultraviolet filter was
not sufficient to make a signal change detectable from the back-

8 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

JD Hel
2430000+

8867.5775
.9799
5843
.0900
0919
947
0980
.6036
.6075
.6094
.6150
.6221
.6271
.6330
.6353
.6402
.6419
.6474
.6497
.6527
6549
6595
.6618
.6656
.6680
.6739
.6763
.6816
.6873
.6898
.6927
.6962
.6993
£7032
.7107
.7264
.7289
.7330
Miao
7412
7434

Phase

0.4168
0.4259
0.4422
0.4638
0.4707
0.4811
0.4933
0.5146
0.5291
0.5361
0.5569
0.5835
0.6022
0.6242
0.6328
0.6509
0.6574
0.6779
0.6865
0.6976
0.7047
0.7233
0.7319
0.7459
0.7547
0.7770
0.7858
0.8055
0.8270
0.8364
0.8472
0.8602
0.8719
0.8862
0.9142
0.9730
0.9821
0.9976
0.0054
0.0282
0.0363

TABLE 2

Blue Observation of 44i Bootis

Am

—(.663
0.668
0.696
0.649
Or 22,
0.673
0.612
0.664
0.668
0.656
0.646
0.729
0.745
0.759
0.723
0.762
0.779
0.790
0.796
0.768
0.818
ORF
0.783
0.832
0.778
0.789
0.800
0.768
0.802
0.750
0.759
0.782
0.833
0.781
0.741
0.629
0.659
0.629
0.643
0.625

—().654

JD Hel
2430000+-+

8867.7476
£7502
.7566
7603
.7624
.7667
7691
.7840
.7900
ofA
£7962
Moos
8049
8091
8150
8204
8224
8256
8280
6320
8384
8411
8447
8471
8513
8537
.8600
8639
8661
8716
.8770
8794
8852
8884
8905
8939
8950
8993
9012
9045
9064

Phase

0.0521
0.0617
0.0858
0.0995
0.1074
0.1234
0.1325
0.1880
0.2103
0.2183
0.2336
0.2448
0.2660
0.2816
0.3036
0.3238
0.3314
0.3428
0.3524
0.38672
0.3913
0.4011
0.4146
0.4237
0.4393
0.4483
0.4716
().4864
0.4948
0.5150
0.5355
0.5443
0.5660
0.5780
0.5858
0.5984
0.6057
0.6184
0.6257
0.6379
0.6451

Am

—0.690
0.677
0.672
0.793
0.741
0.805
0.787
0.806
0.860
0.959
0.816
0.886
0.810
0.825
0.831
0.856
0.858
0.768
0.817
0.819
0.829
0.828
0.786
0.769
0.755
0.666
0.745
0.647
0.717
0.712
0.758
0.655
0.687
0.774
0.784
0.786
0.774
0.772
0.789
0.842

OMG

CHEN AND REKENTHALER: Photoelectric Photometry 9

2438867.5700 867.7400 867.9100
JD Hel

Tt oe

0.5 0.0 0.5
PHASE

Fig. 4. Yellow and blue observations of 44i Bootis on 16-17 April 1965.

10 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ground noise. Infrequently, the check star HR 5635 was observed,
and also the zero point and the dark current were occasionally
checked. The number of sky readings was sometimes reduced if
no change in the sky was apparent. Records of the time, tempera-
ture, relative humidity, and barometric pressure were periodically
recorded on the continuous chart.

To make the strip-chart data more usable it was later reduced
to tabular form. Since the chart ran continuously at a constant
rate, the Eastern Standard Time of each variable reading could
be determined. This was converted to a heliocentric Julian Date
(JD Hel) by using the American Ephemeris and Nautical Almanac.
The phase of each variable reading was calculated from the set

of light elements given by Schneller (1964):
JD Hel Min = 2437362.6179 + 0.:267814191 E.

The difference in magnitude, 4m, between the variable star and
the comparison star at the time of a variable reading was deter-
mined from the following relation:
1 (comp.)
Am = 2.5 Log ————
] (var.)

where |(comp.) is the apparent luminosity of the comparison star
and | (var.) that of the variable; and this ratio is of course equal
to the ratio of chart-readings of the two stars with sky readings
subtracted. Linear interpolation was employed wherever value
between readings was needed. The values, (JD Hel, Phase, 4m),
for the yellow and blue observations are listed in Tables 1 and 2
respectively, and plotted in Fig. 4.

DISCUSSION AND CONCLUSION

Earlier observations of the triple system 441 Bootis by Schilt,
Oosterhoff, Stebbins and Huffer, Shapley and Calder, Plaut, and
Eggen, as summarized in Binnendijk (1955), have shown that its
mean light curve changes. Eggen (1948) has shown that the light
curve varied from day to day with irregular variation outside the
eclipses and pronounced change of depth of the secondary mini-
mum (in the order of 0".04). The present work (refer to Fig. 4)
shows: (1) the variation of depth of the secondary minimum from
cycle to cycle, approximately 0”.033 in the yellow observation;

CHEN AND REKENTHALER: Photoelectric Photometry 11

(2) conspicuous “brightening up” about phase 0.2 during the time
of observation, approximately 0".1 in the yellow observation; and
(3) difference in light curves ih the two colors around phase 0.25.
The probable error for each yellow observed point is estimated
to be + 0".017, and the same value for each blue observed point.

The eclipsing component of 44i Bootis is of the W Ursae
Majoris type. Variations of light curves of eclipsing binaries of
this type are not uncommon. Here the picture is further compli-
cated by several indications in earlier observations that the bright-
er, non-eclipsing component of the visual pair is slightly variable.
The study of the nature of the variation of its light curve is an in-
teresting, challenging, and unsolved problem. It is not until the
two components of the visual binary are observed separated that
new information may be obtained.

To conclude, the words of Harlan J. Smith (1963) are quoted
here:

“Variable stars will normally occupy amateur or small-scale
professional photoelectric observatories, but a rich array of other
applications exists. Variable stars alone can provide a generation
of work for small telescopes equipped with good modern pho-
tometers, while the super-photocells and photometers ultimately
available will make twelve-inch telescopes equal to present thirty-
inch ones in limiting magnitude, greatly extending their useful life.

“,.., nature has not yet set the limits on what can be achieved
in astronomical photometry—the future is still bounded primarily
by our energy, our ingenuity, and our insight into astronomy.

ACKNOWLEDGMENTS

The program of photoelectric photometry of variable stars at
University of Florida was made possible through funds furnished
by the College of Arts and Sciences and the Graduate School of
the University. The continued support of the National Aero-
nautics and Space Administration for this program is greatly ap-
preciated. Special thanks are due to Drs. S. S. Ballard and A. G.
Smith of the Department of Physics and Astronomy for their ad-
vice and encouragement, to Dr. W. M. Protheroe of the University
of Pennsylvania for his generosity in providing the circuitry of the
DC amplifier, to Messrs. H. W. Schrader and R. B. Warren and the
staff of the Machine Shop, Department of Physics and Astronomy,

12 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

for the constructions of the photometer and the sliding roof of the
observatory, to Mr. J. May for constructing the DC amplifier and
the amplifier power supplies, and to Mrs. B. Keyser for typing the
manuscript.

LITERATURE CITED

BINNENDIJK, L. 1955. The light variation and orbital elements of 44i Boo-
tis. Astron. Jour., vol. 60, pp. 355-363.

EccEn, O. J. 1948. The system of 44i Bootis. Astrophys. Jour., vol. 108,
pp. 15-27.

SCHNELLER, H. 1964. i Bootis. Information Bulletin on Variable Stars, no.
50, Konkoly Observatory, Budapest.

Smiru, H. J. 1963. The place of photometry in astronomy. Photoelectric
Astronomy for Amateurs, ed. by F. B. Wood, Macmillan, New York,
p. 42.

WuitForp, A.E. 1962. “Photoelectric techniques,” Encyclopedia of Physics,
vol. 54, ed. by S. Fliigge, Springer-Verlag, Berlin, p. 262.

Department of Physics and Astronomy, University of Florida,
Gainesville, Florida; Air Force Weapons Laboratory, Air Force
Special Weapons Center, Albuquerque, New Mexico.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

The Mannich Reaction of 2-Naphthol with Secondary Amines
J. E. FERNANDEZ AND W. I. FERREE, JR.

THe Mannich reaction involves the reaction of a non-tertiary
amine, formaldehyde, and a compound having a hydrogen atom
which is labile through proximity to an electron attracting group
(Reichert, 1959): It is an important reaction both synthetically for
the production of compounds with a terminal methylene group,
and biologically because it is thought to be involved in the bio-
genesis of alkaloids (Reichert, 1959). In addition, the Mannich
reaction is interesting chemically because it seems to proceed
through several mechanisms of nearly equal probabilities. There-
fore, the mechanism can change abruptly with subtle changes in
reaction conditions or reactants (Fernandez, 1964; Fernandez,
Fowler, and Glaros, 1965). For this reason, the mechanism of the
Mannich reaction has been studied by several workers who have
found seemingly conflicting results (Burke, Nasutavicus, and
Weatherbee, 1964).

The use of aqueous solvents complicates studies of this reaction
because several species are formed, all of which may act as in-
termediates. It has proved a difficult task to derive meaningful
results from such systems. For example, it has been generally es-
tablished that the reaction proceeds through the initial reaction
of amine with formaldehyde to form two intermediates, (I and II),
either or both of which should be capable of reacting with the
active hydrogen compound (HZ) to form the product (Lieberman
and Wagner, 1945; Butler, 1956; Fernandez, 1964).

RoNH + CH,0 === R,NCH,OH ———= R,NCH,NR, + H,0
(I) (1)

I + HZ——-R,NCH,Z + H,0

Il + H2—~ R,NCH,Z + HNR,

We have therefore been engaged in studying several examples
of this reaction in aprotic solvents. In this way the intermediates
can be studied individually without danger of interconverting
them.

Recently, evidence has been offered to support a mechanism
in which the active hydrogen compound reacts in its enol form

14 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

through a cyclic transition state. The examples studied were
(Burckhalter and Lieb, 1961):

Abe
OH 0”) Cf
GS HNR,
+ RNCH,X ae aa
(X = OH or NR)
0
ce jNR, oo 2NR,

and (Fernandez and Fowler, 1964, 1965):

/ / jo / »
R,CHNO,== RCN. R NCH, X Roe \
On cH 0 :
, On Y
R,NCH,CR,NO, + HX RN YO

In these cases, the cyclic transition state involves hydrogen
bonding between the enolic OH and the X group.

We now wish to add evidence for this proposition through
reactions of some 2-naphthol derivatives in which the phenolic
hydrogen is absent. In addition we wish to report a study of
the formation 1,1’-methylenedi-(2-naphthol) as it is related to the
Mannich reaction of 2-naphthol.

Since 2-naphthol undergoes the Mannich reaction only at the
alpha position, it provides a simple model for studying the role
of hydrogen bonding. We find that in the absence of the phenolic
hydrogen atom, reaction does not occur. Thus, reactions of 2-
methoxynaphthalene fail to yield any Mannich base on reaction
with piperidine or morpholine and formaldehyde (Table 1, runs 11,
12). This may be explained either by the lack of hydrogen bond-
ing, or by the relatively low electron-releasing effect of the meth-
oxy group as compared with the oxy anion which is undoubtedly
present in equilibrium with 2-naphthol under the basic Mannich
conditions. The latter alternative is not attractive in view of the
fact that reactions employing sodium naphthoxide run under an-
hydrous conditions yield only small quantities of Mannich base

FERNANDEZ AND FERREE: Mannich Reaction 15

(Table 1, runs 7, 8, 9). The lack of reactivity of 2-methoxynaph-
thalene is notable since this compound condenses with formalde-
hyde in acid under mild conditions to form 1,1’-methylenedi(2-
methoxynaphthalene) (Runti and Collino, 1959).

IANe Ls, Al

Mannich reactions of 2-naphthol, 2-methoxynaphthalene, and
sodium 2-naphthoxide

Percent Yield of Product*®

2-Naphthol

or PMN MBBN

Run 2-Methoxy- or or
No. Reactants*, ft naphthalene PMMN MBMN
1. Naphthol. + MBP 0 99 0
2, Naphthol + C,H,,N.HCl + CH,O 45 52 0
3. Naphthol + C,H,,N.HCl

peerl5O | 0.009 HCl 0 0 99
4. Naphthol + C,.H,,N + CH,O 0 84 0
5. Naphthol + CH,O 0 0 86
6. MB2N + C;H,,N + 0.009 HCl 0 0 82
7. Sodium 2-naphthoate + MBP De 18 0
8. Sodium 2-naphthoate + MBM 66 0 0
9. Sodium 2-naphthoate + PMN 0 2 95
10. Naphthol + PMN + 0.018 HCl 56 42 0
11. 2-Methoxynaphthalene -+- MBP 99 0 0
12. 2-Methoxynaphthalene +

C.H,,N + CH,O 99 0 0

*MBP = methylene bispiperidine, C;sHi:1N = piperidine, MB2N — meth-
ylene bis(2-naphthol), MBM = methylene bismorpholine, PMN = 1-piperidi-
nomethyl-2-naphthol, PMMN = _ 1-piperidinomethyl-2-methoxynaphthalene,
MBMN = methylene bis(2-methoxynaphthalene).

tAll quantities were 0.01 mole unless otherwise specified. Ten milli-
liters of dioxane was used as solvent in all runs except those employing
sodium 2-naphthoate in which N,N-dimethylformamide was used.

The data in Table 1 indicate that the Mannich reaction em-
ploying 2-naphthol can yield either the expected Mannich base (I)
or the 1,1’-methylenedi(2-naphthol) (II) depending upon conditions.
In the absence of catalysts, the major product is the Mannich base
(runs 1, 4). In the absence of amine, II forms in good yields.

16 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

H_NR.

plesk CHO + HNR, Ge. OH
OH HO

I (I)
aoe

(IL)

The use of amine hydrochloride results in a reduced yield of I
(run 3), while an excess of acid results in a quantitative conversion
to II (run 2). This variation of product with catalyst can be ex-
plained either (a) by the participation of I as an intermediate in
the formation of II, or (b) by assuming that the relative rates
of formation of I and II are greatly altered and II becomes the
predominant product in the presence of excess acid. Alternative
(a) is ruled out by the failure of I to react with 2-naphthol in the
presence of excess acid (run 10). Our conclusion is, therefore, that
I and II are formed in parallel reactions. The irreversibility of the
formation of II is shown by the results of run 6. In basic medium
however, the reaction to form II may proceed through I as an in-
termediate (run 9).

We conclude from these experiments that a hydrogen bonded
species, such as III, is essential to the formation of the Mannich

Cis (X = OH or NR,)

bases of 2-naphthol. In the presence of excess acid, such hydrogen
bonding is prevented by the formation of the ammonium ion of
the secondary amine or methylene bisamine. In the strongly basic
media, hydrogen bonding is prevented by the formation of phen-
olate ion.

An increase in acidity increases the phenol/phenolate ion ratio
while simultaneously decreasing the amine/ammonium ion ratio.
Therefore, the pH at which the maximum amount of free phenol

FERNANDEZ AND FERREE: Mannich Reaction Liz

and free base can coexist should be the most favorable for forma-
tion of the hydrogen bonded species (III). This hypothesis is sup-
ported qualitatively by the kinetic studies of Alexander and Un-
derhill (1949) and Cummings and Shelton (1960) if the enol of the
ketone corresponds to the phenol in our work. These studies dem-
onstrated an optimum pH of ca. 4 when the active hydrogen com-
pound was ethylmalonic acid, and of ca. 11 when the active hy-
drogen compound was cyclohexanone. In both cases dimethyl-
amine and aqueous solutions were employed. In the former case,
the strongly acidic ethylmalonic acid requires a strongly acid so-
lution to insure sufficient quantities of the undissociated acid.
In the latter case, a much more basic solution can be tolerated
while maintaining the cyclohexanone primarily in the undissoci-
ated form.

A similar conclusion was reached by Fernandez and Fowler
(1964) in a study of the reaction of 2-nitropropane with methylene
bisamines.

EXPERIMENTAL

Materials. The chemicals used were commercial products of
the highest purity available; they were further purified when nec-
essary. Temperatures are uncorrected.

Mannich Reactions of 2-Naphthol and Piperidine and. Morpho-
line. Table 1 contains the results of reactions of 2-naphthol, 2-
methoxynaphthalene, and sodium 2-naphthoxide with piperidine
or morpholine and formaldehyde. All the reactions were run by
allowing 0.01 mole of each substrate in 10 ml. dioxane to react at
room temperature for twenty hours, and worked up according to
the procedures outlined below.

Procedure A. Reactions employing 2-naphthol or 2-methoxy-
naphthalene were worked up by adding water dropwise until the
solution became turbid, adjusting to pH 4-6 with dilute HCl, and
extracting with ether. The ether was dried over CaSO, and evap-
orated to yield 2-naphthol, 1,1’-methylenedi(2-naphthol) or 2-
methoxynaphthalene. The HCl layer was neutralized with NaOH
and extracted with ether. The ether was dried over CaSO, and
evaporated to yield Mannich base. The products were identified
by melting point and mixture melting points with authentic sam-
ples.

Procedure B. Reactions employing sodium 2-naphthoxide were

18 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

worked up by evaporating off most of the solvent under vacuum,
then adding ether to precipitate unreacted sodium 2-naphthoxide.
This salt was filtered out, hydrolyzed, the solution adjusted to pH
6-7, and recovered as 2-naphthol. The ether filtrate was extracted
with dilute HCl to remove Mannich base and unreacted amine.
The acidic solution was then neutralized with 1 M NaOH, and
the resulting precipitate or oil extracted with ether. The ether
and amine were then evaporated to leave the Mannich base. The
ether layer resulting from the HCl extraction was also evaporated
to recover unprecipitated 2-naphthol and 1,1’-methylenedi(2-naph-
thol). The products were identified by melting points and mix-
ture melting points with authentic samples.

ACKNOWLEDGMENT

This work was conducted while the junior author was a stu-
dent participant under National Science Foundation science edu-
cation grant No. GE 1210.

LITERATURE CITED

ALEXANDER, E. R., AND E. J. UNDERHILL. 1949. Studies on the mechanism
of the Mannich reaction. I. Ethylmalonic acid, a methynyl compound.
Jour. Amer. Chem. Soc., vol. 71, pp. 4014-4019.

Burke, W. J., H. A. NasutTavicus, AND WEATHERBEE, 1964. Synthesis
and study of Mannich bases from 2-naphthol and primary amines.
Jour. Org. Chem., vol. 29, pp. 407-410.

BuRKHALTER, J. H., AND R. I. Lies. 1961. Amino- and chloromethylation
of 8-quinolinol. Mechanism of preponderant ortho substitution in phe-
nols under Mannich conditions. Jour. Org. Chem., vol. 26, pp. 4078-
4083. “

Butter, G. B. 1956. Studies in the mechanism of the Mannich reaction. I.
The reaction of methylenediamines with 2-methyl-2-nitro-1-propanol.
Jour. Amer. Chem. Soc., vol. 78, pp. 482-484.

CummMIncs, T. F., AND J. R. SHeLron. 1960. Mannich reaction mechanisms.
Jour. Org. Chem., vol. 25, pp. 419-423.

FERNANDEZ, J. E. 1964. A study of the intermediates in the Mannich re-
action of nitroalkanes. Tetrahedron Letters, no. 39, pp. 2889-2893.

FERNANDEZ, J. E., AND J. S. FowLer. 1964. Kinetics of the reaction of 2-
nitropropane with methylene bisamines. A study of the Mannich re-
action. Jour. Org. Chem., vol. 29, pp. 402-407.

FERNANDEZ AND FERREE: Mannich Reaction 19

FERNANDEZ, J. E., J. S. FowLErR, AND S. J. Guaros. 1965. Kinetics of the
reaction of nitroalkanes with methylene bispiperidine. A study of the
Mannich reaction. Jour. Org. Chem., vol. 30, (In Press) August, 1965.

LIEBERMAN, S. V., AND E. C. Wacner. 1949. The course of the Mannich
reaction. Jour. Org. Chem., vol. 14, pp. 1001-1011.

ReIcHERT, B. 1959. Die Mannich Reaktion. Springer-Verlag, Berlin, 195
pp.

Runti, C., anp F. Cortino. 1959. Reaction between fB-naphthol and for-
maldehyde in the absence and in the presence of ammonium salts and
hydroxylamine. Ann. Chim. (Rome), vol. 49, pp. 1472-1491.

Department of Chemistry, University of South Florida, Tampa,
Florida.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

Verification of Monotropa hypopithys in Florida
DanrEL B. Warp

J. K. Smatt (1903) appears to have been the first to record the
Pine-sap, Monotropa hypopithys L. (Ericaceae), for Florida, and
although others have since repeated this southern limit for a pre-
dominantly temperate plant, the probability is high that all such
statements are based upon Small’s original report. No Florida col-
lections of this species are in Florida herbaria, and none were en-
countered in the larger northern herbaria examined by Gupton
(1962).

Such unsupported statements of range are not uncommon in
Small’s works, although as more detailed knowledge accumulates
it is also not uncommon to find that Small’s stated ranges are more
accurate than would be predicted from inspection of his collections
and those available to him. This is the case with Monotropa
hypopithys, as verified by the following collection:

Stem and ovary pinkish-red; plants all in fruit; frequent over
several acres of dry Pinus clausa scrub, east edge of Orange Springs,
Marion County, Florida. Richard H. Spielman, 25 November 1965
(deposited in FLAS; duplicates to FSU and GH).

The plants grew near scattered colonies of the much more com-
mon Monotropa uniflora, or Indian Pipes, but could readily be dis-
tinguished from this better known species by the presence on each
stem of three to nine flowers. Although in an area very familiar
to the collector, they had never before been seen by him, suggest-
ing that there may be a marked annual fluctuation in numbers.

The southernmost collections reported by Gupton for Mono-
tropa hypopithys are Clay County, Alabama (Harper, NY), and
Muscogee County, Georgia (Boykin, NY). The present station, in
the north-central portion of the Florida peninsula, thus represents
a range extension to the southeast of approximately 300 miles.

LITERATURE CITED

Gupron, W. O. 1962. An analysis of the taxonomic criteria as applied to
the genus Monotropa. University of North Carolina, unpublished the-
sis, 140 pp.

SMALL, J. K. 1903. Flora of the southeastern United States. New York,
published by the author, 1370 pp.

Department of Botany, University of Florida, Gainesville, Flor-
ida.
Quart. Jour. Florida Acad. Sci. 29(1) 1966

A Pacific Polychaete in Southeastern United States
Joun L. Taytor

THE marine polychaete worm Poecilochaetus johnsoni Hartman
was originally described from a specimen taken near the mouth of
Mission Bay, southern California (Hartman, 1939), and has not
been previously reported from the Gulf of Mexico or the Atlantic
seaboard (Hartman, 1965). This report records the occurrence of
P. johnsoni in the southeastern United States and presents informa-
tion on the biology of this rarely collected species. Specimens
were collected in Florida at Seahorse Key, Tampa Bay, Naples,
and Biscayne Bay in 1958-65. During the same period, the worm
was also taken in North Carolina at Cape Lookout and in Core and
Bogue Sounds.

EXTERNAL MORPHOLOGY

Specimens of P. johnsoni from southeastern United States cor-
respond with the species description (Hartman, 1939) except for a
greater variability in spinous setae and anal cirri. The original
account reported three, four, or perhaps five falcate spines on
the second and third setigers and described three simple and sub-
equal cirri along the ventral border of the anus, and one on its
middorsal border.

Most specimens collected in the southeast have four falcate
spines on the second setiger and three on the third. Exceptions
are fairly common, however, and some individuals have three to
six spines on either setiger. Anal cirri usually form a ventral series
of three, consisting of one median and two lateral processes. The
midventral cirrus is simple, but the lateral pair may be divided
distally once or several times. Only one specimen has a single
middorsal cirrus; some have two dorsal cirri arising separately or
jointly at a common base; and on others, the dorsal border of
the anus appears plain. These features are evidently labile for
eastern races of the species and do not seem of sufficient meristic
importance to warrant establishment of a distinct taxonomic sep-
aration between southeastern and Pacific forms of the worm.

ECOLOGY

Environmental conditions in areas where P. johnsoni has been
collected are summarized in Table 1. These records show that

2, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

the worm commonly occurs in coastal situations having temperate
or subtropical characteristics, moderate tidal flow, and a substrate
of silty sand.

At most localities the mean annual salinity is above 30 0/oo and
fairly stable, but at Seahorse Key, Florida, and in Bogue and Core
Sounds, North Carolina, the mean value is lower and freshets pe-
riodically create brackish conditions. Although the benthic habit
of P. johnsoni may mitigate the effects of dilute seawater, it seems
likely that the worm has some capacity for osmoregulation or other
physiological adjustment during periods of osmotic stress.

Marine grasses are a conspicuous feature in many areas where
P. johnsoni occurs. The friable dwelling tube has been taken
among roots and rhizomes, as well as in clear areas between
patches of vegetation. In May 1965, 189 specimens were collected
in a bottom sample one-half meter square, at the southern end
of Core Sound, North Carolina, adjacent to Whitehurst Island, in
tidal flats densely vegetated with Zostera marina.

Commensal pinnotherid crabs tentatively identified as juvenile
Pinnixa chaetopterana were found in tubes of P. johnsoni at Sea-
horse Key, Florida, and Whitehurst Island, North Carolina. In
both areas adult P. chaetopterana were collected from tubes of
larger polychaetes. The small tube of P. johnsoni imposes a size
limit on commensal crabs that can be accommodated; the associ-
ation is probably temporary, providing a refuge for young crabs
during early post-larval stages.

Reproduction of P. johnsoni apparently takes place in spring,
summer, and fall when water temperature is about 20 C or above.
Plankton from Tampa Bay contained larval stages of P. johnsoni
in April, May and August (John A. Kelly, Jr., and Alexander Drago-
vich, unpublished). At Cape Lookout, North Carolina, a specimen
taken by Lawrence McCloskey in November had ruptured median
and posterior parapods, presumably damaged during release of
gametes.

In comparison with other species of Poecilochaetidae (Hannerz,
1956), P. johnsoni has been considered most closely related to
Poecilochaetus tropicus Okuda from the Pacific (Hartman, 1939 and
1959). From an ecological standpoint, however, P. johnsoni re-
sembles in many respects the eastern Atlantic species, Poecilochae-
tus serpen Allen, studied on the English coast (Allen, 1904). The
population concentration of P. johnsoni at Whitehurst Island,

Taytor: Pacific Polychaete Worm 23

North Carolina, provides an excellent opportunity for detailed
studies and clarification of the biology of this unusual and little-
known spioniform polychaete.

MATERIAL EXAMINED

Seahorse Key, Florida: 2 specimens (anterior portion only), E.
Lowe Pierce, coll., 1958, U. S. National Museum No. 32616; 8
specimens (anterior portion only), John L. Taylor, coll., September
22, 1960; 1 specimen (anterior portion only), John L. Taylor and
Carl H. Saloman, coll., August 5, 1964.

Tampa Bay, Florida: 2 specimens (median fragments only),
John L. Taylor and Car] H. Saloman, colls., August 10, 1964; 4
specimens (juvenile, meroplankton), Alexander Dragovich and John
Kelly, Jr., colls., April 11, 1961, May 8, 1962, August 16, 1962 (2).

Naples, Florida: 1 specimen (anterior portion only), Charlene
Long, coll., December 1958.

Biscayne Bay, Florida: 1 specimen (anterior portion only), Rob-
ert T. Paine, coll., January 26, 1960.

Cape Lookout, North Carolina: 1 specimen (anterior portion
only), Lawrence McCloskey, coll., November 4, 1963.

Core Sound, North Carolina: 189 specimens (entire specimens
and fragments), John L. Taylor, coll., May 4, 1965.

ACKNOWLEDGMENTS

Many people contributed to this report. Dr. Marian Pettibone,
Division of Marine Worms, U. S. National Museum, identified the
first specimens of P. johnsoni collected on the east coast of the
United States. Dr. Meredith Jones, Division of Marine Worms,
U. S. National Museum, reviewed the original manuscript and for-
warded specimens that he identified from the collections at Naples,
Biscayne Bay, and Cape Lookout. Carl Caloman, Alexander Drag-
ovich, and John Kelly, Jr., Bureau of Commercial Fisheries Biologi-
cal Station, St. Petersburg Beach, Florida, provided specimens and
ecological data from the west coast of Florida. Miss Charlene
Long, Museum of Comparative Zoology, Harvard University, pro-
vided field data for the Naples area. Dr. Robert Paine, Depart-
ment of Zoology, University of Washington, and Durbin Tabb,
Marine Laboratory, University of Miami, supplied ecological data
for Bear Cut in Biscayne Bay. Mrs. Mary Kay and Lawrence Mc-

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

24

wnuIpnysa} DISsD]vY J,
auofyy wniposurihs
niysun dDiayjunjdiq

pyep ON

oe

oP

unulpnysay
DISSD]DY I,

vyep ON

UOT}LJIGIA
jewtxo1g

88-S&

68-CE

8E-EE

SESE

ce-OL

CS-0
osuvy

(00/0)

cs

o8€

oeVE

osVE

Ve

urs

A}UITeS

ve-Ol

Gc-Ol

(D0)

91n}e19duts}
IayeM jo
osuel
jenuuy

T WidVvi

I pues snooreojey
ByeEp ON puvs-AjIS
Avo pur
WS %TL TWyeus
rk pue pues osivop
rG op
Avo pur
WIS %S Toys
c’0 pue pues osirv0p
vyep ON ={;purs-AqIS
(s}oU) adA}
qUIOIIMO JUSWUTPIS
yUSIquIy

wosuyol snjanyoojva0g JO UWOde][09 Jo syutod ye suONTPUOd [RvoIso]OOy

EPMO lel
‘Aeq oudvosig

qny Ieog

eplop yy
‘sopdeny

ey re) ttsl
‘Avg vdurvy,
‘ssegq suyo[

1 O28 (0) A |

‘Avq vduey,
‘sstq sooung

epHolay
‘AOY ISIOYRIS

RIULOFIVD
‘Avg UOISSIJ

A}VIO'T

Pacific Polychaete Worm

TAYLOR:

‘IOGI ‘Yyoraoseiq, ‘FOBT ‘AT[PN pur ‘ouvonury ‘ueuto
1S, ‘FOBT ‘AeA OAPI ® 3SkOD 'S “, ‘O9BT ‘APAING OHOAPOID W% ysvOD 'S “A, ‘TOG “URWUAIeH, ‘GE6T “UeUTIeH,

nyysn Avo pure }Is eUl[OIvD) YWON
piayqupji did Co-OT Og Oe-9 =Gi() YP ‘pues wumnipea{ ‘punosg onsog
evul[oOIeD WON

Avo pure ‘punog 9109

WES %OT ‘TUS ‘purysy

DULLDUL DLIJSOZ, Ce-OT OS 0&-9 I pue pues osreop ysINYouUY MA,

pinIsnqip DUYNIOC
{e100 SUIAT]T UO ~—- PUT[OIVD) _ YION

eyep ON 9e-0& Ge QZ-CI T peysodep IIS ‘ynoyoo'T edep
UOT}L}JIBIA osuery uevoy (oy) (sjoux) adA} AY[VIO'T
[eUlxolg —_— orm pvoddic0} yUsIM9) JUSUTIPIS
(00/0) AWUTTVS ToyeM FO qusIquiy
osuel
jenuuy

wosuyol snyapyso}1920g JO UOTE [OO Jo syuIod ye suUOIpUOd. [ROIsO[OOW ©

(ponuru0s) [| ATAVL

26 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Closkey, Duke University Marine Laboratory, and Hugh Porter,
University of North Carolina Institute of Fisheries Research, gave
notes and records for environmental conditions in Bogue Sound,
Core Sound, and at Cape Lookout. Dr. John Bradshaw, Scripps
Institution of Oceanography, furnished temperature and _ salinity
records for Mission Bay, California, and vicinity.

LITERATURE CITED

ALLEN, E. J. 1904. The anatomy of Poecilochaetus, Claparede. Quart. Jour.
Microscope. Sci., vol. 48, pt. 1, pp. 79-151.

Dracovicu, A. 1961. Relative abundance of plankton off Naples, Florida,
and associated hydrographic data, 1956-57. U. S. Fish Wildl. Serv.,
Spec. Sci. Rep.—Fish. 372, 41 pp.

HANNERZ, L. 1956. Larval development of the polychaete families Spioni-
dae Sars, Disomidae Mesnil, and Poecilochaetidae n. fam. in the Gull-
mar Fjord (Sweden). Zoologiska Bidrag, vol. 31, pp. 1-204.

Hartman, O. 1939. New species of polychaetous annelids from southern
California. Allan Hancock Pacific Expeditions, vol. 7, no. 2, pp. 159-
LO}

—. 1959. Catalogue of the polychaetous annelids of the world. Allan
Hancock Foundation Publications, Occasional Paper, no. 23, Pt. II,
Univ. Southern California Press, Los Angeles, pp. 355-628.

—. 1961. Polychaetous annelids from California. Allan Hancock Pa-
cific Expeditions, vol. 25, 226 pp.

1965. Deep-water benthic polychaetous annelids off New England
to Bermuda and other north Atlantic areas. Allan Hancock Founda-
tion Publications, Occasional Paper, no. 28, Univ. Southern California
Press, Los Angeles, pp. 1-378.

SALOMAN, C. H., J. H. Finucane, anp J. A. KeEtty, Jr. 1964. Hydro-
graphic observations of Tampa Bay, Florida, and adjacent waters, Au-
gust 1961 through December 1962. U. S. Fish Wildl. Serv., Data
Report 4, 6 microfiches, 112 pp.

U. S. Coast AND GEOpETIC SuRvEY. 1960. Surface water temperature and
salinity Atlantic Coast North and South America. U. S. Government
Printing Office, Washington, D. C., 76 pp.

—. 1964. Tidal current tables, Atlantic Coast of North America.
U. S. Government Printing Office, Washington, D. C., 188 pp.

Bureau of Commercial Fisheries Biological Station, St. Peters-
burg Beach, Florida. Contribution No. 24.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

A New Burrowing Barnacle from the Western Atlantic
Harry W. WELLS AND JACK T. TOMLINSON

In the course of an ecological study of sessile marine inverte-
brates of the northeastern Gulf of Mexico, numerous specimens of
an undescribed species of burrowing barnacle have been collected
by the senior author. These specimens were found embedded in
characteristic, easily recognized burrows in dredged shell material,
especially in shells of the turkey wing ark, Arca zebra Swainson,
and the mossy ark, Arca imbricata Bruguicre (= Arca umbonata
Lamarck), and in masses formed of calcareous red algae (Litho-
thamnium and Goniolithon species). Independently, the junior au-
thor had found this species in shells from North Carolina and
Florida and in dead coral from Puerto Rico.

These barnacles are members of the suborder Acrothoracica,
which are recognized by their having a soft mantle without cal-
careous plates, reduced cirri, no abdomen (except appendages in
cyprid stages), and dwarf males. Under the system of classifica-
tion of the Acrothoracica proposed by Berndt (1907), they would
be placed in the genus Kochlorine, which was established by Noll
(1875) for a species he described from Cadiz, on the Atlantic coast
of Spain. Later Noll (1883) described another species of Kochlo-
rine from the Cape of Good Hope, South Africa; and Tomlinson
(1963) has recently described a species from Japan. The descrip-
tion of a fourth species follows. |

Genus Kochlorine Noll 1875

Diagnosis (emended). Mouth cirri biramous, may be weakly
segmented; 3 pairs of terminal cirri; 1 pair of 2-segmented caudal
appendages; lateral bar often present.

Kochlorine floridana, new species

Diagnosis. Kochlorine with 1 pair of short conical spines and
rows of tack-shaped teeth on mantle aperture, and with lateral bar;
attachment process moderately developed.

Type locality. At 8-10 meters depth in the Gulf of Mexico off
Dog Island, Florida, approximately 8 miles southeast of Carrabelle,
Franklin Co., Florida.

28 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Type material. Female holotype (USNM cat. no. 1138221) with
one attached male, collected 16 May 1963, and some paratypes
(USNM cat. nos. 113222, 113223, 113224, and 118225), deposited
in the Division of Crustacea, U. S. National Museum. Other para-
types and additional material in the author's collections.

Additional material. ¥YLorma: SE of Pensacola, from the bry-
ozoan Hippoporidra edax (Busk), California Academy of Sciences,
R. Kiwala, coll.; St. Andrews Bay entrance, near Panama City,
from Murex fulvescens Sowerby, G. Bertrand, coll.; off Cape St.
George (Gulf of Mexico) from Aequipecten gibbus (Linne), H. W.
and M. J. Wells, colls.; Fernandina Beach from Arca imbricata,
A. Ross, coll.; off St. Augustine, from Murex fulvescens and other
shell material, H. W. and M. J. Wells, colls. cEorcta: off Sapelo
Island, from Arca imbricata and other shell material, J. Kraeuter,
coll. sourH CAROLINA: off Charleston, M/V Silver Bay Sta. 1389,
from Arca imbricata and calcareous algae; off Georgetown, M/V
Silver Bay Stas. 1360, 1392, from Arca zebra. NORTH CAROLINA:
Shackleford Banks, from Murex fulvescens, H. W. and M. J. Wells,
colls., V. Zullo, coll.; Back Sound, from shelly bottom, H. W.
Wells, coll.; Cape Lookout jetty, A. Ross, coll.; Diamond Shoals,
from Murex fulvescens and Cassis madagascariensis (Lamarck,
H. W. Wells, coll. In addition, the “Alcippe lampas” recorded for
the Beaufort area by Wells (1961) is in reality K. floridana. PUERTO
RICO: from dead coral fragments, W. A. Newman, coll. Other ma-
terial found in Murex microphyllus Lamarck from Tuléar, Mada-
gascar (by courtesy of the Institut Royal des Sciences Naturelles de
Belgique), is regarded by the junior author as being identical.

DESCRIPTION

Female. Body compressed, sac-shaped in lateral view (Fig. 1),
and slightly asymmetrical with the posterior end twisted slightly
to the side. Holotype length, 2.10 mm; width, 1.25 mm; thick-
ness, about 0.75 mm. In a series of 18 additional females, 0.53
to 3.33 mm long, body width 0.518 to 0.702 times body length,
with the smallest specimens proportionately narrowest.

Mantle surface with irregularly scattered small bi- or tricuspate
hooks, slightly more numerous in lateral patches near site of at-
tachment of males; a series of Jarger tack-shaped teeth around
aperture margin. Mantle containing superficial longitudinal mus-
cle bands and deeper circular bands; suffused with red pigment

WELLS AND TOMLINSON: New Burrowing Barnacle 29

Fig. 1. Kochlorine floridana, n. sp., female, lateral view, with attached
male. Abbreviations: AP - attachment process, C - mantale cavity, CA - cau-
dal appendage, DG - digestive glands, E - esophagus, EM - embryos, G - gan-.
glion, K-knob, LB - lateral bar, M-male, MC-mouth cirrus, MP - mouth
parts, PH - pharynx, S - stomach, SP - spine.

30 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

near aperture. Aperture gently arched, bearing a pair of short,
blunt spines, usually studded with strong teeth. Apertural length
0.484 to 0.767 times body width, proportionately longer in smallest
specimens. Apertural rim and spines ornamented in a characteristic
pattern of three rows of heavy teeth. Teeth in outer row bifid, in
inner two rows broad and tack-shaped, with single acute dorsal
and ventral spines and lobate lateral margins to the crown, thus
resembling a flower. Comb-like row of spinules guarding aper-
tural inner edge at ventral end; rounded spinous knob at dorsal
end. Knob of large specimens prominent, bearing up to ten ven-
trally curved spines; spines and knob reduced or absent in juvenile
specimens.

Attachment process moderately developed, more or less pro-
truding, mid-dorsal and posterior to knob. Body normally lodged
in surrounding shell by projection of attachment process into a
small depression in burrow wall, disk posterior to attachment
process, attached to walls of burrows in mollusk shells. Pads of
shed cuticle often retained in the form of a cone at disk site in
specimens removed from calcareous algae (Fig. 3a). One or two
pairs of dissociated antennules occasionally attached to this cuti-
cle, from former attachment of males at this site. A lateral bar,
finely reticulated, on each side extending posteriorly from aper-
tural margin. Distinct muscle bands extending from attachment
process along dorsal body wall to posterior end, from attachment
process to point posterior to stomach, and from attachment proc-
ess to mouth region. Another muscle band extending from mouth
region to point posterior to stomach.

Body enclosed within mantle, tapering and, in the contracted
state, curving dorsally; posterior end bending sharply ventrally;
two pairs of small, fleshy ventral protuberances in midregion.
Head rounded, studded with one or more rows of fine hairs. Di-
gestive tract: muscular pharynx extending dorsally from mouth;
long esophagus extending posteriorly; broad stomach; pair of small,
round digestive glands attached to stomach at juncture with esoph-
agus; anus situated at end of short intestine, between terminal
ClITi.

Paired mouthparts of mandibles with palps and two pairs of
maxillae. Mandible (Fig. 2a) with three simple teeth of decreas-
ing size, a small bifid tooth at the tip of its cutting edge, numerous
spines, and numerous rows of bristles; apodeme extension short,

WELLS AND TomMLINsON: New Burrowing Barnacle 31

Fig. 2. Kochlorine floridana, n. sp., female: a. mandible, b. first: maxilla,
c. second maxillae, d. mouth cirrus, e. caudal appendage and basal segments
of third terminal cirrus. (a-c to same scale; d-e to same scale.)

32 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

not hooked. Palp edged by row of hairs, with a tapering end ex-
tending beyond mandible tip, closely appressed to labrum. First
maxilla (Fig. 2b) bilobate, with many bristles; two strong, curved
teeth at tip of superior lobe; long curved or hooked apodeme at-
tached at base. Second maxilla (Fig. 2c) relatively large and soft,
with long flexible bristles at tip and along edges.

Mouth cirri (Fig. 2d) posterior to second maxillae, capable of
extending anteriorly well beyond apertural margin; with two short
rami attached to a long 2-segmented pedicle. Anterior ramus
longer, 4-segmented; posterior ramus indistinctly 3- or 4-segment-
ed; both provided with numerous flexible, usually feathery bris-
tles.

Three pairs of biramous terminal cirri and one pair of uni-
ramous caudal appendages attached to posterior end of body.
Rami of terminal cirri subequal, with numerous, distinctly seg-
mented articles with long setae along the inner side of their curva-
ture. Setal arrangement (Fig. 3d) characteristic: a distal pair of
long setae (usually with a minute median seta) and a middle pair
of short setae on the inner side of each segment, with one or two
setae of intermediate size situated at intervals on opposite (outer)
face of cirrus. Intermediate sized outer setae spaced from 1 to 5
segments apart, with mean number of intervening segments being
1.69 in cirrus 2 (for 1 specimen), 2.90 in cirrus 3, anderen
rus 4. Mean number of segments (for 5 specimens) in the anterior
and posterior rami respectively: 27.0 and 37.0 for cirrus 2, 40.5
and 45.6 for cirrus 3, and 42.0 and 44.8 for cirrus 4. Pedicles edged
with bristles, 2-segmented with proximal segment much the long-
er, and with oblique sutures between segments. Caudal append-
ages (Fig. 2e) 2-segmented, their tips not reaching the first articu-
lation of cirral pedicles; terminal segment with three feathery
setae and a minute spine at tip; proximal segment minutely ridged
ventrally, with one dorsal seta.

Although all specimens 0.962 mm or more in length show the
cirral complement described above, specimens smaller than 0.8
mm exhibit an incomplete cirral complement. Two specimens,
0.611 and 0.767 mm long, bear two pairs of cirri and a pair of
short, stout cirral buds anterior to the cirri, as well as caudal ap-
pendages. Two other specimens, 0.572 and 0.598 mm long, possess
only two pairs of cirri and the caudal appendages. The smallest
specimen, 0.533 mm long, bears only a single pair of well devel-

WELLS AND TOMLINSON: New Burrowing Barnacle 33

oped cirri and a pair of short cirral buds, as well as the more
slender caudal appendages.

The excavation produced in shell material is smooth-walled and
somewhat variable, corresponding to the shape and size of the
inhabitants body. The excavation slants obliquely in a dorsal
direction and expands to its greatest width in the lower half.
This lower part of the burrow is usually twisted slightly to one
side. Typically, a small depression occurs immediately below the
dorsal lip, in a position corresponding to the attachment process
of the barnacle. The orifice (Fig. 3e) is a narrow slit in surface
view, broader at the ventral end, and is considerably shorter than
the maximum height of the burrow. Particularly at the narrow,
dorsal end, a delicate ridge composed of opaque calcium carbonate
may be built up around the orifice edge. In certain mollusk shells
(e.g. Atrina species), these deposits around the orifice and the sim-
ilar opaque lining of the excavation stand out in marked contrast
to the translucent crystalline structure of the surrounding. shell.

01mm

Fig. 3. Kochlorine floridana, n. sp.: a. lateral view of female removed
from calcareous algae by acid treatment, showing cone of shed cuticle, b.
male, c. cyprid larva, d. setation of segments of terminal cirrus, e. orifices of
burrows produced by large and small individuals. (b-c to same scale.)

34 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Male. Body lobate (Fig. 3b), 0.41 to 0.46 mm long, 0.11 to 0.14
mm wide. One lobe more cylindrical and bluntly rounded than
the others, with a small pigment spot, projecting into a depression
in mantle of female. Attachment by a pair of stout antennules ap-
proximately 0.11 mm long connected to body by a slender neck
region 0.07 to 0.10 mm long (Fig. 3b). Penis tapering, 0.985 mm
long by 0.020 to 0.015 mm wide, extending from posterior end of
one male.

These dwarf males attach to the wall of a burrow occupied
by a female or to the mantle of the female on one or both sides.
Supernumerary males are common, with as many as five males
associated with a single female. Additional pairs of male anten-
nules occasionally remain attached to the cuticular material. Thus,
each female may be served by a number of relatively short-lived
males.

Cyprid larva. Body elongate, tapered at both ends (Fig. 3c),
0.55 mm long and 0.19 mm wide, with a pair of prominent an-
tennules near middle of body, and a pair of pigmented eyespots.

Remarks. This species is placed in the genus Kochlorine Noll
because it possesses three pairs of terminal cirri and a pair of cau-
dal appendages in the adult female. These characters distinguish
Kochlorine from Balanodytes Utinomi, Lithoglyptes Aurivillius,
and Berndtia Utinomi, related acrothoracican genera known to bur-
row into calcareous shells and coral. Balanodytes has four pairs
of terminal cirri (Utinomi, 1949b); Lithoglyptes has four pairs of
terminal cirri and caudal appendages (Tomlinson and Newman,
1960); and Berndtia has five pairs of terminal cirri (Utinomi, 1949a).
From other species described in the genus Kochlorine, K. flori-
dana differs in the shape or number of projections on the aperture
of the female, having a single pair of short conical spines. Kochlo-
rine bihamata Noll (1883) from the Cape of Good Hope, South
Africa, and K. habei Tomlinson (1963) from Japan bear a pair of
hook-like projections on the apertural margin. Kochlorine hamata
Noll (1875) from Spain bears a single hook-like projection on the
aperture as well as a pair of stout conical projections similar to
those found on K. floridana. Kochlorine floridana also differs from
K. habei in possessing many large tack-shaped teeth around the
aperture, and from K. hamata and bihamata in possessing a more
or less prominent attachment process, corresponding to the attach-

WELLS AND Tomurnson: New Burrowing Barnacle 35

ment process described for Balanodytes taiwanus Utinomi (1949b)
from Formosa.

The burrows produced by Kochlorine floridana show some re-
semblance to acrothoracican barnacle burrows described by Ross
(1965) from Florida Miocene material. The discovery of Kochlo-
rine floridana in western Atlantic waters adds this species to the
Recent Caribbean and western Atlantic fauna considered by Ross.
However, an attempt to relate the excavations in Miocene material
to this species does not seem justified by the evidence at hand.

Kochlorine floridana is named in allusion to the collection of
the type specimens from Florida waters.

DISCUSSION

At the type locality, K. floridana is an abundant inhabitant of
shell material. Its burrows occur chiefly in shells of the turkey
wing ark, Arca zebra, whether or not the mollusk is alive. The
burrows are most common in the oldest part of the shell, i.e. in
the umbonal region. At the type locality, Kochlorine floridana
also occurs in masses of calcareous algae, calcareous serpulid poly-
chaete tubes, compartments of the acorn barnacle (Balanus calidus
Pilsbry), dead coral, and shells of the mossy ark (Arca imbricata),
the wing oyster (Pteria colymbus Roding), the rock oyster (Chama
macerophylla Gmelin), and pen shells of the genus Atrina.

By its burrowing activity, K. floridana contributes to the erosion
and destruction of shells that accumulate in nearshore waters. In
these waters, it shares its ecological role with boring sponges (Cli-
ona species) and several species of burrowing pelecypods and poly-
chaetes, which are also abundant in calcareous material at the type
locality.

The classical interpretation of the boring mechanism in boring
barnacles relates their penetrating ability solely to mechanical
erosion by teeth and hooks of the mantle as the mantle is moved
by muscular contraction (for Trypetesa, Darwin, 1854; for Balano-
dytes, Utinomi, 1949b; for Lithotrya, Darwin, 1854, and Yonge,
1963). The deposits of calcareous material found at the narrow,
dorsal end of Kochlorine burrows fit the description of similar de-
posits in the burrow of Trypetesa lampas (Hancock) by Darwin
(1854), who attributed this deposition to a chemical phenomenon
rather than to biological secretion. An opaque calcium carbonate

36 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

layer lines the dorsal wall of the Kochlorine burrow and forms a
ridge at the dorsal end of the orifice. To what degree the barnacle
participates in shaping these deposits is not known.

ACKNOWLEDGMENTS

We wish to thank Mary Jane Wells for her preparation of the
figures and her many contributions to this study. This work has
been supported in part by grants from the National Science Foun-
dation (GB-819) and from the Florida State University Research
Council to H. W. Wells, and in part by a grant from the National
Institutes of Health (Public Health Research Grant GM 09953-04}
to J. T. Tomlinson. In addition, we wish to thank those persons
who contributed specimens to our study, and the Bureau of Com-
mercial Fisheries for the specimens collected by the M/V Silver
Bay.

LITERATURE CITED

BERNDT, W. 1907. Uber das System der Acrothcracica. Arch. Naturg., vol.
73, no. 1, pp. 287-289.

Darwin, C. 1854. A monograph on the subclass Cirripedia. Vol. 2. Ray
Society, London, 684 pp., 30 pls.

Nou, F. C. 1875. Kochlorine hamata N., ein bohrendes Cirriped. Z. Wiss.
Zool. vols 20;,pp.) LI4=I28. pl

——. 1883. Zur Verbreitung von Kochlorine N. Zool. Anz., vol. 6, pp.
471-472.

Ross, A. 1965. Acrothoracican barnacle burrows from the Florida Miocene.
Crustaceana, vol. 9, pp. 317-318.

TomMLuinson, J. T. 1963. Two new acrothoracican cirripeds from Japan.
Publ. Seto Mar. Biol. Lab., vol. 11, no. 2, pp. 263-280.

TOMLINSON, J. T., AnD W. A. NEwMaANn. 1960. Lithoglyptes spinatus, a
burrowing barnacle from Jamaica. Proc. U. S. Nat. Mus., vol. 112, pp.
517-526.

Utinomi, H. 1949a. A new remarkable coral-boring acrothoracican cirriped.
Mem. Coll. Sci., Univ. Kyoto, ser. B, vol. 19, no. 3, pp. 87-98.

——. 1949b. On another form of Acrothoracica, newly found from For-
mosa. Mem. Coll. Sci., Univ. Kyoto, ser. B, vol. 19, no. 3, pp. 95-
100.

Wetts, H. W. 1961. The fauna of oyster beds, with special reference to
the salinity factor. Ecol. Monogr., vol. 31, pp. 239-266.

WELLS AND TOMLINSON: New Burrowing Barnacle 37

YoncE, C. M. 1963. Rock boring organisms. In Mechanisms of hard tissue
destruction (R. F. Sognnaes, ed.), pp. 1-24. Amer. Assoc. Adv. Sci.,
& /, DPE
Washington, D. C.

Department of Biological Sciences, Florida State University,

Tallahassee, Florida; and Biology Department, San Francisco State
College, San Francisco, California.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

Acetes Shrimp on the Florida East Coast
Epwin A. JOYCE, JR.

Durinc spring when commercial shrimp are known to be spawn-
ing, shrimp boat operators are often much concerned over the
many small shrimp of less than an inch in length which become
entangled in the meshes of the nets. These shrimp are transparent
when first brought on board, but as they dry out they become
opaque and drop from the nets in large numbers. Concern is
sometimes so great that the shrimpers wish to have the grounds
closed at this time of year to avoid the unnecessary killing of
“millions of baby white shrimp.”

Recently, however, it was learned that these are not the young
of the commercial shrimp but are instead adult members of the
family Sergestidae (Joyce, 1965). To verify this, specimens were
sent to Dr. L. B. Holthuis of the Rijksmuseum van Natuurlijke
Historie, Leiden, Netherlands, who kindly identified them as
Acetes americanus carolinae Hansen.

This sergestid occurs from North Carolina throughout the Gulf
of Mexico and south to Brazil. There are two subspecies, of
which A. a. carolinae is the northern form (Holthuis, 1959). Con-
sequently, the appearance of these small adult shrimp should in
no way hinder or cause concern to shrimping operations on the
east coast of Florida.

LITERATURE CITED

Ho.truuis, L. B. 1959. Crustacea Decapoda of Suriname (Dutch Guiana).
Zool. Verh. Leiden, ao. 44, pp. 1-296.

Joyce, Epwin A., Jr. 1965. The commercial shrimps of the northeast coast
of Florida. Florida Board of Conservation, Prof. Paper Ser., no. 6,
pp. 1-224.

Florida Board of Conservation Marine Laboratory, St. Peters-
burg, Florida. Contribution No. 91.

Quart. Jour. Florida Acad. Sci. 29(1) 1966:

A New Frog of the Genus Hyla from British Guiana
COLEMAN J. GoIN

A sMALL, brilliantly marked tree frog from British Guiana has
been called to my attention by Miss A. G. C. Grandison, for whom
this attractive little species is named.

Hyla grandisonae, sp. nov.

Type. British Museum (Nat. Hist.) 1938.10.3.25, collected from
a leaf of a shrub in the forest at Mazaruni, British Guiana, by
J. Smart.

Diagnosis, A small Hyla with reduced webbing between the
fingers, a well developed patagium, and brilliant white dorsal sur-
faces of the upper arms. Perhaps most closely related to Hyla
rondoniae Bokermann (1963) and Hyla bokermanni Goin (1960),
but readily distinguished from both of these by the more strongly
developed vomerine teeth, the larger choanae, the bright white
dorsal surfaces of the upper arms, and the lack of oval white
spots on the anterior faces of the thighs.

Description of type. Vomerine teeth in two rather long, heavy,
series, lying close together between the posterior halves of the
rather large rounded choanae; tongue three-fourths as wide as
mouth-opening, wider than long, its posterior border fused and
nearly straight. Snout short, rounded when viewed from above,
markedly truncate in profile, the upper jaw extending slightly be-
yond lower; nostrils more lateral than superior, considerably pro-
jecting, their distance from end of snout about one-third that
from eye, separated from each other by an interval nearly equal
to their distance from eye. Canthus rostralis well defined, curved;
loreal region concave and nearly vertical, the upper lip flaring out
slightly below it. Eye large, very prominent, its diameter equal to
its distance from nostril; palpebral membrane not reticulate; inter-
orbital distance about equal to width of upper eyelid, which is
relatively wide and about equal to distance between nostrils.
Tympanum very distinct, about three-fifth the diameter of eye,
separated from eye by a distance equal to about one-half its own
diameter. Fingers webbed at base, fourth a disk’s length longer
than second, just reaching to disk of third, which covers about two-
thirds the tympanic area; no projecting rudiment of a pollex; no

40 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ulnar ridge; toes more than one-half webbed, the web on fourth
toe reaching the base of the penultimate phalanx, and that on
third and fifth toes reaching the distal end of their penultimate
phalanges; third and fifth toes subequal, disk of fourth covering
about one-third the tympanic area; a distinct oval inner, but no
distinct outer, metatarsal tubercle; no tarsal ridge; no dermal ap-
pendage on heel. Body not elongate, in post-axillary region dis-
tinctly narrower than greatest width of head; when hind leg is
adpressed, heel reaches almost to nostril; when limbs are laid along
the side, knee and elbow considerably overlap; when hind legs
are bent at right angles to body, heels overlap slightly. A well
developed patagium extends from the back of the upper arm to
the side of the body. Skin of upper parts smooth; a distinct gland-
ular ridge passes above tympanum where it then turns downward
and terminates; skin of throat and chest smooth, that of belly and
lower surface of thigh uniformly and very finely granular; no
traces of a skinfold across chest; adult male vocal sac apparently
internal. Skin of head not coossified with skull, roof of skull not
exostosed.,

Dimensions. Head and body 20.8; head length 9.4; head width
8.3; femur 9.8; tibia 10.4; heel-to-toe 11.0 mm.

Color in alcohol. A very strikingly marked little frog. The
dorsal ground color is dark brown with the under surfaces of the
chin, throat, and body gray. There is a rather distinct rounded
white spot on the occipital region, and the dorsal surfaces of the
upper arms are clear white. The patagium, while not white like
the dorsal surfaces of the upper arms, is unpigmented. The left
heel has been injured, but there is a short white line on the right
Ince

Discussion. This species is certainly related to the group of
small, dark, brightly patterned South American hylas, of which
H. parviceps Boulenger (1882) seems to be the best known mem-
ber. H. grandisonae seems structurally most like H. rondoniae
and H. bokermanni but it differs from these in having slightly
larger legs, a larger tympanum, heavier rows of vomerine teeth,
and larger choanae. It differs from all these in dorsal pattern and
in lacking oval white spots on the anterior faces of the thighs. From
parviceps it differs further in having a well developed patagium,
which parviceps lacks.

If, as I suspect, the development of a patagium is more indica-

Gorm: New Guianan Frog Al

Fig. 1. Dorsal view, side
and foot of the type of Hyla grandisonae. Actual snout to vent length 20.8
mm.

of head, roof of mouth, and underside of hand

42 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

tive of relationships than is pattern, we would find H. rondoniae,
bokermanni, and grandisonae most closely related, with H. parvi-
ceps a more distantly related form. On the other hand H. rondo-
niae, bokermanni, and parviceps each have oval white spots on
the anterior faces of their thighs, but the thighs of H. grandisonae
are uniformly dark anteriorly.

Acknowledgments. J wish to thank Miss A. G. C. Grandison
for the privilege of describing this species; Mr. Paul Laessle for
the fine figure of the type specimen; and the National Science
Foundation, who have supported my past work on South American
tree frogs, for grant GB-3644 which made possible my work dur-
ing the summer of 1965.

LITERATURE CITED

BOKERMANN, W. C. A. 1963. Duas novas espécies de “Hyla” de Rondonia,
Brasil (Amphibia, Salientia). Rev. Brasil. Biol., vol. 23, pp. 247-250,
6 figs.

BouLENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata
in the collection of the British Museum. Ed. 2, pp. xvi + 503, 30
pls., many text figs.

Gorn, C. J. 1960. Description of a new frog of the genus Hyla from north-
western Brazil. Ann. Mag. Nat. Hist., ser. 13, vol. 2, pp. 721-724,
Dec., 1959, 1 fig.

Department of Zoology, University of Florida, Gainesville,
Florida.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

A Second Specimen of Ophisaurus ceroni
J. Atan HoL_Man

The glass lizard Ophisaurus ceroni was described on a single
individual with a partially broken tail, found dead on the road,
in coastal dune-scrub within the city limits of Veracruz, Veracruz,
Mexico (Holman, 1964). The capture of another O. ceroni is thus
of interest, as it represents the first Ophisaurus taken alive and un-
damaged in Mexico and the second known specimen of O. ceroni

(Figs. 1-2).

Fig. 1. Ophisaurus ceroni Holman. Photograph from life.

This topotype, Museum of Natural History Illinois State Uni-
versity No. 391, was taken in coastal dune-scrub (Fig. 3) at Vera-
cruz, August 31, 1965, by J. Alan Holman and Donna Rae Holman.

DESCRIPTION

Snout-vent length 181 mm; tail length 335 mm; snout-vent/tail
.040; length of regenerated portion of tail 43 mm; head width
10.8 mm; eye diameter 3.5 mm; dorsal scales in 14 longitudinal
series; scales around parietal 7-7; upper labials 11-11; preoculars

Ad QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

3-3; postnasals 2-2; scales around tail 18; scales along lateral fold
101.

Fig. 2. Head of Ophisaurus ceroni Holman. Photograph from life.

Frontonasal divided into anterior and posterior frontonasal;
labials separated from orbit by lorilabials and suboculars; pre-
frontals separated by a posterior extension of the frontonasal;
upper postnasal in contact with supracanthal row as well as with
anteriormost canthal; anterior frontonasal separates postinterna-
sals; five supraoculars; canthals extending to just anterior to mid-
dle of eye; frontal broad posteriorly, pointed anteriorly, anterior
end separating prefrontals; interparietal broad anteriorly, tapering
to a point posteriorly; occiptal narrower than interparietal at its
greatest width; frontoparietal in contact with third and fourth
supraoculars; first and second upper labials in contact with nasal.

Body broader than high; dorsal scales keeled, ventrals smooth
and flat. Ear opening oval, larger than round nostril. Dorsal
ground color grayish-brown, interrupted by three distinct dark
stripes, two lateral and one mid-dorsal. Each lateral dark stripe
divided by a light line and occupying parts of two scale rows; mid-
dorsal stripe divided by very thin light line and occupying ad-
jacent halves of two scale rows; discrete white spots lacking on
both back and sides; no stripes or other dark pigmentation below
lateral fold. Head light grayish-brown; both top and sides of
head speckled with dark spots; vertical bars lacking on neck.

Houtman: Mexican Glass Lizard AD5

Fig. 3. Type locality of Ophisaurus ceroni Holman. Coastal dune-scrub
at Veracruz, Veracruz, Mexico, September 3, 1965.

DISCUSSION

The specimen has all the diagnostic characters that separate
O. ceroni from other species of the genus. The few minor differ-
ences between the second specimen and the holotype may be
attributed to individual variation. These differences are as fol-
lows: (1) prefrontals separated by a posterior extension of fronto-
nasal (prefrontals in broad contact in holotype); (2) occipital nar-
rower than interparietal at its greatest width (occipital about as
broad as interparietal at its greatest width in holotype); (3) lateral
and mid-dorsal stripe relatively broad with each stripe divided by
a thin light line (lateral and mid-dorsal stripe somewhat narrower,
not separated by thin light lines in holotype); (4) vertical neck bars
absent (four quite indistinct, vertical bars on neck of holotype).

The new lizard was caught at 9:00 am as it crawled in a
grassy section of coastal dune-scrub only a few feet from where
the holotype was collected. In captivity, the lizard spends much
time partially or fully buried in the sand. The reptile emerges

46 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

occasionally and crawls about with very deliberate motions. Dur-
ing such times it will usually take cockroaches (Nauphota cinerea)
from the end of a forceps. On one occasion it ate 10 roaches with-
in a few minutes, but it has never chased the insects. Once the
lizard attacked a large, sluggish grasshopper that was placed in
the terrarium, but after chewing on the insect for more than half
an hour, the lizard declined to swallow it. Newly born garter
snakes (Thamnophis elegans vagrans) were offered, but were not
attacked by the glass lizard. Carr (1940) states that Florida Ophi-
saurus eats small snakes and lizards.

McConkey (1954) recognized three New World Ophisaurus
species in his comprehensive study of the genus. Since two addi-
tional species have been described from Mexico (see McConkey,
1955; Holman, 1964), I append the following key to the known
New World species of Ophisaurus.

Key tro NEw Wor.up SPECIES OF OPHISAURUS

i Scales along: lateral fold97 or less O. compressus
Scales alone lateral fold’ 98 or more 2 eee Y
2. Dark stripes or dark pigmentation below lateral fold. —_ O. attenuatus
No dark strikes or dark pigmentation below lateral fold. 3
3. Mid-dorsal dark stripe distinct throughout; frontonasal divided;
white spots on dorsum absent. 2.) O. ceroni
Mid-dorsal stripe indistinct or absent; frontonasal usually undivided;
white spots on dorsum present... ee eee 4
4. Distinct vertical white neck bars present.....__.__- =. O. ventralis
Vertical white: neck bans) absent sete aes eee O. incomptus

LITERATURE CITED

Carr, A. F. 1940. A contribution to the herpetology of Florida. Univ.
Florida Publ., Biol. Sci. Ser., vol. 3, no. }, pp. 1-118.

Hotman, J. A. 1964. A new glass lizard from Veracruz, Mexico. Quart.
Jour. Florida Acad. Sci., vol. 27, no. 4, pp. 311-815.

McConkey, E. H. 1954. <A systematic study of North American lizards of
the genus Ophisaurus. Amer. Midland Nat., vol. 51, no. 1, pp. 133-
WAL.

——. 1955. A new lizard of the genus Ophisaurus from Mexico. Nat.
Hist. Misc., Chicago Acad. Sci., no. 145, pp.. 1-2.

Department of Biological Sciences, Illinois State University,
Normal, Illinois 61761.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

An Extinct Lizard of the Genus Leiocephalus from Jamaica
RicHARD ETHERIDGE

Tue iguanid lizard genus Leiocephalus is now confined to the
West Indies where it occurs on almost all islands except Jamaica
and those islands to the south and east of Hispaniola. An extinct
species has been described from the Late Pleistocene of Barbuda,
and another form apparently became extinct on Martinique in this
century (Etheridge, 1964a). The former existence of these two
species led me to suggest that Leiocephalus might have occurred
throughout the Lesser Antillies and on Puerto Rico in the not-too-
remote past. An extinct species is also known from the Late Pleis-
tocene of Hispaniola (Etheridge, 1965) and from the Early Miocene
of Florida (Estes, 1963).

The discovery of an extinct species of Leiocephalus in several
Jamaican cave deposits indicates that the range of the genus dur-
ing the Late Pleistocene may actually have included virtually all
islands of the West Indies.

JAMAICAN LOCALITIES

Dairy Cave. The best known fossil deposits in Jamaica are
found in Dairy Cave (Fig. 1), about 2.5 km west of Dry Harbour
(now known as Discovery Bay) in St. Anns Parish; 18°27’25” N,
77° 22/38” W. Koopman and Williams (1951) and Hecht (1951)
have given general descriptions of the cave. Some fossils of Leio-
cephalus were collected here by Dr. Harold E. Anthony in 1919
and 1920. The probable site of Anthony's collection has been de-
termined by Dr. Walter Auffenberg at about 200 feet west south-
west of the main entrance. It is designated in Auffenberg’s field
notes for 1958 and on the map (Fig. 1) as Locality E.

In 1958 Dr. Auffenberg obtained additional fossils of Leioceph-
alus from two other localities within Dairy Cave. One of these,
designated Locality G, is about 25 feet south southwest of An-
thony’s site. The bones were recovered from a small amount of
matrix taken from two solution holes in the wall, about 12 feet
above the cave floor. The second site, Locality D, was discovered
in a roughly circular chamber about 15 feet across and 10 feet
high, near the eastern end of the cave. According to Auffenberg’s
field notes much of the matrix has been removed in phosphate min-
ing. The original floor of flow stone is about three feet below the

48 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

limestone floor, and is provided with pillars of travertine connect-
ing the old floor with the ceiling. Below this is a series of beds,
more or less consolidated near the top, more clayey near the bot-
tom. About four feet above the original floor is a layer of shell.
Below this the shell layer gradually gives way to layers of dark
reddish clay. The fossils of Leiocephalus, together with those of
other lizards and mammals, from this locality are from the lower
layers of clay. In his field notes Auffenberg suggests the likelihood
that this layer was contemporaneous with Anthonyss “lizard layer’.

main
entrance

vy
Po cee

ee a a
He

J

Fig. 1. Map of Dairy Cave, St. Anns Parish, Jamaica. Prepared from a
map originally drawn and surveyed by the Jamaica Geological Survey. En-
trances to the cave are indicated by arrows. Localities from which fossils
have been obtained are indicated by crosses and are designated A through G.

Montego Bay Airport Cave. This site was discovered by Auf-
fenberg on June 28, 1957, near the west end of the air strip at
Montego Bay, St. James Parish; 18°30’22” N, 77°5411” W. Ac
cording to his field notes the cave is fairly large but much of it
has been dug out in mining for phosphate. In one of the large
chambers at the eastern edge of the main entrance to the cave he
located a bone-bearing deposit below the former travertine floor.
The cave earth there is semiconsolidated, reddish in color and
contained an abundance of broken shells and bone. The bone
is yellowish in color and chalky in consistency. The verk dark red
color of the beds led him to believe that these deposits are very

ETHERIDGE: Extinct Jamaican Lizard 49

old, with no possibility of recent mixture as seems to be the case
for some fossil lizards in Dairy Cave (Auftenberg, field notes, 1957).
The fossil lizards were collected by Auffenberg during this visit,
and again in June 1958.

Portland Ridge Caves. Several fossiliferous caves have been
explored in Portland Ridge, Clarendon Parish. One of these, des-
ignated Portland Cave I by Auffenberg, was discovered by Dr.
H. E. Anthony in 1919 or 1920, and has been described by Koop-
man and Williams (1951). The fossil Leiocephalus from this cave
were obtained by Dr. Auffenberg on July 7, 1957. According to
his field notes of that year the fossils were taken at Locality E,
near the end of the left, i.e., south, passage. Three beds may be
distinguished at this locality: Bed 3, about 2 feet deep from
the surface, is composed of fine soil, grayish at the surface and
grading to reddish at its base; Bed 2 lies below this, 3 to 8 inches
deep, comprised of pebbly, red, slightly consolidated earth, con-
taining many bones, most of which are of white color; and Bed 1,
about 8 inches deep, whitish and extending to the limestone, con-
taining bones that are blackish. The bones of Leiocephalus were
taken from Bed 2.

A second cave in Portland Ridge, designated Portland Cave
III, is located about 1000 feet southwest of Portland Cave I.
Fossil Leiocephalus were collected here by Dr. Ernest E. Williams
in 1953. According to Auffenberg’s field notes of 1957 the cave
is small and provided with two openings, one at the north end and
one at the south end. To the east it is connected with a very large
and deep cave on a lower level. In a test pit Auffenberg located
a bone-bearing layer about 4 feet down from the surface, just
above a layer of white earth. The Leiocephalus bones collected
by Williams near the south entrance may have come from this
layer.

Leiocephalus jamaicensis new species

Holotype. A left dentary, No. 2311 in the vertebrate paleon-
tology collections of the American Museum of Natural History.
Collected by H. E. Anthony in 1919 or 1920.

Type Locality. Dairy Cave, about 200 feet west southwest
of the main entrance (Locality E), 2.6 km west of Dry Harbour,
Seams Parish, Jamaica; 18°27°25” North Latitude, 77°22’38”
West Longitude. Probably Late Pleistocene.

50 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Referred Specimens. (AMNH = American Museum of Natural
History; UF = University of Florida, the Florida State Museum).
Dairy Cave, Loc. E: maxillae AMNH 2312-3, parietal AMNH 23814,
caudal vertebrae AMNH 2315. Dairy Cave, Loc. D: dentaries UF
8503-4. Dairy Cave, Loc. G: dentary UF 8502. Montego Bay
Airport Cave: dentary UF 8505, parietal UF 8506, frontal UF
8508, pterygoid UF 8509, caudal vertebrae UF 8510. Portland
Cave I, Loc. E: dentaries UF 8511-2, maxilla UF 8513, pterygoid
UF 8514, body vertebrae UF 8516 (2), caudal vertebrae UF 8518
(3). Portland Cave III: dentaries UF 8489-90, maxillae UF 8491-3,
articular-surangular UF 8494, premaxillae 8495 (2), frontal UF
8496, caudal vertebrae UF 8497 (5), 8500, body vertebrae UF
8501 (2).

Diagnosis. A large species of Leiocephalus apparently allied
to L. melanochloris, L. screibersi and L. inaguae, from which it
differs only in having attained a greater maximum size. Its max-
imum snout-vent length was at least 130 mm, as great as the
largest living form, L. carinatus microcyon, and may have been
as large as 170 mm. The maximum recorded snout-vent lengths
of melanochloris, schreibersi and inaguae are 108 mm, 90 mm and
73 mm respectively.

Leiocephalus jamaicensis also differs from L. carinatus, L. punc-
tatus, and L. loxogrammus in lacking pterygoid teeth, from L.
personatus and L. pratensis in that the surface rugosities of the
frontal and parietal bones do not form a pattern of the overlying
scales, from L. apertosulcus in having a closed Meckelian canal,
from psammodromus, greenwayi, cubensis, punctatus and _sticto-
gaster in that the anterior border of the angular process of the
articular bones forms an obtuse angle rather than a right or acute
angle with the medial border of the articular, from macropus,
loxogrammus, personatus and raviceps in that the main axis of the
articular process projects medially rather than posteromedially
from the articular condyle, and from cuneus in that the articular
and retroarticular processes are not separated by a distinct in-
dentation.

Description of Type. The nearly complete left dentary is 16.5
mm long from its broken posterior border to the symphysis. Its
tooth row, measured in a straight line from the projected anterior
border of the first tooth to the projected posterior border of the
last tooth (both are missing) is 12.8 mm long. The height of the

Eruerwce: Extinct Jamaican Lizard 51

dentary at the position of the next-to-last tooth is 3.1 mm. There
are 24 teeth or vacant alveoli; missing are (front to back) teeth
number 1, 6, 8, 10, 17, 22 and 24. The second and third teeth are
bluntly conical and curve inward and slightly backward. The
fourth and fifth teeth are similar to these except that there is a
slight posterior cusp near the base of the crown in each. The
crown of the seventh tooth is very slightly flared, compressed
linguo-labially, and has a small anterior and posterior cusp below
the median one. The eleventh tooth and all of the more posterior
ones have a crown that is flared in an anterior-posterior direction,
linguo-labially compressed to form a moderately sharp cutting
edge, strongly tricuspid and curved inward. The anterior and
posterior cusps of each tooth are smaller than the median cusp
and separated from it by a wide groove that fades out at the base
of the crown. The occlusial edge of each of these teeth is slight-
ly oblique to the main axis of the tooth row. All of the tricuspid
teeth are crowded closely together so that the anterior cusp of each
tooth is overlapped labially by the posterior cusp of the preceding
tooth. Below their flared crowns the shaft of each tooth expands
gradually toward its ankylosed base. About 40 percent of each
tooth rises above the alveolar border of the dentary. A vertical
row of very small foramina is visible in the lingual face of the
dentary in the narrow spaces between the teeth.

The labial and ventral faces of the dentary are smooth, except
where the most superficial layers of bone have been erroded away
or scraped in preparation. The lower half of the labial face and
the ventral face are convex. The upper half of the labial face
is convex anteriorly, becoming flattened to slightly concave and
lightly scored posteriorly. A large, shallow, triangular and slightly
concave depression in the upper half of the posterior labial face of
the dentary extends forward to the level of the twentyfirst tooth,
marking the former position of the anterolateral process of the
coronoid. Four mental foramina form a row in the labial face
between the second and thirteenth tooth.

A large, > shaped indentation in the posterior border of the
lingual face of the dentary extends forward to the level of the
posterior border of the twentyfirst tooth, marking the former posi-
tion of the angular and splenial. Anterior to the apex of this
indentation (the former anterior border of the anterior inferior
alveolar foramen) the lingual face is shallowly concave to the

a2, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

level of the nineteenth tooth, and smoothly convex beyond this
point. Meckel’s canal penetrates the anterior end of the dentary
as a small opening just below and behind the symphysial surface.
Additional Dentaries. Three additional dentaries from Dary
Cave, two from Portland Cave I, two from Portland Cave III, and
one from Montego Bay Cave are referred to this species. Two
complete dentaries measure 12.8 mm and 12.4 mm along their
tooth rows and contain 24 and 21 teeth and vacant alveoli respec-
tively. The first 11 teeth are missing in one of these specimens
and the remaining teeth have flared, strongly tricuspid crowns as
in the type. In the other the crowns of the first seven teeth are
simply pointed, the eighth tooth is tapered and weakly tricuspid,
the ninth tooth is missing and the tenth and all of the more
posterior teeth have flared and strongly tricuspid crowns. In
a partial dentary the crowns of the first five teeth are simply
pointed, the sixth is missing, the seventh is tapered and weakly
tricuspid, the eighth and ninth are missing and the tenth and all
of the more posterior teeth are flared and strongly tricuspid. Thus,
the transition from simply pointed anterior teeth to flared and
strongly tricuspid posterior teeth may begin as far forward as
the fourth tooth and be completed as far posteriorly as the ninth.
Maxillae. Two maxillae from the type locality, 3 from Portland
Cave III and one from Portland Cave I are referred to this spe-
cies. ‘I'wo complete specimens measure 8.8 mm and 11.9 mm
along their tooth rows and contain 18 and 20 teeth plus vacant
alveoli respectively. In the smaller specimen the first two teeth
are simply pointed, the third is missing, the fourth is tapered and
weakly tricuspid and the fifth and all of the more posterior teeth
are flared and strongly tricuspid. In the larger specimen the first
three teeth are simply pointed, the fourth is tapered and weakly
tricuspid, the fifth and sixth are missing and the seventh and all
more posterior tooth crowns are flared and _ strongly tricuspid.
Premaxillae. Two premaxillae from Portland Cave III contain
seven teeth or vacant alveoli; one median tooth flanked by three
on each side. The crowns of all of the teeth are simply pointed
and curved slightly backward. They measure 4.0 and 4.5 mm
across the widest part of the rostrum. The former positions of the
overlapping nasal bones are indicated by smooth sutural scars on
the dorsal surface of the slender nasal process.
Frontals. One frontal from Portland Cave III and another

ETHERIDGE: Extinct Jamaican Lizard 58

from the Montego Bay Airport Cave may be definitely assigned
to the genus Leiocephalus because of their sharply projecting crista
cranii, a feature which distinguishes this genus from Anolis, the
only other iguanid genus in the caves. However, the two frontals
differ in their shape and ornamentation and possibly do not come
from the same species. If they do not, then it is uncertain which
one of them should be assigned to L. jamaicensis.

The Portland Cave specimen is 3.8 mm wide across its nar-
rowest interorbital distance and is estimated to have been about
§.6 mm wide across its parietal border. Heavy rugosities orna-
ment the entire dorsal surface and orbital margins of the bone
but do not form a regular pattern corresponding to the scales
which overlayed them. A deep, narrow, oval notch in the parietal
border indicates the former position of the parietal foramen.

The Montego Bay specimen is 2.5 mm wide across its narrow-
est interorbital width and is estimated to have been about 9.6 mm
wide across its parietal border. Thus, the orbital borders are more
strongly concave than in the Portland Cave specimen. In addi-
tion, the dorsal surface of the element is smooth, and a wide,
shallow indentation in the parietal border indicates the former
position of the parietal foramen.

Parietals. A nearly complete parietal, broken in preparation,
from the type locality and a fragment of another from the Montego
Bay Airport Cave are referred to this species. The Dairy Cave
specimen is 10.6 mm wide across its frontal border. The lateral
borders of the roof converge strongly toward the occipital border,
which has been broken away. Strong rugosities, similar to those
of the frontal from Portland Cave, ornament the dorsal surface
of the roof. The Montego Bay specimen, though fragmentary,
is obviously from a smaller individual. The lateral roof borders
converge posteriorly but remain widely separated at the occipital
border. Its upper surface, though not completely smooth, is
much less rugose than that of the Dairy Cave specimen.

Pterygoids. A nearly complete pterygoid from Portland Cave
and part of another from Montego Bay Airport Cave are referred
to this species. The anterior end of the palatine process and the
posterior end of the pterygoid process are broken away in the
Portland Cave specimen. It is 5.2 mm wide between the postero-
medial corner of the palatine plate and the lateral extremity of
the ectopterygoid process, and 1.5 mm wide across the narrowest

54 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

part where the palatine plate meets the pterygoid process. The
ventral surface of the palatine plate is perfectly smooth, without
the slightest trace of teeth or alveoli.

The Montego Bay specimen lacks the distal parts of the pala-
tine, pterygoid and ectopterygoid processes. It is 1.9 mm wide
across the narrowest part where the palatine plate meets the ptery-
goid process. The ventral surface of its palatine plate is also
smooth and lacks all evidence of teeth or alveoli.

Articular + Surangular. A fused articular + surangular from
Portland Cave III is 16.6 mm long from the posterior end of the
retroarticular process to the broken anterior end of the surangular;
about one mm should be added to this for the length of the intact
element.

The short, robust angular process projects medially from the
articular condyle for a distance of 2.4 mm. Its upper surface is
smoothly convex. Distally the process curves slightly upward.
The anterior border of the process curves irregularly forward to-
ward the medial border of the articular. The retroarticular process
projects posteriorly from the condyle for a distance of 2.6 mm and
is 2.6 mm wide at the condyle. Its medial, lateral and posterior
borders are somewhat raised, surrounding a shallow, more or less
rectangular depression. The retroarticular and angular processes
are united posteromedially by a thin shelf whose margin forms a
very shallow sigmoid curve between the distal extremities of the
two processes.

Caudal Vertebrae. Five posterior segments of autotomic cau-
dal vertebrae from the type locality, four posterior segments, one
anterior segment, an intact autotomic vertebra and nine vertebrae
from the anterior, nonautotomic part of the tail from Portland
Cave III, and one posterior segment from the Montego Bay Air-
port Cave are referred to this species.

The centrum of the largest nonautotomic vertebra is 5.0 mm
long, including its condyle. Well developed zygosphenes are
present on a median projection of the neural arch between the
prezygapophyses, and corresponding zygantra are present between
the postzygapophyses. The neural spine is strongly compressed
and tapered. A thin, median crest rises above the neural arch,
sloping steeply downward from the neural spine to the interzy-
gapophyseal projection of the arch. The centrum is strongly com-
pressed ventrally to form a blunt, median keel below. The trans-

ETHERIDGE: Extinct Jamaican Lizard 55

verse processes are wide and flat, slightly tapered toward their
distal extremities, and oriented posterolaterally.

Measurements of the intact autotomic vertebrae are: greatest
length of the centrum, including its condyle 6.0 mm, length of the
anterior segment from the anterior ventral border of the centrum
to the fracture plane 1.0 mm, length of the posterior segment from
the fracture plane to the anterior ventral border of the condyle
3.2 mm, width of the centrum posterior to the plane of fracture
1.2mm. Moderately well developed zygosphenes and zygantra
are present on a small median projection of the neural arch be-
tween the prezygapophyses and corresponding zygantra are pres-
ent between the postzygapophyses. A thin, median crest rises
vertically above the neural arch between the neural spine and the
fracture plane. It ascends abruptly above the plane as a thin,
spine-like projection, then abruptly descends to the neural arch
of the anterior segment. The fracture plane passes more or less
vertically through the spine, down through the neural arch, curves
posteriorly around the bases of the transverse processes and con-
tinues vertically down through the centrum. The dissociated seg-
ments of the other autotomic caudal vertebrae are similarly con-
structed; however their proportions differ greatly, due, no doubt,
to their different points of origin from the caudal vertebral column.

Body Vertebrae. Two body vertebrae from Portland Cave I
and two from Portland Cave III are referred to this species. The
largest, apparently from near the middle of the body, has a cen-
trum, including its condyle, 5.9 mm long. Zygosphenes and zy-
gantra are moderately well developed. The neural spine is ro-
bust, strongly compressed and wider at the top than at the base.
The sharp anterior border of the spine slopes steeply downward
to the neural arch, and continues forward as a low, median neural
crest. The centrum is strongly compressed ventrally to form a
blunt, median keel.

Maximum Size. The snout-vent lengths of the animals from
which the fossils came have been calculated by multiplying var-
ious measurements of the fossils by the ratio of measurements of
the same elements of skeletons of modern species to their snout-
vent lengths. Assuming that the proportions which existed in the
extinct species fall within the limits of those which exist among
all living forms, a minimum and maximum estimate of snout-vent
length may be obtained for each fossil of the extinct species. The

56 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

minimum and maximum estimates for the largest fossil of each
of several elements are as follows: dentary 90-120 mm, maxilla
111-120 mm, frontal 109-134 mm, parietal 107-131 mm, and articu-
lar-surangular 130-170 mm. Thus, L. jamaicensis probably reached
a maximum size of at least 130 mm and may have reached a max-
imum size as great as 170 mm.

Comparisons. The fossils described here are referred to the
genus Leiocephalus on the basis of numerous osteological char-
acteristics which distinguish that genus from the related South
American genera Stenocercus, Tropidurus, Ctenoblepharis, Lio-
laemus, Urocentron, Proctotretus, Platynotus, Plica, and Ophryoes-
soides (Estes, 1963; Etheridge, 1964a, 1965, 1966a). The osteolog-
ical characteristics which distinguish these genera, the tropidurines,
from other groups of iguanid lizards have been given by Etheridge
(1964b).

Slight but consistent structural differences in the skeleton may
be used to differentiate most of the living and extinct species of
Leiocephalus. Those which distinguish L. jamaicensis are given
in the diagnosis. Although these differences are useful for the
identification of fossil bones, most of them are not of the sort
that may be used to indicate the phylogenetic relationships of the
species to one another. For this reason little can be said as to the
probable relationships of L. jamaicensis to other forms, except that
since it can be distinguished from L. schreibersi, L. melanochloris,
and L. inaguae only by its larger maximum size, it is probably
most closely allied to those forms.

The maximum snout-vent length attained by any living species
is 130 mm, recorded for L. carinatus microcyon (Schwartz, 1959).
L. hermimeri, which apparently became extinct on Martinique in
this century, reached 139 mm (Boulenger, 1885). The extinct forms
L. cuneus of Barbuda and L. apertosulcus of Hispaniola appar-
ently reached a maximum snout-vent length of about 200 mm
(Etheridge, 1964a, 1965). Only two living species have been
reported as fossils: L. personatus from Cerro de San Francisco, Do-
minican Republic, (Etheridge, 1965) and L. carinatus from New
Providence Island (Etheridge, 1966b). The maximum size of these
fossils is approximately equal to that attained by modern popula-
tions of the same species at those localities today.

Other living species of West Indian lizards are known as fos-
sils and a number of them grew to a larger size than in modern

ETHERIDGE: Extinct Jamaican Lizard 57

populations from the same area; e.g. Anolis leachii on Barbuda,
Anolis ricordi on Hispaniola, and Anolis sagrei on New Providence.
In these species the ontogenetic gradients known to exist in modern
populations are continued in the fossils to a greater maximum size.
Hecht (1951) described an extinct Jamaican gecko, Aristelliger
titan, which attained a maximum size greater than the largest liv-
ing species. However, there appears to be no structural differ-
ence between A. titan and the modern Hispaniolan species A. lar
that cannot be accounted for solely on the basis of the larger size
of A. titan (Etheridge, 1965). Thus there are no anatomical rea-
sons for not considering A. titan to have been a Jamaican popula-
tion of A. lar that grew to a larger size than does the present pop-
ulation of the species on Hispaniola. Leiocephalus jamaicensis
may be distinguished from schreibersi, melanochloris, and inaguae
only by its greater maximum size. As is the case of A. titan and
A. lar, decision to recognize L. jamaicensis as an extinct species,
rather than as a large Jamaicana representative of L. melanochloris,
L. schreibersi, or L. inaguae, must be made on some grounds other
than anatomical comparisons.

In my description of the Barbudan form, L. cuneus (Etheridge,
1964a), I failed to come to grips with the problem of distinguish-
ing it from L. herminieri of Martinique. L. cuneus differs from all
living species in the extreme anterior position of the transition
from simple to tricuspid teeth; however, I have no information
as to the position of this transition in the now extinct L. her-
minieri. L. cuneus grew considerably larger than L. herminieri,
but the possibility remains that L. cuneus merely represents a pop-
ulation of L. herminieri that lived on another island and grew to
a larger size.

Discussion

Liocephalus, Stenocercus, Tropidurus, Ctenoblepharis, Liolae-
mus, Urocentron, Proctotretus, Platynotus, Plica, and Ophryoes-
soides appear to form a group of related genera, the tropidurines,
within the family Iguanidae, and their closest allies appear to be
the sceloporine genera of North America, Sceloporus, Uta, Uro-
saurus, Petrosaurus, Sator, Uma, Callisaurus and Holbrookia (Eth-
eridge, 1964b). Leiocephalus is the most distinctive genus among
the tropidurines; that is, the skeletal and integummentary differ-
ences between it and the other tropidurine genera are greater than

58 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

those between any other two tropidurine genera (Etheridge, 1966a).
The existence of Leiocephalus in the Early Miocene of Florida
(Estes, 1963) indicates that the peculiar specializations of the genus
were attained by at least mid-Cenozoic, and therefore the diver-
gence between the tropidurines and sceloporines must have oc-
curred at a still earlier time.

Two possibilities are apparent. 1) Sceloporines and tropidurines
may have coexisted in North and Middle America during the mid-
Cenozoic, and that following the migration of Leiocephalus into
the West Indies the continental tropidurines became extinct north
of Panama; or 2) that tropidurines and sceloporines diverged in
South and North America, respectively, following some event
which caused the separation of their common ancestral stock, e.g.,
the Eocene opening of the Panamanian portal, that Leiocephalus
evolved in the West Indies from a South American tropidurine im-
migrant, and that the Florida fossils represent an invasion from
the islands to the North American mainland.

Because of the close geographic proximity of Florida to the
West Indies, and the inconclusive evidence (Estes and Tihen,
1964, p. 466) tor the former existence of tropidurines elsewhere
on the mainland north of Panama, the latter alternative appears
more reasonable. If this is correct, then Leiocephalus must have
achieved its distinctiveness in the West Indies at least by mid-
Cenozoic. The former existence of Leiocephalus on Jamaica and
Barbuda clearly indicates that the genus must have occurred
throughout most if not all of the West Indies in the past. Further-
more, the presence of extinct species of Leiocephalus in three of
the four large West Indian lizard faunas that have been studied
to date indicates that the current distribution of the genus may
not reflect the extent of its range and structural diversity even in
the very recent past.

ACKNOWLEDGMENTS

Most of the material reported here was collected by Dr. Walter
Auffenberg with financial assistance from National Science Foun-
dation grant G-3896. I am grateful to Dr. J. C. Dickinson, Jr.,
Director of the Florida State Museum, and Dr. Clayton E. Ray,
former Assistant Curator of Natural Sciences, for providing me
with facilities and financial support during the summer of 1963,
when this study was begun. I am also grateful to Dr. Ernest E.

ETHERIDGE: Extinct Jamaican Lizard 59

Williams of the Museum of Comparative Zoology at Harvard and
to Dr. Charles Walker of the University of Michigan Museum of
Zoology for the loan of skeletons. Correspondence with Dr. Auf-
fenberg on various aspects of this study has been extremely helpful.

LITERATURE CITED

BOULENGER, GEorRGE A. 1885. Catalogue of the lizards in the British Mu-
seum (Natural History). London, vol. 2, pp. 1-497.

Estes, RicHarp. 1963. Early Miocene salamanders and lizards from Flor-
ida. Quart. Jour. Florida Acad. Sci., vol. 26, no. 3, pp. 234-256.

Estes, RicHARD, AND JosEPH A. TIHEN. 1964. Lower vertebrates from the
Valentine Formation of Nebraska. Amer. Midland Naturalist, vol. 72,
pp. 453-472.

ETHERIDGE, Ricuarp. 1964a. Late Pleistocene lizards from Barbuda, Brit-
ish West Indies. Bull. Florida State Mus., vol. 9, no. 2, pp. 43-75.

1964b. The skeletal morphology and systematic relationships of
sceloporine lizards. Copeia, no. 4, pp. 610-631.

—. 1965. Fossil lizards from the Dominican Republic. Quart. Jour.
Florida Acad, Sci., vol. 28, no. 1, pp. 83-105.

—. 1966a. The systematic relationships of West Indian and South
American lizards referred to the iguanid lizard genus Leiocephalus.
Copeia, no. 1, pp. 79-90.

1966b. Pleistocene lizards from New Providence. Quart. Jour.
Florida Acad. Sci., vol. 28, no. 4, 1965 (1966), pp. 349-358.

Hecut, Max k. 1951. Fossil lizards of the West Indian genus Aristelliger
(Gekkonidae). Amer. Mus. Novitates, no. 1538, pp. 1-33.

KoopMAN, Karu F., AND ERNEST EF. WitiiaMs. 1951. Fossial Chiroptera
collected by H. E. Anthony in Jamaica, 1919-1920. Amer. Mus. Novi-
tates, no. 1519, pp. 1-29.

ScHwartz, ALBERT. 1959. The Cuban lizards of the species Leiocephalus
carinatus (Gray). Reading Publ. Mus. and Art Gallery Sci. Publ., no.
10, pp. 1-497.

San Diego State College, San Diego, California.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

Growth and Longevity of Rats Fed Different Diets
L. R. ARRINGTON AND C. B. AMMERMAN

THE average life span of the male laboratory rat is considered
to be about 700 days; females are reported to live about 10 per
cent longer (Carlson and Hoelzel, 1947; Sherman et. al., 1949;
French et al., 1953; and Sperling et al., 1955): Maximum life may
reach 1200 days in the male and 1300 days in the female (Berg and
Harmison, 1957). Many factors, including nutrition, are known to
influence longevity. Considerable evidence is available to show
that certain types of food restriction or undernutrition may result
in longer life span than high nutrient intakes which permit maxi-
mum growth (McCay, et al., 1935; McCay, 1947; Silberberg and
Silberberg, 1955; Ross, 1959; Berg and Simms, 1960; and Berg,
1960).

The rat is considered to remain in a continuous state of growth
(Dunn et al., 1947), but maximum body weight may be expected
just beyond 400 days. A decline in weight accompanies aging and
may equal 30 per cent during the last 300 days (Everitt, 1957; Ev-
eritt et al., 1957).

Many different strains of laboratory rats and different dietary
regimes are used in research. The purpose of the study reported
here was to determine the growth and longevity of Long-Evans
rats fed natural, semi-purified, and purified diets.

PROCEDURE

Weanling rats of the Long-Evans strain produced in this lab-
oratory from a closed colony were used as experimental animals.
The experimental young were weaned at 23-24 days of age from
females which had been fed a complete commercial pelleted rat
diet (Purina Laboratory Chow). At the time of weaning they were
placed on the respective dietary regimes and housed individually
in stainless steel cages maintained in an air conditioned room at
77-79° F. Experimental diets and fresh tap water were supplied
ad libitum. No other treatments were given throughout the life
span other than periodic removal for weighing and transfer to
clean cages. No exercise was enforced and the only activity was
voluntary movement within the individual cage.

The diet referred to as the natural diet was a complete com-

ARRINGTON AND AMMERMAN: Diet and Longevity 61

mercial pelleted ration considered to supply nutrient requirements
of the rat.

The semi-purified diet contained, in per cent: corn meal, 38;
corn starch, 10; vitamin free casein, 10; zein, 10; sucrose, 25; min-
erals (USP XIV), 3; corn oil, 2; NaCl, 0.5; and vitamins (Vitamin
Diet Fortification, Nutritional Biochemicals Corp.), 1. This diet
was also modified for an additional treatment by replacement of
the casein with zein in order to provide a diet deficient in the
amino acids lysine and tryptophane. Fourteen rats were fed this
amino acid deficient diet, and 14 were fed the control with casein.
In addition, 138 rats which had been fed the amino acid deficient
ration for 4 to 8 weeks were changed to the control diet. In pre-
liminary studies, this deficiency had resulted in growth failure and
early mortality of the rats. The change to a diet supplemented
with lysine and tryptophane was made in order to observe the
effects of this early dietary deficiency upon longevity after the
change to a normal diet. The transfers to the control diet were
made at 4 to 8 weeks after consuming the deficient diet, and the
time of change was selected for each rat when severe symptoms
of deficiency were so evident that it apparently would not have
survived without amino acid supplementation.

The control purified diet contained, in per cent: corn starch, 40;
sucrose, 22; casein, 26; corn oil, 4; cellulose, 3; minerals, 4; and
vitamins, 1. Two similar diets containing less casein (8 and 14
per cent) and corresponding increases in carbohydrates were fed
to additional rats in order to study the effect of level of protein
intake upon longevity. The control diet supplied 22 per cent pro-
tein, considered to be adequate for normal growth; the two lower
levels of casein were considered inadequate.

Growth data were obtained from weekly body weight meas-
urements during the first two months after weaning and at ap-
proximately monthly intervals throughout the life span. Statisti-
cal comparisons were based upon the analysis of variance.

RESULTS AND DISCUSSION

Data representing the average and range of life span of rats
fed the three diets are recorded in table 1. Longevity was not
affected by the different types of diets. Male rats consuming the
semi-purified diet appeared to live longer than those consuming

62

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Diet

Natural

Purified

Semi-purified

Ay. all treatments

IN GRAMS

BODY WEIGHT

600
500
400
300
200

100

Bice 1

TABLE 1

Longevity of Rats Fed Different Diets

No. Life Span (days)

Sex Rats Av. Range

re) 5 657 379-993
2 5 889 640-1045
re) 8 667 227-1298
2 10 901 511-1211
} 7 805 340-1154
Q il 897 552-1270
3 19 718 227-1293
2 20 897 511-1270

AGE IN DAYS

Excess
2 over

Days

232

Weight changes of Long-Evans rats during life span.

cy

%
35

35

19

25

ARRINGTON AND AMMERMAN: Diet and Longevity 63

the natural or purified diets, but considerable variation was ob-
served and the difference was not statistically significant. Within
all treatments, females lived longer than males. The average ex-
cess of females over males was 179 days or approximately 25 per
cent longer. This additional survival time of females is greater
than that reported by others (Carlson et al., 1947; French et al.,
1953; Sperling et al., 1955). No evidence of specific infections was
observed and all deaths recorded were included in the average.

Average body weights of the two sexes through the major por-
tion of the life span are plotted in figure 1. The weights repre-
sent the combined average of those consuming the natural, control
semi-purified, and highest level of protein in the purified diet. It
should be noted that the number of rats surviving the latter por-
tion of the period had been decreased and the body weight aver-
ages represent a progressively smaller number of animals. During
the terminal stages of life most of the rats lost weight for periods
which ranged from a few days to several months. Using the
highest weight attained and the lowest near death, the average
decreases for males was 18 per cent, for females 13 per cent.

Rats fed the amino acid deficient diet gained very little and
had a very short life span (table 2, figure 2). When other rats
which had been fed this diet for 4 to 8 weeks were changed to
the contro] diet containing casein, body weight increased rapidly
and the life span was essentially equal to that of rats consuming
the adequate diet continuously. The average life span of the
former group was numerically less, but considerable variation was
observed and the difference was not statistically significant.

The purified diets which were inadequate in quantity of pro-
tein (7 and 12 per cent) promoted a slower rate of weight gain
than the control, but length of life spran was not affected (table 2).
Although the gain of rats consuming the lowest level was much
less during the early months of the treatment period, after 16
months the average weight approximated 85 per cent of the higher
level. These observations are in general agreement with those
of others that certain systems of restricted feed or nutrient intake
do not decrease life span (McCay et al., 1935; McCay, 1947; Sil-
berberg et al., 1955; Ross, 1959). The quality of protein fed in
this trial was considered satisfactory and quantity only was re-
stricted.

64 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 2

Effect of Amino Acid Deficiency, Change from Amino Acid Deficient
to Adequate Diet and of Protein Intake Upon Longevity

No. Life Span (days)

Rats Av. Range

Amino acid deficient* 14 187 30-431
Amino acid deficient

changed to adequate* 13 737 397-1082

22% Protein** 6 754 253-1059

12% Protein** 6 745 227-1002

7% Protein** 6 796 449-1293

*Semi-purified diet.
**Purified diet, all protein supplied by casein.

600

500

400 (851)
G37)

300

® CONTROL

* AMINO ACID DEFICIENT,
CHANGED TO CONTROL

© AMINO ACID DEFICIENT

200

BODY WEIGHT IN GRAMS

100
CTS)

FIGURES IN PARENTHESIS REPRESENT
AVERAGE LIFE SPAN

O 100 200 300 400 500 600 700 800

AGE IN DAYS

Fig. 2. Growth and longevity of rats fed a semi-purified control diet,
amino acid deficient diet, and amino acid deficient changed to control diet.

ARRINGTON AND AMMERMAN: Diet and Longevity 65

SUMMARY

Weanling Long-Evans rats were fed natural, semi-purified, or
purified diets in a growth and longevity study. For additional
studies, the semi-purified diet was modified to be deficient in the
amino acids lysine and tryptophane, and the purified diet included
three levels of protein, two of which were inadequate for normal
growth. The average life span of rats on the different types of
complete diets was not significantly different. Combined average
for longevity of the males from each group was 718 days; fe-
males, 897 days. Growth of rats fed the amino acid deficient diet
was severely retarded and average life span was 187 days. Rats
fed the amino acid deficient diet for 30 to 60 days and changed
to the control increased rapidly in weight and lived essentially a
normal life span. Purified diets containing 7 or 12 per cent pro-
tein as casein promoted slower growth than the diet with 22 per
cent protein but did not significantly affect life span.

LITERATURE CITED

Berc, B. N. 1960. Nutrition and longevity in the rat. I. Food intake in
relation to size, health and fertility. Jour. Nutr., vol. 71, pp. 242-254.

Berc, B. N., AND H. S. Sms. 1960. Nutrition and longevity in the rat. II.
Longevity and onset of disease with different levels of food intake.
jou Nutr vol) 71, pp. 255-268.

Bere, B. N., AND C. R. Harmison. 1957. Growth, disease and aging in the
rat. Jour. Gerontology, vol. 12, pp. 370-377.

Carson, A. J., AND F. Hoeizex. 1947. Growth and longevity of rats fed
omnivorous and vegetarian diets. Jour. Nutr., vol. 34, pp. 81-96.

Dunn, M. S., E. A. Murpuy, AnD L. B. Rockianp. 1947. Optimal growth
of the rat. Physiol. Rev., vol. 27, pp. 72-94.

Everitt, A. V. 1957. The senescent loss of body weight in male rats.
Jour. Gerontology, vol. 12, pp. 382-387.

Everitt, A. V., AND C. Wess. 1957. The relation between body weight
changes and life duration in male rats. Jour. Gerontology, vol. 12,
pp. 128-135.

FRENCH, C. E., R. H. INcram, J. A. UrAM, G. P. Barron, AnD R. W. SwIrFT.
1953. The influence of dietary fat and carbohydrate on growth and
longevity in rats. Jour. Nutr., vol. 51, pp. 329-339.

66 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

McCay, C. M. 1947. Effect of restricted feeding upon aging and chronic
diseases in rats and dogs. Amer. Jour. Pub. Health, vol 37, pp.
DPSS),

McCay, C. M., M. M. CroweLit, AND L. A. Maynarp. 1935. The effect
of retarded growth upon the length of life span and upon ultimate
body size. Jour. Nutr., vol. 10, pp. 63-79.

Ross, M. H. 1959. Protein, calories and life expectancy. Fed. Proc., vol.
18, pp. 1190-1207.

SHERMAN, H. C., H. L. CampsBett, anp M. S. Racan. 1949. Analytical
and experimental study of the effects of increased protein with liberal

calcium and riboflavin intakes: Complete life cycles. Jour. Nutr., vol.
37, pp. 317-327.

SILBERBERG, H., AND R. SILBERBERG. 1955. Diet and life span. Physiol.

Rev., vol. 35, pp. 347-362.

SPERLING, G., F. LovEntace, L. L. BAarnss, C. A. H. Smitn, J. A. SAXON, Jr.,
AND C. M. McCay. 1955. Effect of long time feeding of whole milk
diets to white rats. Jour. Nutr., vol. 55, pp. 399-414.

Department of Animal Science, University of Florida, Gaines-

ville, Florida. Florida Agricultural Experiment Stations Journal
Series No. 2165.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

Effect of Vitamins A and E on Copper in Heart of Cattle
he. SrIRLEY, H. L. CHAPMAN, Jr., J. F. Eastey, AND T. J. CUNHA

CERTAIN dietary factors are known to affect the concentration
of copper in tissues of animals. McCall and Davis (1961) demon-
strated that high dietary protein levels markedly decreased the
deposition of dietary copper in the liver of rats. High dietary
levels of copper resulting in increased levels of vitamin A in the
liver of swine demonstrated an interrelationship between these
nutrients (Shirley, et al., 1962). Kneta (1963) found 0.5 and 0.39
mg per cent copper in the myocardium of calves and cattle, re-
spectively, and that heart contained more copper than skeletal
tissue.

The present study was made to determine if (a) high levels of
vitamins A and E given orally during November through March,
(b) high levels of vitamins A and E given orally during June through
September, and (c) high levels of vitamins A and E given by intra-
muscular injection, affected the concentration of copper, ash, and
water in the ventricle of the heart of cattle.

PROCEDURE

Three trials were made.

Trial 1. High dietary levels of vitamins A and E in concen-
trate rations on winter pastures. Seventy-two Brahman-British
crossbred steers, with an average age of ten months, were fed ad
libitum a ration consisting of ground snapped corn, dried citrus
pulp, cottonseed meal, urea, and minerals. By analysis, the ration
contained approximately 8 ppm copper. During the experimental
period they grazed Roselawn St. Augustine pasture from Novem-
ber through March for 136 days. Monthly analyses of the pasture
during this period gave average values of approximately 4 ppm
copper, 2.5 ppm molybdenum, 40 mg carotene per kg dry weight
and 0.2 per cent nitrate. Twenty-four steers were randomly al-
lotted to each of three vitamin A palmitate supplementation levels
of 0, 25,000, and 50,000 IU (International Units) per day. The
steers fed each level of vitamin A were subdivided into groups of
eight steers each and fed 0, 50, and 250 IU of vitamin E (alpha-
tocopherol acetate) per day. The steers were slaughtered after
136 days on the treatments, and the Jeft ventricle was frozen im-

68 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

mediately at —8 C and analyzed within a few days for copper, ash,
and water.

Trial 2. High dietary levels of vitamins A and E in concen-
trate rations on summer pastures. Thirty-two Brahman-Angus
crossbred steers approximately ten months of age were fed the
same basal ration ad libitum as those in trial 1 above, during June
through September for 98 days. Monthly analyses of the pasture
during this period gave average values of 6.3 ppm copper, 3.0 ppm
molybdenum, 60 mg carotene per lb. dry weight and 0.2 per cent
nitrate. Half were supplemented with 25,000 IU of vitamin A per
day. Each of these two vitamin A dietary groups was subdivided
and half fed 50 IU of vitamin E per day. The steers were slaugh-
tered after 98 days on the treatments, and the left ventricle was
frozen as above prior to analysis for copper, ash, and water.

Trial 3. High levels of vitamins A and E injected intramuscu-
larly. Thirty-two Brahman-British crossbred steers approximately
ten months old were fed the ration of trials 1 and 2 above ad libitum
for 160 days. They were divided into four equal groups on the
basis of weight and each group given one of the following treat-
ments: (a) no supplementary vitamins A and E, (b) 720,000 IU of
vitamin A, (c) 1,400 IU of vitamin E (alpha-tocopherol acetate), and
(d) 720,000 IU of vitamin A plus 1,400 IU of vitamin E per 28
days per steer by intramuscular injection in the rump muscle.
They were slaughtered at the end of the trial and the left ven-
tricle frozen immediately at —8 C and analyzed for copper, ash,
and water.

Water and ash were determined by the A.O.A.C. (1960) oven
and furnace methods, respectively; and copper by the dithiocarba-
mate method of Sandel (1959). Statistical analyses were made
according to Duncan’s Multiple Range Test (1955).

RESULTS AND DISCUSSION

Trial 1. High dietary levels of vitamins A and E in concen-
trate rations on winter pasture. The effects of vitamins A and E
on the concentration of water, ash, and copper in the ventricle of
the steers are shown in Table 1. The treatments had no signifi-
cant effect on these constituents. Average values for water in the
heart ventricle varied among the treatment groups from 77.4 to
78.7 per cent. Corresponding values for percentages of ash varied

SHIRLEY ET AL.: Vitamins and Copper 69

from 4.7 to 5.1, and copper concentrations ranged from 17.5 to
24.7 ppm on the dry weight basis.
TABLE 1

Copper level in heart of cattle on winter pasture with fattening ration
and supplemental vitamins A and E (8 steers per treatment)

Vitamin A Vitamin E Water Ash Copper
IU IU % % dry wt ppm dry wt
0 0 TE 4.8 19.7
0 50 Tso 4.8 20.2
0) 250 TSoa 4.9 DAT
25,000 0 717.9 5.0 aS
25,000 50 78.4 4.7 17.8
25,000 250 78.2 4.8 Tile,
50,000 0 78.6 Halt Pa Tl
50,000 50 77.8 4.9 21.8
50,000 250 11.4 5.0 DOD,

Trial 2. High dietary levels of vitamins A and E in concen-
trate rations on summer pasture. The levels of supplementation
of vitamins A and E on summer pasture in this trial had no effect
on water, ash, and copper in the ventricle as shown in Table 2.
These values are very similar to those of Table 1. Weight gains,
carcass data, and feed consumption of the steers of trials 1 and 2
were reported by Chapman et al. (1964).

TABLE 2

Copper level in heart of cattle on summer pasture with fattening ration
and supplemental vitamins A and E (8 steers per treatment)

Vitamin A Vitamin E Water Ash Copper
IU IU % % dry wt ppm dry wt
0 0 79.9 5.0 20
25,000 0 80.0 or 21
0 50= 79.8 Del Wi

25,000 50 79.9 5.1 20

70 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Trial 3. High levels of vitamins A and E injected intramuscu-
larly. Data are presented in Table 3 for the concentration of wa-
ter, ash and copper in the ventricle of steers given vitamins A and
E by intramuscular injection. Average water values for the treat-
ments varied only from 79.1 to 79.2 per cent; corresponding ash
values ranged from 5.3 to 5.4; and average copper concentrations
varied from 16 to 19 ppm on the dry weight basis. In the liver,
average copper values among the treatment groups varied from
360 to 434 ppm on the dry weight basis at the time the steers
were slaughtered. The injected vitamin A and E treatments had
no significant effect on the concentration of the water, ash, and
copper in these tissues. Injection of vitamins A and E intramuscu-
larly resulted in essentially the same tissue concentrations of water,
ash, and copper as when these vitamins were administered orally
at high levels as in trials 1 and 2.

TABLE 3

Copper level in heart of steers injected intramuscularly with vitamins
A and E (8 steers per treatment)

Vitamin A Vitamin E Water Ash Copper

IU IU %o % dry wt ppm dry wt
0 0 79.1 5.3 19
720,000* 0 79.2 5.3 16
0 1,400* 79.1 5.4 i)

~720,000* 1,400* UDP 5.4 18

"Injected intramuscularly each 28 days during 160-day feeding trial.

SUMMARY

Separate groups of Brahman-British crossbred steers approxi-
mately 10 months of age were fed ad libitum a ration consisting
of ground snapped corn, dried citrus pulp, cottonseed meal, urea,
and minerals for 136 days on winter pastures, and 98 days on sum-
mer pastures. During the winter trial, 24 steers were randomly
allotted to each of three vitamin A palmitate oral supplementation
levels of 0, 25,000, and 50,000 IU per day. The steers fed each
level of vitamin A were subdivided into groups of eight steers
each and fed 0, 50, and 250 IU of vitamin E (alpha-tocopherol ace-

SHIRLEY ET AL.: Vitamins and Copper tl

tate) per day. During the summer trial the same treatments were
made as during the winter except those treatment groups given
50,000 IU of vitamin A and 250 IU of vitamin E were deleted.
Vitamins A and E were injected intramuscularly into the rump of
similar animals fed the same ration on pasture as above during the
winter for 160 days. These treatments were (a) controls, (b) 720,000
IU of vitamin A every 28 days, (c) 1,400 IU of vitamin E, and (d)
720,000 IU of vitamin A plus 1,400 IU of vitamin E per 28 days
per steer.

The treatments had no significant effect on the concentration
of copper, ash, and water in the ventricle of the steers. Those
given the oral doses of vitamins A and E during the winter had
average values for copper in the ventricle that varied among the
treatment groups from 17.5 to 24.7 ppm; and corresponding values
for percentages of ash that varied from 4.7 to 5.1. Average values
for water varied from 77.4 to 78.7 per cent. Similar values were
obtained in those steers given oral doses of the vitamins during
the summer, and those injected intramuscularly with large doses
of vitamins A and E.

ACKNOWLEDGMENTS

This study was supported in part by grants-in-aid from the
National Heart Institute, NIH HE-01318 and from the Moorman
Manufacturing Company. The authors are indebted to A. Z.
Palmer for technical assistance and to G. K. Davis for valuable
suggestions during the study.

LITERATURE CITED

ASSOCIATION OF OFFICIAL AGRICULTURAL CHEMists. 1960. Official methods
of analysis. Washington, D. C. 832 pp.

CHAPMAN, H. L., Jr., R. L. SHrriey, A. Z. PALMER, C. E. HAIngEs, J. W.
CARPENTER, AND T. J. CuNnHA. 1964. Vitamins A and E in steer fat-
tening rations on pasture. Jour. Animal Sci., vol. 23, pp. 669-673.

Duncan, D. B. 1955. Multiple range and multiple F tests. Biometrics,
vol. 11, pp. 1-8.

Kneta, Z. 1963. Trace element content of the myocardium. Latvijas PSR
Zinatnu Akad. Vestis, vol. 9, pp. 115-122. (Chem. Abstr.)

McCa tu, J. T., AND G. K. Davis. 1961. Effect of dietary protein and zinc
on the absorption and liver deposition of radioactive and total copper.
Jour. Nutrition, vol. 74, pp. 45-50.

72 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SANDELL, E. B. 1959. Colorimetric determination of traces of metals. In-
terscience Publishers, Inc., New York, N. Y. Vol. 3, pp. 1032.

SHIRLEY, R. L., T. N. Mracnam, A. C. Warnick, H. DD. WALLACE ieee
Eastey, G. K. Davis, anp T. J. CunHa. 1962. Gamma irradiation
and interrelation of dietary vitamin A and copper on their deposition
in the liver of swine. Jour. Nutrition, vol. 78, pp. 454-456.

Everglades Experiment Station, Belle Glade, and Department

of Animal Science, University of Florida, Gainesville, Florida.
Florida Agricultural Experiment Stations Journal Article No. 2155.

Quart. Jour. Florida Acad. Sci. 29(1) 1966

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1966

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Presbyterian College
Florida Southern College
Florida State University
Jacksonville University
Marymount College
Miami-Dade Junior College
Mound Park Hospital Foundation
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida
University of Tampa

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1966

President: MarGARET GILBERT
Department of Biology, Florida Southern College
Lakeland, Florida

President Elect: Jackson P. SIckELs
Department of Chemistry, University of Miami
Coral Gables, Florida

Secretary: Joun D. KiLBy
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: JAMES B. FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: Prerce BRODKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges

should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

@
he
%iF cs Quarterly Journal

\

oR

ol of the

Florida Academy of Sciences

Vol. 29 June, 1966 No. 2
CONTENTS

Organotin esters McDonald Moore and Francis C. ie 73

A new monogenetic trematode from Georgia Charles E. Price 77

Barnacles of the northeastern Gulf of Mexico Harry W. Wells 81

Minima of eclipsing binaries VV Orionis and Beta Persei K-Y. Chen 96

New records of Bahamian Odonata Dennis R. Paulson 97

Hermaphroditism in mullet, Mugil cephalus Linnaeus
Martin A. Moe, Jr.

Review of the Lutjanus campechanus complex of red snappers
Luis R. Rivas

Centrarchid spawning in the Florida Everglades James P. Clugston

The exotic herpetofauna of southeast Florida
Wayne King and Thomas Krakauer

The vertebral musculature of Chersydrus (Serpentes) Walter Auffenberg

111

EL7

137

144

155

Mailed September 8, 1966

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes origina] articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 84% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Makraeins should be 1% inches all around.

TirLes should not exceed 42 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTSs in the text and
AppREss in paragraph form following Literature Cited.

LITERATURE CiTED follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLES are charged to authors at $16.70 per page or fraction. Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

Lecenps for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($15.80 per page, $14.00 per half
page for zinc etchings; $14.50 per page, $13.25 per half page for halftones).
Drawincs should be in India ink, on good board or drafting paper, and let-
tered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. PHoToGcRAPHS should be of good contrast, on glossy paper. Do not
write heavily on the backs of photographs.

ProoF must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

Nols 29 June, 1966 No. 2

Organotin Esters
McDonaLp Moore AND FRANCIS C. LANNING

In connection with our current research on acyloxygermanes,
we prepared the heretofore unknown tin tetrabenzoate. ‘Tin tetra-
butyrate was also prepared by a method unreported for it prepa-
ration.

Schmidt, Blohn, and Jander (1947) prepared tin tetracetate by
solvolysis reaction of stannic iodide in acetic anhydride, and An-
derson (1964) prepared tin tera-i-butyrate from stannic chloride and
silver isobutyrate. Henderson and Holliday (1965) prepared the
above two tin esters and tin tetraformate, tin tetrabutyrate, tin
tetralaurate, and tin-tetrastearate from the reaction of tetravinyltin
with the carboxylic acids.

In the present work we carried out the synthesis of two organo-
tin esters from stannic chloride and the sodium salts of organic acid
in anhydrous benzene. Tin tetrabutyrate and tin tetrabenzoate
were prepared in 83.2 and 74.6 per cent yields, respectively (Table
1). Tin tetrabenzoate is a white solid that begins to decompose at
118°C, while tin tetrabutyrate is a colorless semisolid.

Like tin tetracetate (Schmidt, Blohn, and Jander, 1947) these
organotin esters react with water and alcohol. They decompose
when heated.

Infrared spectra of these compounds were determined between
4000 and 700 cm?. Both organotin esters show significant ab-
sorption bands near 1704 and 1262 cm, corresponding to values
reported by Henderson and Holliday (1965) as characteristic of the
carbonyl] stretching bands in organotin esters. No anhydrides were
present, as no bands occur between 1850 and 1720 cm.

A benzene solution of tin tetrabenzoate reacts with twice the
stoichiometric amount of phenylmagnesium bromide to yield after

74 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

hydrolysis tetraphenyltin (43.6 per cent yield), triphenylcarbinol,
and a ketone. The analysis of tetraphenyltin and triphenylcarbinol
is given in the experimental section of this report. A ketone was
indicated by the infrared spectrum and by a positive ketone test
with 2, 4-dinitrophenylhydrazine, but could not be isolated. An
absorption chromatographic method is being worked out to sep-
arate these substances.

It is significant to note that acyloxysilanes react with twice the
stoichiometric amount of Grignard reagents to produce a_ poly-
siloxane, (R.SiO), (Lanning and Moore, 1958), and tetrapropoxy-
germane reacts with twice the stoichiometric amount of ethylmag-
nesium bromide to produce triethylgermanium oxide and _tera-
ethylgermanium, while twice the stoichiometric amount of phenyl-
magnesium produces triphenylgermanium oxide (Lanning and
Moore, unpublished). The mechanism for the reaction of tin tetra-
benzoate with phenylmagnesium bromide will be published later.

Experimental preparation of organotin esters. The apparatus
and method used in preparing the organotin esters were very
similar to those used by Lanning and Moore (1958) to prepare
tetracyloxysilanes. The stannic chloride and sodium salts were
chemically pure reagents. Anhydrous benzene was used as the
diluent.

The preparations were carried out by adding .05 mole of stannic
chloride dissolved in 50 ml of benzene dropwise into a slurry of 1.5
times the calculated amount of anhydrous sodium salts dispersed in
300 ml of benzene. The temperature was lowered to 5° by use
of an ice bath. The stannic chloride was added slowly over a one-
hour period. The mixture was stirred during the addition of the
stannic chloride and for 2 1/2 hours thereafter. When the ben-
zene solution gave no test for chloride ion, the sodium chloride
and excess sodium salts were removed. The benzene was removed
from the filtrate under reduced pressure at 0° leaving nearly pure
organotin esters. The analyses and yield of these esters are given
in Table 1.

The tin analysis was carried out by converting the organotin
esters to SnO», using fuming nitric and sulfuric acids. Best results
were obtained by transfering the organotin ester-benzene solution
to a pre-weighed 50 ml Erlenmeyer flask, removing the benzene
under reduced pressure, and analyzing the organotin ester for tin
in the same flask.

Moore AND LANNING: Organotin Esters 75

TABLE, I

Analysis Of Organotin Esters

Tin %
Compound Yield M.p. Calcd. Found
(C,H,CO,),Sn 83.2 25.41 25.47
(Gali-CO;),Sn 74.6 118 dec 19.68 19.53

The infrared spectra of organotin esters and the products from
tin tetrabenzoate with phenylmagnesium bromide were obtained
with a Perkin-Elmer, model 137, Infrared Spectrometer. The or-
ganotin esters were determined from a benzene solution using
matched cells. All the other products were made into pellets with
potassium bromide.

Reaction of tin tetrabenzoate with phenylmagnesium bromide.
A benzene solution containing 15 g of tin tetrabenzoate was added
through a dropping funnel into twice the stoichiometric amount of
a Grignard reagent prepared from bromobenzene in the usual
manner. The mixture was stirred mechanically and maintained at
the boiling point for one hour after the addition was complete.
The Grignard complex was hydrolyzed in an ammonium chloride
solution containing ice. Some hydrochloric acid was added after-
wards to react with the magnesium halide precipitate.

A large amount of white needles and a small amount of yellow
solid were formed. This material was collected. The yellow solid
was removed with forceps, and the white needles were washed
with petroleum ether. The analysis proved the white needles
to be tetraphenyltin, 2.3 g (43.6%), m.p. 238°, lit .226°. Anal. caled.
HOmGonklegons ©. 67.49: H, 4:72: Sn, 27.29. Found: C, 67.88: H,
4.84; Sn, 27.25. The carbon, hydrogen, and tin analyses were de-
termined by Schwarzkoff Microanalytical Laboratory, Woodside
77, New York. The infrared spectrum was identical to the spec-
trum for tetraphenyltin, which was supplied free by the Peninsular
Chemresearch, Inc.

The organic layer was separated and dried with Drierite. The
benzene-ether solution was removed under pressure leaving a
waxy product. The infrared spectrum of this product indicated
a ketone, alcohol, and tetraphenyltin. Triphenylcarbinol was re-

76 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

moved by extracting this product with ethanol and when recrystal-
lized four times from ethanol gave a m.p. 160-1°, lit. 161-2°.
The infrared spectrum of triphenylcarbinol was identical to the
authentic material, and a mixture melting point with the authentic
material was undepressed. An ethanol solution of the crude prod-
uct gave a positive ketone test with 2, 4-dinitrophenylhydrazine,
but could not be isolated, probably because the ketone was not
produced in sufficient quantity to identify.

ACKNOWLEDGMENTS

The authors are indebted to Hampton Junior College Chem-
istry Club and the National Science Foundation for financial sup-
port of this investigation. They also wish to express their appre-
ciation to Dr. Mary Guy of the University of Florida for the in-
frared spectra, and to Tommy Bright, student at Hampton Junior
College, for his cooperation.

LITERATURE CITED

AnpersonN, H. H. 1964. Volatile n-butyltin and phenyltin tricarboxylates.
Organotin oxymonocarboxylates. Inorg. Chem., vol. 3, pp. 912-914.

HeNpERSON, A., AND A. D. Hotiiway. 1965. The reactions of tetravinyltin
with carboxylic acids: Properties of tin II tetracarboxylates. Jour.
Organometal Chem., vol. 4, pp. 377-381.

Lanninc, F., C., AND M. Moore. 1958. Acyloxysilanes and their reaction
with Grignard reagents. Jour. Org. Chem., vol. 23, pp. 288-291.

Scumipt, H., C. BLoHn, AND’ G. JANDER. 1947. Solvolysis reactions in ace-
tic anhydrides. The preparation of germanium and tin tetracetate.
Agnew Chem., vol. A59, p. 233-237.

Department of Chemistry, Hampton Junior College, Ocala,
Florida (present address: Central Florida Junior College, Ocala);

Kansas State University, Manhattan, Kansas.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

A New Monogenetic Trematode from Georgia
CHARLES E. PRICE

Georcia is a fruitful area for the study of monogenetic trema-
todes, as the state possesses an impressive array of teleost fishes
that serve as hosts. The writer has thus far described new species
of Dactylogyrus, Gyrodactylus, and Cleidodiscus, and the paper
here presented represents the first account of Actinocleidus from
this state.

Actinocleidus is one of 46 genera belonging to the subfamily
Ancyrocephalinae of the family Dactylogyridae. Eight of these
have members in the North American parasite fauna, totaling
approximately 105 species. The new Actinocleidus brings the
total species of this genus to 24.

MATERIALS AND METHODS

A single host specimen of Lepomis auritus (Linnaeus) was ob-
tained from Mr. Emory Milner from his branch near Hollanville,
Georgia, to whom the author wishes to express his sincere thanks.
The branchial material and recovered parasites were treated as
described by Price and Mizelle (1964), and measurements were
performed as outlined by Mizelle and Klucka (1953). Measure-
ments and illustrations were made microscopically with the aid of
a filar micrometer ocular and a camera lucida, respectively. All
measurements are given in microns.

Actinocleidus georgiensis sp. n.

Host and locality. Lepomis auritus (Linnaeus); Milner’s Branch,
three miles southeast of Hollanville, Georgia.

Number of Specimens Studied. Two.

Types. Holotype deposited in the Helminthological Collection,
U. S. National Museum, Washington, D. C. Paratype in author's
collection.

Description. A dactylogyrid of moderate size, provided with
a thin, smooth cuticle devoid of scales or spines. Body length 334,
greatest width of body 55, near midlength. No anterior cephalic
lobes formed; vestigial lateral cephalic lobes present.

Two pairs of eyespots, members of posterior pair much larger
and slightly closer together than members of other pair. A few

78 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

comprising eyespot granules scattered in cephalic region. Head
organs not readily observed; apparently three to five pairs. Pedun-
cle short and narrow, setting the haptor off distinctly from body
proper. Haptor somewhat irregular in outline, two finger-like lobes
of tissue projecting posteriorly from the haptor, accompanying the
anchors. Length of haptor 51, width 74.

Two pairs of anchors, all members quite similar (Figs. 1 and 2),
Each anchor composed of (1) a solid base with prominent deep
and superficial roots, (2) a solid shaft connected to (3) a solid point.
Length of anterior anchor 35, width of base 13; length of posterior
anchor 34, width of base 14. Anchor wings prominent, arising
in a depression on lower portion of base and inserting near junc-
tion of shaft and point.

0.05 mm

=

Fig. 1. Actinocleidus georgiensis sp. n. 1, anterior anchor; 2, posterior
anchor; 3, anterior bar; 4, posterior bar; 5, 6, haptoral hooks; 7, cirri; 8,
accessory pieces.

Two haptoral bars, articulated to each other near their mid-
points, the anterior bar much larger than the posterior one (Figs.
3 and 4). Lateral halves of anterior bar joined together by a cen-
tral portion measuring about one-fifth as wide as the adjoining
lateral halves. Length of anterior bar 43. Posterior bar with
prominent articulating surfaces; length of posterior bar 26.

Haptoral hooks 14 (7 pairs), similar in shape and essentially in
accord with the hook arrangement described by Mizelle and Crane
(1964). Hooks similar in length, but some pairs are of heavier con-
struction than others (Figs. 5 and 6). Each hook composed of (1)
a solid elliptical base well differentiated from (2) a solid shaft,
and (3) a sickle-shaped termination provided with an opposable
piece. A posteriorly projecting structure extends from a point
opposite the opposable piece for a distance of ca. one-half the

Prick: New Monogenetic Trematode 79

Weneth ot the shaft. Hook lengths: nos. 1, 3, and 6—15; nos. 2, 4,
and 5—14; no. 7—16.

Copulatory complex consists of a cirrus and accessory piece
(Figs. 7 and 8). Cirrus arises from an expanded base, the tube
strongly curved and tapering to a sharp termination. The acces-
sory piece, articulated to cirrus base, begins its course as a sclero-
tized rod and is bifid distally. One ramus heavy and with a botuli-
form ending, the other ramus less prominent. Testis postovarian.
Prostate large, folded, of a yellowish color, and filled with small
discrete granules. Vas deferens somewhat undulate, not looped
around intestinal limb. Vagina a chitinized tube, opening laterally
near left body margin; vagina associated with a well-defined sem-
inal receptacle.

Vitellaria poorly developed, with little tendency to form lateral
bands. Intestinal crura confluent posteriorly.

TAXONOMIC CONSIDERATIONS

The species apparently most closely related to the present form
is A. subtriangularis, described by Mizelle and Jaskoski (1942) from
the gills of Lepomis miniatus. Several sclerotized parts are quite
similar in these two parasites, including the haptoral hooks. The
task of comparing these species is made difficult by the apparent
significant degree of distortion of the single specimen used in the
description of A. subtriangularis but an adequate degree of dis-
similarity exists in morphology of accessory pieces (especially in
the manner of joining to cirrus bases), haptoral bars, and vagina
to set the two apart.

ACKNOWLEDGMENT

This work was sponsored by the Faculty Research Fund of
the Woman's College of Georgia.

LITERATURE CITED

MrzeLie, J. D., anp J. W. Crane. 1964. Studies on monogenetic trema-
todes. XXIII. Gill parasites of Micropterus salmoides (Lacepede)
from a California pond. Trans. Amer. Micr. Soc., vol. 83, pp. 343-348.

MizELLE, J. D., AND B. J. JAskosxi. 1942. Studies on monogenetic trema-
todes. VIII. Tetraonchinae infesting Lepomis miniatus Jordan. Amer.
Midland Nat., vol. 27, ppv. 145-153.

80 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MiIzeELLE, J. D., and A. R. Kiucka. 1953. Studies on monogenetic trema-
todes. XIV. Dactylogyridae from Wisconsin fishes. Amer, Midland
Nat., vol. 49, pp. 720-733.

Price, C. E., AND J. D. Mizetir. 1964. Studies on monogenetic trematodes.
XXVI. Dactylogyridae from California with the proposal of a new
genus, Pellucidhaptor. Jour. Parasit., vol. 50, pp. 572-578.

The Woman's College of Georgia, Milledgeville, Georgia.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

Barnacles of the Northeastern Gulf of Mexico
Harry W. WELLS

In the course of research on fouling organisms in the north-
eastern Gulf of Mexico, several species of barnacles (Cirripedia)
have been collected which had not been reported previously from
these waters. As records have accumulated since the study be-
gan in 1961, the gaps in our knowledge of this important group
have become more impressive and more inviting to the attention
of the student of marine biology. Because barnacles grow in
great abundance on most structures and require control measures
costing many millions of dollars annually, they are usually re-
garded as the most important component of the fouling com-
munity. They are well represented in the northeastern Gulf of
Mexico, both in quantity and in numbers of species.

In this report are listed the species of barnacles from inshore
waters of the northeastern Gulf of Mexico with annotated remarks
on their collection and distribution. These records are presented
here to augment our knowledge of the barnacle fauna, and to
alert students and investigators in marine biology to their pres-
ence. A large number exhibit relatively close associations with
specific substrates, particularly with other marine animals, and
provide interesting subjects for research on their symbiotic ex-
istence and related adaptations of morphology, physiology, and
life history. Although these barnacle species are often segregated
among their respective preferred substrates or hosts in juvenile
and adult stages, the larval stages (nauplius and cypris larvae)
occur together with other plankton, and contribute to the great
numbers of larval plankton in the sea. Consequently, a thorough
analysis of the planktonic stages would depend on a recognition
of the numerous species of barnacles present in our waters.

A number of new records are included which extend known
geographic ranges from other areas and from the southern part
of the Gulf of Mexico, and which contribute to our knowledge
of the local fauna and to the zoogeography of the Gulf.

MATERIALS AND METHODS

Collections of barnacles were made principally in the vicinity
of the Florida State University Marine Laboratory at Alligator
Harbor, Franklin County; and in the vicinity of the dredged en-

82 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

trance to St. Andrews Bay, near Panama City, Bay County, Florida,
approximately 90 miles west of the laboratory.

This part of the Gulf coast is dominated by sandy beaches and
low marsh-lands, generally unsuited for attachment of barnacles.
However, hard substrates available in this area include shell ma-
terial, and such man-made structures as jetties, pilings, old boats,
buoys, and bridge piers, as well as several hard bottom rock out-
crops a short distance offshore. These structures present habitats
suitable for a number of sessile marine invertebrates, including
several common species of barnacles. The shells of certain inverte-
brates and the leathery skin of sea turtles present satisfactory at-
tachment sites for a number of more selective species, several of
which appear to be restricted to such symbiotic associations with
other animals. Additional species are highly modified parasitic
forms, penetrating into the tissues of certain species of crabs.
Collections from these several habitats constitute the basis for this
report. For identification, the specimens were examined under a
stereoscopic microscope, and where necessary, their mouthparts and
cirri were examined by means of a compound microscope.

Identifications were made primarily with the aid of Pilsbry’s
monographic treatment of the American species (Pilsbry, 1907,
1916), which is strongly recommended for illustrations and de-
scriptions of most species recorded here. Previous records of bar-
nacle distribution in the Gulf of Mexico have been tabulated and
analyzed by Henry (1954, 1959), whose studies have served as
valuable points of reference. More recent records have been
reported from the Panama City area by Hulings (1961).

An unpublished mimeographed check-list (R. W. Menzel, edi-
tor) and a modest reference collection of marine animals of the
immediate vicinity are provided by the Florida State University
Marine Laboratory for the use of students and investigators. The
check-list, containing nine barnacle species identified by Charles
Yentsch, has provided a point of departure for this study.

In the following systematic treatment, collection data are pro-
vided for at least one occurrence of each species examined. No
attempt was made to account for every collection of the more
common species. Available records from the literature of bar-
nacles from the northeastern Gulf are included in this account.
Representative specimens will be deposited in the collections of
the United States National Museum.

Weis: Barnacles of Gulf of Mexico 83

Order THORACICA

Suborder Balanomorpha
Family BaLanmaE: Subfamily BALANINAE
1. Balanus amphitrite amphitrite Darwin

From rock jetties at the entrance to St. Andrews Bay, near Pan-
ama City, Bay Co., 6 November 1964. This species has been re-
corded from St. Marks, Wakulla Co., and locations on the Florida
West coast by Henry (1959) as B. a. denticulata.

2. Balanus calidus Pilsbry

From shells collected on hard bottom 3 miles SE of Dog Island
(off Carrabelle, Fla.), at 20-30 feet, 29 September 1962; 29 October
1962; 23 November 1962; 16 May 1963; 19 June 1965; from other
barnacles (Chelonibia testudinaria) attached to a loggerhead turtle
(Caretta caretta) collected near Bald Point, 2 May 1965; and from
shells of the calico scallop (Aequipecten gibbus) from off Cape
St. George, 2 October 1962. Apparently this common species
attaches to a variety of substrates in waters of high salinity
(> 30 0/oo). It has been recorded from the Panama City area by
Hulings (1961).

3. Balanus declivus Darwin

From loggerhead sponges (Spheciospongia vesparia) collected
at 6 feet depth at St. Teresa, Franklin Co., 10 February 1962 and
16 May 1964. Several specimens were completely imbedded in
the sponge tissue, with only the aperture remaining visible at the
bottom of a shallow pit. The long, curved rostral plate typical of
this species provides a large surface available for sponge attach-
ment. This barnacle is characteristically imbedded in sponges.
It possesses a special series of rasping spines on the fourth cirri
which would assist in retarding overgrowth by tissues of the host
sponge. This species has not been reported previously from this
part of the Gulf, previous records being those of Pilsbry (1916)
from off Cape Sable in southern Florida, and of Henry (1954)
from Texas.

84 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

4, Balanus eburneus Gould

From rock jetties at the entrance to St. Andrews Bay, near
Panama City, 6 November 1964; and from pilings at St. Teresa,
25 May 1964, by W. Stewart. This common species occurs on a
wide variety of substrates, and is particularly abundant near the
low tide line and in estuarine areas.

5. Balanus galeatus (Linné)

From gorgonians (Leptogorgia virgulata) collected at St. Teresa,
16 May 1964; at the entrance to St. Andrews Bay, near Panama
City, 2 July 1965 (by D. Flescher); at 3 miles SE of Dog Island,
16 May 1963. This species occurs on whip corals (gorgonians),
particularly on the denuded stems where the barnacles may be-
come encrusted by regenerative growth of the gorgonians’ outer
soft layer (coenenchyme). As the barnacle develops, changes in
its growth form tend to elevate the barnacle’s aperture above the
encrustation that may be encroaching upon its basal attachment.

6. Balanus improvisus Darwin

From rock jetties at the entrance to St. Andrews Bay, near Pan-
ama City, 6 November 1964; and from pilings at St. Teresa, 25 May
1965, by W. Stewart. This very abundant species occurs on many
substrates in this area, particularly on pilings and oyster shells.

7. Balanus trigonus Darwin

From the compartments of another barnacle (Chelonibia caret-
ta) on the carapace of a loggerhead turtle (Caretta caretta) off
Alligator Harbor, 22 April 1964. Pilsbry (1916) recorded this
species from sponges; on the third cirri it possesses special spines
that can break sponge tissue from its aperture. Henry (1954)
recorded a single occurrence from the Gulf of Mexico without
any more detailed locality data, but with a notation that it was
associated with deep water crabs or palinurids. As a result of
recent discoveries of this species in the Miami area (Moore and
McPherson, 1963) and on the North Carolina coast (Wells et al.,
1964; Ross et al., 1964), it has been suggested that it may be ex-
tending its geographic range. Transport on sea turtles could con-
tribute materially to its passive dispersal to new areas. However,

Weis: Barnacles of Gulf of Mexico 85

its additional occurrence off Georgia and on the northeast and
west coasts of Florida (personal collections) suggests that it is more
widely distributed along the coast of the southeastern United
States than published records indicate.

8. Balanus venustus niveus Darwin

From rock jetties at the entrance to St. Andrews Bay, near
Panama City, 6 November 1964; from dead shells, St. Teresa, 26
September 1964; and from shells of the calico scallop (Aequipecten
gibbus) from off Cape St. George, 2 October 1962. This subspecies
is abundant and occurs on a wide variety of hard substrates. It
has been recorded previously from the Atlantic and Gulf coasts
as B. amphitrite niveus (Pilsbry, 1916, Henry, 1959), but Harding’s
nomenclature, based on the Darwin types (Harding, 1962), is fol-
lowed here. The colored form described from Sarasota Bay by
Pilsbry (1916, p. 94) is common offshore.

9. Balanus venustus obscurus Darwin

From loggerhead turtles (Caretta caretta) collected near St.
Teresa, 22 April 1964, and near Bald Point, Franklin Co., 2 May
1965. This subspecies has not been reported previously from the
Gulf of Mexico, nor from mainland North American waters, but
it is known from Jamaica (Harding, 1962).

10. Acasta cyathus Darwin

From a sponge (Ircinia fasciculata) from Lighthouse Point,
Franklin Co., 10 March 1962. This species is a common inhab-
itant of certain sponges in the northeastern Gulf: tube sponges
(Callyspongia vaginalis), stinker sponges (Ircinia fasciculata), vase
sponges (Ircinia campana), and sponges of the genus Verongia.
In each case, the barnacle is anchored within the sponge tissue
by recurved spines on the compartmental plates. It possesses
stout spines on the fourth cirri which serve to prevent occlusion of
the barnacle aperture by lateral growth of the host sponge. The
genus Acasta is characterized by these morphological adaptations
associated with life in sponges (Pilsbry, 1916). Although Henry
(1954) recorded this species only from the southern part of the
Gulf, it is contained in the checklist of the Florida State University
Marine Laboratory.

36 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ll. Pyrgoma floridanum Pilsbry

From coral (Siderastrea radians) from St. Teresa, Franklin Co..,
at 2-10 feet depth, 4 November 1961, 10 February 1962, 16 May
1964. This species commonly occurs on and imbedded within
the coral mass, usually marked by swollen, gall-like knobs on the
corallum. One coral colony 11 cm in diameter bore 8 living speci-
mens at its surface, and the empty plates of several other spec-
imens completely imbedded in the interior of the corallum. Pils-
bry (1931) described this species from specimens associated with
another coral (Manicina areolata) from off Tarpon Springs, approx-
imately 160 miles SE of St. Teresa. Members of this genus are
typically associated with corals (Pilsbry, 1916).

Subfamily CHELONIBIINAE
12. Chelonibia caretta (Spengler)

From the carapace of a loggerhead turtle (Caretta caretta) at
St. Teresa, Franklin Co., 22 April 1964. This species has not been
reported from the Gulf of Mexico, but has been reported previ-
ously from New Jersey and the West Indies (Pilsbry, 1916).

13. Chelonibia manati lobatibasis Pilsbry

From a loggerhead turtle (Caretta caretta) from near Bald
Point, Franklin Co., 2 May 1965. The specimen was partially
imbedded, the basal lobes of the parietes penetrating 3-4 mm into
the skin of the turtle. Pilsbry (1916) described this subspecies
from a loggerhead turtle collected at Osprey, Florida, approxi-
mately 50 miles S of Tampa. It apparently has been known only
from his original discovery.

14. Chelonibia patula (Ranzani)

From a blue crab (Callinectes sapidus) collected in Alligator
Harbor, 11 October 1964. This species is typically associated with
arthropods; in this area it is common on the carapace of older,
mature blue crabs and on “horseshoe crabs” (Limulus polyphemus).

15. Chelonibia testudinaria (Linne)

From loggerhead turtles (Caretta caretta) at St. Teresa, 22 April
1964, and near Bald Point, Franklin Co., 2 May 1965. This species

Wetts: Barnacles of Gulf of Mexico 87

is characteristically found on the carapace of the loggerhead tur-
tle; it frequently supports a secondary layer of encrusting organ-
isms. Although it has been recorded from several locations in the
Gulf (Pilsbry, 1916), the nearest recorded locality for this species
has been Pensacola, 100 miles W of Panama City.

16. Platylepas hexastylos (Fabricius)

From a loggerhead turtle (Caretta caretta) near Bald Point,
2 May 1965. These barnacles were depressed and partially im-
bedded in the skin, producing a convex basal surface. Struc-
turally, this species is well adapted to life on sea turtles; each
compartmental plate is reinforced by a calcareous strut. Pilsbry
(1916) has recorded this species from a loggerhead turtle from
Osprey, approximately 50 miles S of Tampa. He also described
a variety (or subspecies), Platylepas hexastylos ichthyophila, from
a garfish caught in brackish water in Hernando County, approxi-
mately 40 miles N of Tampa.

17. Platylepas hexastylos variety

From a loggerhead turtle near Bald Point, Franklin Co., 2 May
1965. More than twenty specimens similar to the form described
and illustrated by Pilsbry (1916: p. 287; Pl. 67, fig. 4) as a variety
from Sicily. This variety is not depressed but has a cylindrical
form, with lobate bases of the parietes projecting downward into
the turtle’s skin.

18. Stomatolepas praegustator Pilsbry

From a loggerhead turtle (Caretta caretta) near Bald Point,
Franklin Co., 2 May 1965. Forty specimens were found in the
mouth cavity and upper gullet, where they were partially imbed-
ded. An additional group of 55 specimens were removed from
the soft dermis of the dorsal aspect of the hind legs, where they
were also partially imbedded. Pilsbry (1910) described this species
from the gullet of a loggerhead turtle from the Tortugas. A second
collection, made by W. L. Schmitt from the tongue of a loggerhead
turtle at the same locality, is contained in the U. S. National Mu-
seum (usNM No. 79178). The present account is apparently but
the third record of this species, and the first report of its occur-

88 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

rence on the outside of a turtle’s body. The beautifully sculptured
compartmental plates provide anchorage in the host’s tissue.

Family CHTHAMALIDAE

19. Chthamalus fragilis (Darwin)

From rock jetties at the entrance to St. Andrews Bay, near
Panama City, Bay Co., 12 October 1963 and 6 November 1964;
and from Alligator Harbor, 22 April 1964. This species is widely
distributed in the northeastern Gulf where it is common on suitable
substrates in the intertidal zone, on wood, stone, metal, and fre-
quently on salt marsh grass (Spartina patens). Is usually occurs
higher in the intertidal zone than any other macroscopic sessile
marine invertebrate.

20. Chthamalus stellatus bisinuatus Pilsbry

From rock jetties at the entrance to St. Andrews Bay, near
Panama City, 12 October 1963, 30 April 1964, and 8 May 1965,
by W. Stewart. Numerous specimens were widely distributed in
the intertidal zone interspersed among specimens of Chthamalus
fragilis, which outnumber them. This subspecies has been reported
previously from Brazil (Pilsbry, 1916; Oliveira, 1941) and West
Africa (Stubbings, 1961).

Suborder Lepadomorpha
Family LEPADIDAE

21. Lepas anatifera Linné

From shells of purple sea snails (Janthina species) collected at
St. Andrews State Park, near Panama City, Bay Co., 28 April 1963,
and from driftwood near Panama City, 11 April 1965, by H. Mat-
thews. This species is characteristically attached to driftwood
and other floating substrates.

22. Lepas pectinata Spengler

From floating Gulfweed (Sargassum species) and shells of pur-
ple sea snails (Janthina species) washed ashore at St. Andrews

We tts: Barnacles of Gulf of Mexico 89

State Park, near Panama City, 28 April 1963. This species is also
characteristically associated with floating substrates.

Family TRmLASMATIDAE

23. Octolasmis hoeki (Stebbing)

Reported as a commensal in the branchial chamber of an oxy-
stomatous crab (Calappa flammea) from a nearshore area near
Panama City (Hulings, 1961).

24. Octolasmis lowei (Darwin)

From the branchial chambers of blue crabs (Callinectes sapidus)
collected at Alligator Harbor, 10 July 1965, by T. Borkowski; and
the branchial chamber of a stone crab (Menippe mercenaria) at
St. Teresa, 30 August 1965. This common species is a commensal
in the branchial chambers of crabs, particularly in older, mature
blue crabs. It has been reported previously from the Alligator
Harbor area by Pearse (1952) as O. mulleri, which is recognized
today as a synonym of O. lowei (Causey, 1961).

Family ALEPADIDAE

25. Conchoderma virgatum (Spengler)

From the dorsal fin on an orange filefish (Alutera schoepft) col-
lected 31 December 1964, by H. Matthews; and from the head
of a cowfish (Lactophrys tricornis) collected 9 April 1966, by T.
Scanland, both at the entrance to St. Andrews Bay, near Panama
City. Although not recorded from the Gulf of Mexico, this species
has been reported elsewhere from seaweeds, sea turtles, and large
fish, as well as from the bottoms of ships (Pilsbry, 1907).

Family SCALPELLIDAE

26. Scalpellum arietinum Pilsbry

From the calico scallop (Aequipecten gibbus) beds off Cape
St. George, 2 October 1962; and from materials dredged 15 miles
S of Alligator Point, 14 November 1965. This species has been
reported from a nearshore area near Panama City (Hulings, 1961).

90 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Order RHIZOCEPHALA
Family SACCULINIDAE

27. Sacculina pustulata Boschma

Reported as a parasite on a spider crab (Hemus cristulipes)
collected 10 miles SE of Alligator Point (Reinhard, 1955).

28. Loxocephalus texanus Boschma

From blue crabs (Callinectes sapidus) collected in Alligator
Harbor, 11 October 1964. This species is not uncommon as a par-
asite of blue crabs in this area.

29. Loxothylacus panopaei (Gissler)

From a hairy crab (Pilumnus sayi) from 15 miles S of Alligator
Harbor, at 60 feet depth, 14 November 1965 (by T. Scanland):
and from mud crabs (Neopanope packardi) from St. Andrews Bay,
near Panama City, 20 September 1957 (Oceanographic Institute
collection). This species is a parasite of mud crabs (Xanthidae).
and has been reported in the Gulf from Tampa and from Engle-
wood, approximately 75 miles S of Tampa, and in Louisiana and
Texas (Reinhard and Reischman, 1958).

30. Peltogaster species

From hermit crabs (Pagurus longicarpus) from Live Oak Island,
Wakulla Co., 3 November 1962 and 21 November 1963. To my
knowledge, this sacculinid parasite of hermit crabs has not been
reported previously from the Gulf of Mexico.

Order ACROTHORACICA
Family KocHLORINDAE

31. Kochlorine floridana Wells and Tomlinson

From shells of the mossy ark (Arca imbricata = A. umbonata),
the turkey wing ark (Arca zebra), and a variety of other molluscs,
and from calcareous bryozoans, coral, barnacle compartments
(Balanus calidus), tubes of serpulid polychaetes, and calcareous red
algae; 3-5 miles SE of Dog Island, 30-35 feet depth, 29 September
1962, 29 October 1962, 23 November 1962, 16 May 1963 (Wells and

We tts: Barnacles of Gulf of Mexico 91

Tomlinson, 1966), and 19 June 1965. Also from shells of the calico
scallop (Aequipecten gibbus) from off Cape St. George, 60 feet
depth, 4 October 1962; and from shells of the giant eastern murex
(Murex fulvescens) from rock jetties at the entrance to St. Andrews
Bay, near Panama City, 5 November 1965, by G. Bertrand.

GEOGRAPHIC CONSIDERATIONS

Of the 31 taxa included in this report, a high percentage are
newly recorded from the northeastern Gulf of Mexico. Six taxa
have not been recorded previously from the Gulf: Balanus ven-
ustus obscurus, Chelonibia caretta, Platylepas hexastylos variety,
Chthamalus stellatus bisinuatus, Conchoderma virgatum, and Pelto-
gaster species. Present records of the following species extend their
known distribution 400 miles northward into the shallow waters
of western Florida: Balanus declivus, Acasta cyathus, and Stomato-
lepas praegustator. Records of four species extend their known
geographic range at least 200 miles northward along the Florida
west coast; and additional records represent new localities for
species that have been collected at other localities around the
Gulf of Mexico. These new records are not to be taken as evi-
dence of recent movement of many barnacle species into the
study area. Instead, they reflect an increasing interest in the local
fauna by marine biologists. It is to be expected that further re-
search on this group will extend the known distribution of sev-
eral of these barnacle species even further.

This barnacle fauna includes a number of widespread, warm-
temperature species that are also distributed along the Atlantic
coast of the United States as far north as New Jersey or Massa-
chusetts: Balanus venustus niveus, B. eburneus, B. improvisus,
Chelonibia patula, and Chthamalus fragilis. The barnacles most
often found in estuaries and inshore areas of brackish water are
these warm-temperate species. Most exhibit a continuous distri-
bution around the tip of peninsular Florida through waters of
distinctly tropical characteristics (Moore and Frue, 1959; Stephen-
son and Stephenson, 1950). For an analysis of geographic affini-
ties, Conchoderma virgatum and the Lepas species, which are
nearly cosmopolitan, should be added to these widespread species.

Species of tropical or subtropical distribution which do not
extend north of Cape Canaveral or Cape Hatteras on the east coast
far outnumber the widespread species. They include Balanus

92 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

amphitrite amphitrite, B. calidus, B. declivus, B. galeatus, B. tri-
gonus, B. venustus obscurus, Acasta cyathus, Chthamalus stellatus
bisinuatus, Octolasmis species, Scalpellum arietinum, Saculina pus-
tulata, Loxothylacus manopaei, and Kochlorine floridana. The spe-
cies associated with sea turtles (Platylepas, Stomatolepas, and Che-
lonibia species) should be included here, for they are usually con-
sidered to be tropical although subject to being transported by
their hosts to extratropical shores. This sizeable group of species
is an indication of the importance of the tropical element in the
fauna of the northeastern Gulf. These barnacles generally are
found in offshore or higher salinity waters in this area.

The case of Chthamalus stellatus bisinuatus warrants special
comment, for this tropical form has been discovered in numbers
near Panama City, but to our knowledge has not been recorded
previously from the Gulf of Mexico. Although another subspecies,
C. s. angustitergum, has been reported from the Florida Keys (Pils-
bry, 1916; Stephenson and Stephenson, 1950), C. s. bisinuatus has
been recorded previously only from Brazil (Pilsbry, 1916; Oliveira,
1941) and West Africa (Stubbings, 1961). However, C. s. bisinuatus
evidently has a wide distribution in the eastern Gulf of Mexico,
for it also occurs in suitable habitats on the southern west coast
of Florida (personal collections). The abscence of previous records
from this region probably is due to the inconspicuous appearance
of this species and its apparent preference for intertidally situated
hard substrates. Henry (1954) notes that members of the genus
Chthamalus are often overlooked by collectors because of their
small size and inconspicuous form. Alternatively, modern man
has created many suitable habitats for this species by his con-
struction of jetties, bridge pilings, and navigational aids, and this
species may enjoy a wider distribution as a consequence. Its
presence in the Panama City area might have resulted originally
from its transport on the hulls of ships that use this port.

In summary, the barnacle fauna of the northeastern Gulf con-
sists of a mixture of warm temperate and tropical species.

SUBSTRATE

Of the barnacles recorded from the northeastern Gulf, a ma-
jority (17) exhibit relatively specific symbiotic associations with
particular substrates or hosts. One large group attaches to the
exposed surfaces of sea turtles, particularly to the carapace, but

Wetts: Barnacles of Gulf of Mexico 93

also to the skin of the head, neck, legs, and ventral surfaces:
Chelonibia caretta, C. manati lobatibasis, C. testudinaria, Platy-
lepas hexastylos, P. hexastylos variety, and Stomatolepas praegus-
tator (which also attaches in the posterior part of the turtle’s mouth
cavity). Heavy accumulations of Chelonibia species may occur on
the turtle’s carapace, especially near the posterolateral margins,
providing suitable surfaces for attachment by a secondary layer
of fouling organisms. Such secondary encrustations may include
barnacles of less specific substrate requirements, hydroids, ane-
mones, bryozoans, serpulid polychaetes, molluscs, and algae Sa-
bellid and spionid annelids and molluscs which burrow into the
calcareous plates of the Chelonibia contribute further to the fauna
of the sea turtle shell.

Another large group exhibiting specific substrate requirements
is comprised of the barnacles associated with crabs: Chelonibia
patula, which typically attaches to the carapace; Octolasmis hoeki
and O. lowei, which attach to crabs’ gills; and the parasitic species
Sacculina pustulata, Loxocephalus texanus, Loxothylacus panopaei,
and Peltogaster species, which invade the inner tissues of their
hosts and extend to the outside as sacs attached to the abdomens
of their hosts. Other barnacles are characteristically imbedded in
sponges (Balanus declivus and Acasta cyathus) or in coelenterates
(Balanus galeatus and Pyrgoma floridanum).

Another species that exhibits a relatively specific affinity for
a special substrate is Kochlorine floridana, which perforates only
calcareous materials.

The remaining 13 species, including the barnacles most likely
to be seen by biologists or laymen, are relatively non-specific in
their choice of substrate and have been recorded from a wide
variety of objects. Generally, these species attach to non-living
structures, but they may also occur wherever living organisms pre-
sent a suitable firm substrate. Thus, the shells of arthropods and
molluscs may provide attachment for such barnacles, whether the
arthropod or mollusc is alive or not. Balanus amphitrite amphi-
trite, B. calidus, B. eburneus, B. improvisus, B. trigonus, B. venus-
tus niveus, B. c. obscurus, Chthamalus fragilis, C. stellatus bisinu-
atus, and Scalpellum arietinum are species of the northeastern Gulf
that exhibit such non-specificity. Addititonal examples of bar-
nacles of non-specific habits would include Lepas anatifera, L.
pectinata, and Conchoderma virgatum which however, are pri-

94 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

marily associated with floating objects, whether living or inani-
mate.
SUMMARY

Thirty-one taxa of barnacles are recorded from inshore waters
of the northeastern Gulf of Mexico, including 16 which are newly
reported from this area. Six taxa are newly recorded for the Gulf
of Mexico, including Balanus venustus obscurus, Chthamalus stel-
latus bisinuatus, Chelonibia caretta, Platylepas hexastylos variety,
Conchoderma virgatum, and Peltogaster species. This barnacle
fauna consists of a mixture of warm-temperate and tropical species.
Many species exhibiting specific symbiotic relationships with sea
turtles, with crabs, and with other marine organisms are repre-
sented.

ACKNOWLEDGMENTS

I wish to express my sincere appreciation to the students who
have contributed specimens of interesting species, and to Mary
Jane Wells who has ably assisted in the collection and identification
of the materials included in this report. This study has been sup-
ported by National Science Foundation grant GB-819 to Florida
State University.

LITERATURE CITED

Causey, D. 1961. The barnacle genus Octolasmis in the Gulf of Mexico.
Turtox News, vol. 39, pp. 51-55.

Harpinc, J. P. 1962. Darwin’s type specimens of varieties of Balanus
amphitrite. Bull. Brit. Mus. (Nat. Hist.) Zool., vol. 9, no. 7, pp. 273-
296, 10 pls.

Henry, Dora P. 1954. Cirripedia; the barnacles of the Gulf of Mexico.
In: Gulf of Mexico: Its origin, waters, and marine life. U. S. Fish and
Wildl. Serv., Fishery Bull. 89, vol. 55, pp. 443-446.

———. 1959. The distribution of the amphitrite series of Balanus in North
American waters. In: Marine boring and fouling organisms. Univ.
Washington, Friday Harbor Symposium, pp. 190-203.

Hutines, N. C. 1961. The barnacle and decapod fauna from the nearshore
area of Panama City, Florida. Quart. Jour. Florida Acad. Sci., vol. 24,
pp. 215-222.

Moore, H. B., ano A. C. Frurz. 1959. The settlement and growth of Balanus
improvisus, B. eburneus, and B. amphitrite in the Miami area. Bull.
Mar. Sci. Gulf & Carib., vol. 9, pp. 421-440.

Moore, H. B., ANp B. F. McPuerson. 1963. Colonization of the Miami area
by the barnacle Balanus trigonus Darwin, and a note on its occurrence

Wetts: Barnacles of Gulf of Mexico 95

on the test of an echinoid. Bull. Mar. Sci. Gulf & Carib., vol. 13,
pp. 418-421.

OxiverrA, L. P. H. pe. 1941. Contribucao ao conhecimento dos crustaceos
do Rio de Janeiro sub-ordem “Balanomorpha” (Cirripedia: Thoracica).
Mem. Inst. Oswaldo Cruz, vol. 36, pp. 1-36.

PrarsrE, A. S. 1952. Parasitic crustaceans from Alligator Harbor, Florida.
Quart. Jour. Florida Acad. Sci., vol. 15, pp. 187-248.

Prrspry, H. A. 1907. The barnacles (Cirripedia) contained in the collections
of the U. S. National Museum. Bull. U. S. Nat. Mus., vol. 60, pp.
LEQ Ie pls:

—. 1910. Stomatolepas, a barnacle commensal in the throat of the log-

gerhead turtle. Amer. Nat., vol. 44, pp. 304-306.

1916. The sessile barnacles (Cirripedia) contained in the collections
of the U. S. National Museum, including a monograph of the American
species. Bull. U. S. Nat. Mus., vol. 98, pp. 1-366, 76 pls.

1931. The cirriped genus Pyrgoma in American waters. Proc.
Acad. Nat. Sci., Philadelphia, vol. 88, pp. 81-83.

REINHARD, FE. G. 1955. Some Rhizocephala found on brachyuran crabs in
the West Indian region. Jour. Wash. Acad. Sci., vol. 45, pp. 75-80.

REINHARD, FE. G., AND P. G. Reiscuman. 1958. Variation in Loxothylacus
panopaei (Gissler), a common sacculinid parasite of mud crabs, with
the description of Loxothylacus perarmatus, n. sp. Jour. Parasitol.,
vol. 44, pp. 93-97.

Ross, A., M. J. CeraMe-Vivas, AND L. R. McCuioskry. 1964. New barnacle
records for the North Carolina coast. Crustaceana, vol. 7, pp. 312-313.

STEPHENSON, T. A., AND ANNE STEPHENSON. 1950. Life between tide marks
in North America. I. The Florida Keys. Jour. Ecol., vol. 38, pp. 345-
402, pls. 9-15.

StusBincs, H. G. 1961. Cirripedia Thoracica from tropical West Africa.
Atlantide Report, Copenhagen, No. 6, pp. 7-41.

WELLs, H. W., anv J. T. Tomuinson. 1966. A new burrowing barnacle
from the Western Atlantic. Quart. Jour. Florida Acad. Sci., vol. 29,
Pwo Tes. 1-3.

WeEtus, H. W., Mary JANE WELLS, AND I. E. Gray. 1964. The calico scal-
lop community in North Carolina. Bull. Mar. Sci. Gulf & Carib., vol.
14, pp. 561-593.

Department of Biological Science, Florida State University,
Tallahassee.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

Minima of Eclipsing Binaries VV Orionis and Beta Persei
K-Y. CHEN

PHOTOELECTRIC observation of VV Orionis was made on Novem-
ber 20-21, 1965, in blue (effective wave length — 0.425 micron),
B, and in visual (effective wave length — 0.545 micron), V, wave
length regions, using the 12:5-inch reflecting telescope at the Uni-
versity of Florida (Chen and Rekenthaler, 1966). The star BD-1°-
949 was observed for comparison. During the seven hour period,
27 individually observed points were obtained in B and 25 points
in V. The minimum light was determined by Hertzsprung’s
method to have occurred at 2439085.7867 and 2439085.7979 in B
and V, respectively, both values being in heliocentric Julian Date.
The average value then is 2439085.7923.

Beta Persei was observed on November 24-25, 1965, with Pi
Persei as the comparison star. In the five-hour period 35 individ-
ually observed points were obtained in B and 33 in V. The times
of minimum in heliocentric Julian Date were determined to be
2439089.6899 and 2439089.6888 in B and V, respectively. The av-
erage of these values is 2439089.6894.

ACKNOWLEDGMENT
This research was supported in part by the National Aeronau-

tics and Space Administration.

LITERATURE CITED

CuEeN, K-Y., aNnD D. A. REKENTHALER. 1966. Photoelectric photometry of
44i Bootis. Quart. Jour. Florida Acad. Sci., vol. 29, no, 1) pp. I-12:

Department of Physics and Astronomy, University of Florida,
Gainesville, Florida.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

New Records of Bahamian Odonata
Dennis R. PAULSON

Tue dragonfly fauna of the Bahama Islands remained virtually
undocumented until Westfall (1960) listed the Bahamian odonate
specimens in the collection of the American Museum of Natural
History. Vaurie (1952) and Rabb and Hayden (1957) had _ previ-
ously discussed the collecting of some of these specimens in their
general accounts.

I was fortunate in being able to collect dragonflies in the Ba-
hamas during three visits there. These trips allowed me to sample
the faunas of South Bimini (17-19 November 1962), Cat Island (27
November-2 December 1963), and San Salvador (26 December
1963-2 January 1964). In addition, I received collections made by
Dr. Fred G. Thompson on Andros (30 October-1 November 1963)
and by Cecil R. Warren on South and East Bimini (16-22 August
1964). My wife, Mary Lynn, collected a single specimen on Grand
Bahama Island (11 April 1963). Dr. Minter J. Westfall, Jr., called
my attention to a Bahamian specimen of interest in the University
of Florida Collections and allowed me to borrow specimens in his
care. Dr. Oliver Flint permitted me to examine pertinent speci-
mens of Anax longipes in the United States National Museum, and
Dr. Thomas W. Donnelly furnished a specimen and addititonal in-
formation of the same species. The following list includes all
specimens obtained in the above-mentioned collections.

Lestes scalaris Gundlach. car IsLANpD, 1 mi. S Tea Bay, 29
November (2); sAN SALVADOR, 6.6 mi. S Cockburn Town, | January
(3). A mating pair and a single male were observed at a small
pond surrounded by buttonwoods on Cat Island, all two to three
feet above the water. The species was common at the San Salva-
dor locality, and several mating pairs were seen. They perched
low in the dense stand of Phragmites occupying most of the shal-
lower parts of the pond. None was observed in the coppice, where
L. spumarius was often encountered.

The appendages of the four males collected agree in detail with
those figured by Calvert (1909: Pl. I, Figs. 17, 18). They have
been compared with specimens from Cuba, Jamaica, and Puerto
Rico, and agree in all respects, with the exception of one aspect of
the wing venation, which apparently varies geographically. <A

98 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

postnodal index was devised by totalling the postnodals of all four
wings for each specimen. Bahamian specimens have fewer post-
nodals (31-40 in five specimens) than do those from Jamaica (40-48
in seven specimens). A Cuban individual agrees best with those
from the Bahamas (29 postnodals), and one from Puerto Rico is
intermediate (39 postnodals).

L. scalaris has been recorded from Cuba, Puerto Rico (Calvert,
1909), and Hispaniola (Needham, 1941). The present records add
Jamaica (by the courtesy of M. J. Westfall, Jr.) and the Bahamas to
the known range, which is thus the entire Greater Antilles and
Bahamas. It is completely sympatric with the following species.

Lestes spumarius Selys. souTH BrMINt, 16-22 August (1); car
ISLAND, hills above The Bight, 28 November (1); Orange Creek, 1
December (1); SAN SALVADOR, 3.6 mi. S Cockburn Town, 1 January
(3). At the last locality, several mating pairs were seen in low
weeds around a small pond that was completely surrounded by
coppice. Other adults were observed in grass at a pond 6.6 mi. S
Cockburn Town; when disturbed, they flew immediately into the
adjacent low coppice. Only three were found on Cat Island, al-
though many hours were spent in their favored habitat. The
species was rather common in the coppice on South Bimini in
both August and November. On all three islands where I ob-
served spumarius, individuals were seen in the densest forest pres-
ent, in which they hung vertically from small twigs five to seven
feet above the ground. When flushed, they flew only a short dis-
tance (ten to thirty feet) but were so well camouflaged that I usu-
ally lost sight of them until I disturbed them again.

As in L. scalaris, individuals from the Bahamas and Cuba have
fewer postnodals than do those from elsewhere in the Greater An-
tilles. A Cuban specimen has 39, thirteen Bahamian specimens
have 40-47, eleven Jamaican specimens have 45-56, and two His-
paniolan specimens have 45-48 postnodals. In this species a cor-
related variation in size is evident. The hind wings of ten Baha-
mian males vary from 19-21 mm, those of two males from Hispani-
ola and six males from Jamaica from 22-23 mm. No such variation
was noted in scalaris, in which the range of the hind wing length
is 17-19 mm. Larvae and exuviae of Lestes were collected on An-
dros, Cat Island, and San Salvador, but I have been unable thus
far to determine their specific identity.

Argiallagma minutum Selys. Car IstANp, 1 mi. S Tea Bay, 29

PauLson: Odonata of Bahamas 99

November (1); sAN saLvapor, 3 mi. S Cockburn Town, 26 Decem-
ber (1); 3.6 mi. S$. Cockburn Town, 1 January (2). This species was
common at ponds on both islands. Larvae, exuviae, and tenerals
were numerous at shallow marshes with abundant sedge (Eleo-
charis spp.) growth near Tea Bay. Another larva was collected
from a pond 5 mi. NW Fresh Creek, Andros.

Ischnura ramburi Selys. anpros, Andros Town, 31 October
(12); 2.5 mi. SE Andros Town, 31 October (1); car IsLaNnp, 1 mi. S
Tea Bay, 29 November (4); sAN sALvApoR, 1 mi. NE Cockburn
Town, 28 December (1); 6.6 mi. S Cockburn Town, 1 January (1).
I. ramburi was locally common on Cat Island and uncommon on
San Salvador. Only one was seen on South Bimini in November
1962. Large series of larvae were collected on Andros and in a
fresh-water pond south of Tea Bay on Cat Island.

Westfall (1960) reported both “typical ramburi with the dorsum
of segment 9 entirely black, and also the subspecies credula Hagen”
from the Bahamas. I have re-examined most of the specimens
listed by Westfall and categorized them and the ones listed above.
Individuals with more than half of the dorsum of segment nine
black were called “ramburi,’ those with more than half of that
segment blue were called “credula,’ and those with about half
black and half blue were called “intermediate.” Both males and
homochromatic females were included. On South Bimini, Andros,
and New Providence, ramburi is the predominant form (10 of 12
specimens). On Stanyard Cay, San Salvador, Crooked Island,
Great Inagua, and South Caicos, credula is the predominant form
(11 of 13 specimens). Mixed populations are present on Eleuthera
(5 ramburi, 3 intermediate, 1 credula) and Cat Island (3 ramburi,
2 intermediate, 8 credula). Thus a geographic correlation is clear-
ly evident, the northern Bahamas inhabited by the ramburi-type
and the southern islands by the credula-type, with intermediate
populations on the intervening islands.

Credula is the dominant or exclusive form elsewhere in the
West Indies (Calvert, 1901-1908; Klots, 1932; Whitehouse, 1943)
and ramburi the dominant form in Florida (Byers, 1930; and my
examination of over 800 specimens from south Florida), and the
two may be distributed more-or-less in the pattern of allopatric
subspecies in these areas. However, Calvert (1901-1908) reported
much mixing of the two types, and I have taken both throughout
southern Mexico and examined credula from as far north as Sinaloa

100 = QuarTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

and ramburi from as far south as Colima on the Pacific coast and
Guatemala on the Caribbean coast. Considering the extremely
broad area of overlap of the two forms and the common occur-
rence of both chromotypes in far-flung populations, no biogeo-
graphic purpose is served by the formal recognition of credula
as a subspecies, and I consider it a strict synonym of ramburi.

Anomalagrion hastatum Say. SAN SALVADOR, 6.6 mi. S Cockburn
Town, 30 December (1). No others were seen on any of the is-
lands, surprising in light of the abundance and wide distribution
of the species from Westfall’s (1960) records.

Anax junius Drury. anpros, Andros Town, 31 October (1);
CAT ISLAND, 1 mi. S Tea Bay, 29 November (1). A few were seen
on South Bimini at a fresh-water rockpit. Last instar larvae were
very common in another rockpit on the same island and were found
emerging not long after dark on 18 November. Adults were scat-
tered over Cat Island, breeding in the ponds near Tea Bay. Two
females oviposited (unattended by males) at one pond, and a very
young Anax larva was collected there. A half-grown junius larva
was taken in a pond 2 mi. SE Andros Town, Andros.

Anax longipes concolor Brauer. SAN SALVADOR, 1 mi. N Cock-
burn Town, 26 December (1); 6.6 mi. S Cockburn Town, 1 January
(1). A half-grown larva of this species was collected from a small
pond at Knowles, Cat Island. Several males flew over a small
pond south of Cockburn Town, and four or five individuals cruised
over the scrub at Watling’s Castle, the highest hill in the area.
When first seen in the field, these dragonflies were a source of
puzzlement to me, as they looked like no Anax known from the
northern hemisphere. When in the hand, it could be seen that
they were structurally much like A. longipes from Florida, but
smaller and differently colored. Notes on the coloration of the
male in life were taken: frons and thorax lime green, abdominal
segments 1 and 2 same color, 2 porcelain blue distally; 3 to 10
blackish with pale green spots (small transverse ones on anterior
third and larger rhombic ones on posterior third of middle seg-
ments); appendages brown; femora reddish, remainder of legs
black. The female was quite similar, although lacking the blue
on the second segment. Both are fully hardened adults.

Brauer described Anax concolor from Brazil in 1865, and Hagen
(1890) and Martin (1908) considered it a race of A. longipes. Dur-
ing the same period, Kirby (1890) and Muttkowski (1910) listed it

Pautson: Odonata of Bahamas 101

as a full species in their catalogues, the latter mistakenly reporting
it from throughout the known range of longipes. The same authors
were divided on their ideas of the range of longipes, Kirby record-
ing it from only the United States and the other three listing it
as well from Haiti and Mexico, Muttkowski from Jamaica, and
Martin from Brazil. Calvert (1901-1908) did not discuss concolor
but recorded longipes from the United States, Mexico, Brazil, and
Haiti. He then (1909) added the Bahamas to the known range of
the species from a specimen taken on Eleuthera. Thus no real
consistency is evident in the treatment of these forms at the turn
of the century. Since that time, no mention of the relationship
of longipes and concolor has appeared in the literature, and noth-
ing has been said of the biology of concolor.

My specimens from San Salvador agreed with the diagnosis
of the Brazilian concolor rather than the northern longipes, prompt-
ing an analysis of the distribution and relationship of the two forms.
The following specimens were examined: Arkansas, 3; Tennessee,
1; North Carolina, 5; South Carolina, 6; Georgia, 6; Florida, 11;
San Salvador, 2; Dominica, 4; Mexico, 1; Costa Rica, 1; Bolivia, 5;
anal |Binevall, 2

Hind wing lengths of these specimens in millimeters are as
follows: United States, 49-51; San Salvador, 46-48; Dominica,
46.5-49: Mexico, 45; Costa Rica, 46; Bolivia, 45-46; and Brazil, 40-
44. Measurements of the hind femora and tibiae of United States
specimens (respectively 13.5-15 mm and 12-13 mm) do not overlap
those of tropical specimens (respectively 10-13 mm and 9-11 mm).
The abdominal coloration is difficult to assess in many of the dried
specimens. All United States individuals, insofar as can be de-
termined, have reddish abdomens. The male abdomens are un-
patterned, those of the females obscurely marked with paler spots.
Teneral females (two reared from North Carolina) are spotted
exactly as described above for the San Salvador male, except the
spots are blue on segments 3 to 8 and yellowish to greenish on
nine. I have seen no teneral males, but the very conspicuous spot-
ting of the teneral females in longipes, later replaced by red-brown
overall, indicates concolor as the ancestral form. Some specimens
from outside the United States appear to have reddish abdomens,
but this may be an artifact of drying. The abdomens of two males
which I collected (San Salvador and Veracruz, Mexico) were bright-
ly marked in life but faded to brown after drying. Probably all

102. QuarTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

tropical populations of the species have dark brown or black ab-
domens with bluish or greenish spots.

Although published records are valueless at the subspecies level
(all West Indian and Mexican records, for example, having been
listed merely as longipes), I consider the two forms allopatric rep-
resentatives of one another, the relationship best expressed by the
use of trinomials. Provisionally, A. 1. longipes is restricted to the
United States, occurring along the Atlantic and Gulf coastal plains
from Massachusetts (Hagen, 1890) to eastern Texas (T. W. Don-
nelly, in litt.). At least a few individuals have been found west
of the Appalachians in Ohio (Borror, 1937; Borror and Epstein,
1942) and Indiana (Montgomery, 1949). A. I. concolor probably
occurs throughout the West Indies and from Mexico (Veracruz
and Guerrero) through Central America to Brazil. Thus the two
races are separated by a salt-water gap between the Bahamas and
Florida and a desert gap between central Texas and northeastern
Mexico, both of which are important biogeographic barriers for
terrestrial organisms. Either gap might be crossed with ease by
a strong-flying species like longipes, but breeding success should
be poor to nonexistent in the intervening areas. Even assuming
the sporadic incursion of individuals of one race into the territory
inhabited by the other, interbreeding may be prevented by the
visual dissimilarity of the races. If indeed odonates choose their
mates visually, at least in groups which bear bright, species-specific
patterns, then longipes and concolor are as effectively isolated re-
productively as any two species of Anax. I am at present retaining
them as subspecies because of their structural identity and obvious
very close relationship.

Gynacantha ereagris Gundlach. ANnpros, Andros Town, 31 Oc-
tober (2). Westfall (1960) overlooked a Bahamian record of this
species (Calvert, 1919), a male collected on Crooked Island on 24
November 1890 by the University of Pennsylvania Expedition. On
Cat Island ereagris was fairly common in low scrub adjacent to
a marshy area east of the coastal coppice at Orange Creek. One
was shot after observing it at close range, but a large snake (Al-
sophis), which darted out from the spot where it fell, diverted my
attention long enough for the dragonfly to flutter away. On San
Salvador, a few brown aeshnids were seen which were probably
this species. From those observed on Cat Island, my impression
is that this species frequents more open areas than does G. nervosa.

Pautson: Odonata of Bahamas 103

Only two were observed in the dense coppice, whereas many were
seen in the open scrub nearby.

Gynacantha nervosa Rambur. soutH Brmint, 18 November (1).
Five or six were seen in the coppice in as many hours of field work
there. As none was found on the other islands by me or previous
collectors, it may be that nervosa has reached the Biminis from
southern Florida, where it is very common.

Triacanthagyna trifida Rambur. SAN sALvaApor, 7.1 mi. N Cock-
burn Town, 27 December (1). This species swarmed at dusk in
small numbers at two localities on San Salvador.

Orthemis ferruginea Fabricius. sourH BrMINi, 16-22 August
(1); ANDROS, 2.5 mi. SE Andros Town, 31 October (4); CaT ISLAND,
1 mi. S Tea Bay, 29 November (2); sAN SALVADOR, 6.6 mi. S Cock-
burn Town, 1 January (1). Both sexes were observed in numbers
on all islands visited, being exceeded in general abundance only
by Erythrodiplax umbrata.

Libellula needhami Westtall. NEW PROVIDENCE, July-August
1961 (1). Dr. Westfall has asked me to include this specimen, a
male collected by Martin Dickinson, in the present paper. It
furnishes the first Bahamian record for needhami.

Macrodiplax balteata Hagen. cat IsLAND, 1 mi. S Tea Bay,
30 November (1). Males were seen at a number of open Chara-
filled fresh-water ponds on Cat Island, and a pair in tandem flew
over one of the ponds for some time. A last instar larva was col-
lected at Andros Town, Andros.

Idiataphe longipes cubensis Scudder. ANpDROs, 5 mi. NW Fresh
Creek, 1 November (1).

Brachymesia furcata Hagen. CAT ISLAND, 1 mi. S Tea Bay, 29
November (2). This species was fairly common at ponds on Cat
Island, and one male was observed at a pond on San Salvador.

Pachydiplax longipennis Burmeister. ANpDRos, Andros Town, 31
October (1); 2.5 mi. SE Andros Town, 31 October (2).

Micrathyria didyma Selys. caT IsLAND, 1 mi. S Tea Bay, 29
November (1); Tea Bay, 1 December (1); SAN saLvapor, 6.6 mi. S
Cockburn Town, 1 January (1). A few of each sex were seen at
small ponds and in the coppice near Tea Bay. Exuviae were col-
lected there and at Knowles on stems of Typha within a foot of the
water. A few adults perched on Phragmites at the pond on San
Salvador.

Erythrodiplax berenice naeva Hagen. soUTH BIMINI, 16-22 Au-

104. QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

gust (23); EAST BIMINI, 21 August (4); ANDROS, 2.5 mi. SE Andros
Town, 31 October (1). Westfall (1960) recorded E. b. berenice
Drury from the Bimini Islands and E. b. naeva from elsewhere in
the Bahamas, following determinations made by Dr. Donald J.
Borror. My Bimini material is rather intermediate between the
two races. The small stigma and relatively open venation ally it
with naeva, as does the lack of pigment in the wings of all females.
The penes are not the much angulated structures of naeva, but
they diverge from Floridian berenice in the direction of naeva.
The single Andros male is identical in venation and stigma size to
the Bimini males, but its penis is more typical of naeva, although
still not as extreme as those of Greater Antillean males or even
many from the Florida Keys. In general, I believe that all Ba-
hamian specimens of the species should be referred to naeva,
even though the structure of the penis varies as much as it does.
Bimini specimens are quite distinct from E. b. berenice as typified
by specimens from North Carolina, Florida, Mississippi, and Texas.

Erythrodiplax connata Burmeister. ANDROS, 2.5 mi. SE Andros
Town, 31 October (1). This female is apparently the same as spe-
cimens from Andros and New Providence associated with E. con-
nata by Borror (in Westfall, 1960).

Erythrodiplax justiniana Selys. anpros, 2.5 mi. S Andros Town,
30 October (1).

Erythrodiplax umbrata Linnaeus. SOUTH BIMINI, 16-22 August
(14); anpros, Andros Town, 31 October (1); 2.5 mi. SE Andros
Town, 31 October (3); 2.7 mi. SE Andros Town, 31 October (8);
2 mi. NW Fresh Creek, 1 November (1); sAN SALVADOR, 6.6 mi. S
Cockburn Town, 1 January (1). Males were seen in numbers in
late November at an area where leakage from a water pipe pro-
duced marshy conditions on South Bimini. They were also com-
mon around a few ponds on Cat Island and one on San Salvador.
Females were widespread in scrub and coppice on South Bimini
and Cat Island, greatly outnumbering males. The series from
Andros, collected at random around several ponds, included more
young than mature individuals. Larvae were collected at the same
ponds, and exuviae were found within a few inches of the water in
a sinkhole pond on Cat Island.

Lepthemis simplicicollis Say. car 1sLAND, 1 mi. S Tea Bay, 29
November (1). This species was fairly common at a few localities
on Cat Island but was not observed on San Salvador, where it was

PAULSON: Odonata of Bahamas 105

apparently common in March 1953 (Westfall, 1960). Two exuviae
and a few small larvae were taken on Cat Island.

Lepthemis vesiculosa Fabricius. This species is uncommon in
the Bahamas; the Van Voast-American Museum of Natural History
Expedition collected none, and I saw no adults on my fall and win-
ter visits at a season when vesiculosa is common in south Florida.
However, an exuvia and last instar larvae were collected from small
ponds at two localities on Cat Island.

Tramea abdominalis Rambur. cat IsLAND, Tea Bay, 30 Novem-
ber (1); Orange Creek, 1 December (1); san sauvapor, 6.6 mi.
S. Cockburn Town, 1 January (1). Only scattered individuals of
this and the following species were seen.

Tramea binotata Rambur. GRAND BAHAMA ISLAND, Freeport,
11 April (1); ANpRos, 2.5 mi. SE Andros Town, 81 October (3); 1.5
mi. S. Andros Town, 1 November (1); Cat 1sLaNnp, 1 mi. S Tea Bay,
30 November (2); san saLvapor, 7.1 mi. N Cockburn Town, 28
December (1). This species was slightly more common than T.
abdominalis on the islands visited. Among small numbers of tra-
meas with narrow wing spots observed on South Bimini, several
were approached closely enough to associate them definitely with
this species.

Lramea lacerata Hagen. One female of this species seen on
South Bimini in late November was easily identified by its overall
black color, large wing patches, and conspicuous yellow abdominal
spots. These occupy most of segment seven of the abdomen and
are an excellent field recognition character for T. lacerata.

Tramea onusta Hagen. soutTH Bimini, 16-22 August (3); CAT
ISLAND, The Bight, 28 November (2); Knowles, 1 December (3);
SAN SALVADOR, 6.6 mi. S Cockburn Town, 1 January (1). A few
onusta were seen on South Bimini and San Salvador, and on Cat
Island it was everywhere the common species. Males and mated
pairs were present at all the fresh-water ponds, the pairs oviposit-
ing in typical Tramea fashion. Immature individuals swarmed
over the road from one end of the island to the other, and larvae
of all sizes and exuviae were common at most ponds. Three larvae
were reared.

Pantala flavescens Fabricius. soUTH BIMINI, 16-22 August (1);
19 November (1); anpRos, 2.5 mi. S Andros Town, 30 October (2);
SAN SALVADOR, 3.3 mi. S Cockburn Town, 1 January (1). A few
adults were seen on South Bimini, and full-grown larvae were com-

106 QvuARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

mon in a small fresh-water rockpit along with those of Anax junius.
An exuvia was found on a grass stem at an almost dry pond in
pineland on New Providence, 26 November 1963, and a few adults
were seen at the same place. Two larvae were collected at 5 mi.
NW Fresh Creek on Andros. A few adults were seen on Cat Island
and more on San Salvador, where one was collected in late after-
noon hanging from a twig at the edge of a path through the cop-
pice. It moved up the path ahead of me in a manner reminiscent
of a Gynacantha.

Discussion

Westfall (1960) listed 26 species of Odonata from the Bahama
Islands, two of which (Epiaeschna heros, Pachydiplax longipennis)
were without specific locality data. Gynacantha ereagris, listed
by Calvert (1919) from the islands, brought the list to 27. The
present paper cites a definite locality for Pachydiplax and adds six
species to the known fauna (Lestes scalaris, Gynacantha nervosa,
Libellula needhami, Macrodiplax balteata, Lepthemis vesiculosa,
and Tramea lacerata), bringing the total to 33. Of this list, at least
six species (Coryphaeschna ingens, Epiaeschna heros, Gynacantha
nervosa, Celithemis eponina, Libellula needhami, and Tramea
lacerata) may be only vagrants from the nearby Florida peninsula,
but the others are probably breeding residents. All previous rec-
ords of Bahamian dragonflies referred only to the adults, but the
present paper records the collection of larvae of at least twelve
species.

As presently known, the Bahamian odonate fauna indicates
affinities with both the Greater Antilles and the southeastern United
States. Records of Coryphaeschna ingens, Epiaeschna heros,
Celithemis eponina, and Tramea lacerata indicate mainland affin-
ities, but these isolated records of stragglers are of little zoogeo-
graphical importance. Pachydiplax longipennis remains the only
species which faunistically ties the Bahamas to Florida rather than
the Antilles. From the south the Bahamas have received popula-
tions of Lestes scalaris, L. spumarius, Argiallagma minutum, Anax
longipes concolor, Gynacantha ereagris, Micrathyria didyma, Ery-
throdiplax connata cf. connata, E. justiniana, and Dythemis ru-
finervis, none of which occur in Florida, as well as eleven tropical
species which have reached Florida as well. Of special interest
is the presence in south Florida of Antillean species which have

PauLson: Odonata of Bahamas 107

not colonized the Bahamas so far as is known. These include
Lestes tenuatus, Enallagma cardenium, Neoerythromma cultella-
tum, Coryphaeschna viriditas, Miathyria marcella, and Tauriphila
australis. Anax amazili and Brachymesia herbida, both wanderers
to south Florida, do not breed there and cannot be considered
important in this connection. The stream-dwelling habits of E.
cardenium may have precluded its establishment in the Bahamas,
as would the breeding habits of Miathyria and Tauriphila (Paul-
son, 1966), but L. tenuatus, N. cultellatum, and C. viriditas should
find satisfactory breeding sites in the Bahamas. The fortuitousness
of colonization may be the most important factor in the differences
between the tropical odonate faunas of south Florida and the
Bahamas.

The seasonal aspects of Neotropical dragonfly faunas have re-
ceived little attention in the literature. This has been because
collections were made on short visits of nonresident collectors who
could not draw a picture of seasonal changes. Calvert attempted
to correlate the seasonal distribution of a few forms with the
rainy season (1908, 1931; Calvert and Calvert, 1917), but he un-
fortunately never published the seasonal data he accumulated
for many species in his year in Costa Rica. The writer (1966)
found that as high as 29 per cent of the anisopteran and 42 per
cent of the zygopteran fauna of southern Florida were found as
adults throughout, or at least during, mid-winter. As on the Trop-
ical mainland and elsewhere in the West Indies, the Bahamian
fauna is too poorly known seasonally to allow the conclusions pos-
sible to workers in better documeted areas. However, available
records indicate that a very high proportion of Bahamian species
fly throughout the year. All of the widespread and common resi-
dents (Lestes spumarius, Argiallagma minutum, Ischnura ramburi,
Anomalagrion hastatum, Orthemis ferruginea, Idiataphe longipes,
Brachymesia furcata, Erythrodiplax berenice, E. umbrata, Tramea
abdominalis, T. binotata, T. onusta, and Pantala flavescens) have
been taken in the winter months (December to February). Drag-
onfly collecting in the Bahamas has been concentrated in the win-
ter (October to April), and mid-summer records are scattered.
Enough of the latter are available for the less common species,
most of which have been taken in the winter, to indicate an all-
year season for them as well. Mid-winter occurrence has been
reported for Anax longipes, Brachymesia furcata, and Erythrodiplax

108 QuvaRTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

berenice, species usually absent at that season in southern Florida.
The southern Bahamas, south of the Florida peninsula, are the
source of some of these records. On both Cat Island and San Sal-
vador I was impressed by the abundance of Odonata relative to
their numbers in the Florida Keys at the same season. The Keys
are climatically much like the Bahamas and could support adult
Odonata in numbers throughout the year, yet an extreme mid-
winter reduction in individuals if not species in the former area
has been brought about by some extrinsic factor(s) apparently not
present in the Bahamas. These tropical species might be expected
to be relatively intolerant of temperate conditions, yet it is note-
worthy that they are the long-season species in southern Florida,
rather than the temperate species which have moved into that
area from the north. Although poorly documented, it appears
that some of the Bahamian species have flight seasons like those
which they occupy in southern Florida. Thus the gynacanthines
fly during the fall and Pantala hymenaea in mid-summer.

A final word on the subject of Bahamian Odonata concerns fur-
ther problems in the area. A thorough knowledge of the fauna
of the major islands, especially Grand Bahama, Andros, and Great
Inagua, will be necessary to elucidate the present zoogeographic
status of the area. The relative proximity of Andros to Florida
and its extensive fresh-water marshes make it a likely target for
invasion by Floridian species, especially those of the Everglades.
Coryphaeschna ingens, Libellula needhami, Celithemis eponina,
and Brachymesia gravida, for instance, should find their ecological
requirements adequately satisfied on that island as Pachydiplax
apparently has. At least ten species of Bahamian birds, mostly
restricted to the northern islands, show direct affinities with Flor-
ida forms rather than Antillean forms (Bond, 1956), and a similar
phenomenon may yet be discovered in the Odonata. In addition,
the Bahamas from a practical standpoint represent a very access-
ible area to study life histories and habits of tropical dragonflies,
amply justifying repeated visits by the serious investigator.

LITERATURE CITED

Bono, JAMES. 1956. Check-list of birds of the West Indies. Acad. Nat.
Sci. Phila., Philadelphia, ix + 214 pp.

Borror, Donatp J. 1937. An annotated list of the dragonflies (Odonata)
of Ohio. Ohio Jour. Sci., vol. 37, pp. 185-196.

Pautson: Odonata of Bahamas 109
Borror, DONALD J., AND ERwIN J. Epstein. 1942. New records of Ohio
dragonflies (Odonata). Ohio Jour. Sci., vol. 42, pp. 81-83.

Byers, C. Francis. 1930. A contribution to the knowledge of Florida Odo-
natin, Ela. Publ., Biol. Sci. Series, vol. 1, pp. 1-327.

CALVERT, AMELIA S., AND Puitie P. Catvert. 1917. A year of Costa Rican
natural history. The MacMillan Co., New York, xix + 577 pp.

Catvert, Pumir P. 1901-1908. Biologia Centrali-Americana. Vol. 50: Neu-
roptera. Porter & Dulau, London, xxx + 420 pp., 10 pls.

1908. The composition and ecological relations of the odonate
fauna of Mexico and Central America. Proc. Acad. Nat. Sci. Phila.,
1908, pp. 460-491, pl. 26.

—. 1909. Contributions to a knowledge of the Odonata of the Neo-
tropical region exclusive of Mexico and Central America. Ann. Car-
negie Mus., vol. 6, pr. 73-281.

—————. _ 1919. Gundlach’s work on the Odonata of Cuba. Trans. Amer.
Ent. Soc., vol. 45, pp. 335-396.

—. 1931. The generic characters and the species of Palaemnema (Odo-
mata: Agrionidae). Trans. Amer. Ent. Soc., vol. 57, pp. 1-111, pls. 1-21.

Hacen, HERMANN A. 1890. Synopsis of the Odonata of North America,
No. 2. The genus Anax. Psyche, vol. 5, pp. 303-308.

Kirsy, W. F. 1890. A synonymic catalogue of Neuroptera Odonata, or
dragonflies. Gurney & Jackson, London, ix + 202 pp.

Kiots, Evste B. 1932. Insects of Porto Rico and the Virgin Islands. Odo-
nata or dragon flies. Sci. Surv. Porto Rico and Virgin I., vol. 14, pp.
LOW, wie, lee

Martin, RENE. 1908-1909. Coll. Zool. du Baron Edm. de Selys Long-
champs. Cat. System. et Descrip. Fasc. 18-20, Aeschnines. Bruxelles,
223 pp., 6 pls.

MontTcoMErRY, B. Etwoop. 1949. Anax longipes Hagen (Odonata: Aeshni-
dae), an Atlantic Coastal species in Indiana. Proc. Indiana Acad. Sci.,
VOM DONOR ELL OIL.

Muttkowskt, RicHarp A. 1910. Catalogue of the Odonata of North Amer-
ica. Bull. Publ. Mus. Milwaukee, vol. 1, pp. 1-207.

NEEDHAM, JAMES G. 1941. Life history notes on some West Indian
coenagrionine dragonflies (Odonata). Jour. Agric. Univ. Puerto Rico,
Ol DMD l= 19)

Pautson, Dennis R. 1966. The dragonflies (Odonata: Anisoptera) of south-
ern Florida. Unpubl. doctoral dissertation, University of Miami,
603 pp.

11G QuaRTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Rass, GeorcE B., AND Exuis B. HAaypEn, Jr. 1957. The Van Voast-Amer-
ican Museum of Natural History Bahama Islands Expedition. Record
of the expedition and general features of the islands. Amer. Mus.
Novitates, no. 1836, pp. 1-53.

VaurnigE, Parricta. 1952. Insect collecting in the Bimini Island group, Ba-
hama Islands. Amer. Mus. Novitates, no. 1565, pp. 1-24.

WESTFALL, MINTER J., JR. 1960. The Odonata of the Bahama Islands, the
West Indies. Amer. Mus. Novitates, no. 2020, pp. 1-12.

WHITEHOUSE, Francis C. 1943. A guide to the study of dragonflies of

Jamaica. Bull. Inst. of Jamaica, Sci. Series, no. 3, pp. 1-69.

7450 S. W. 102 Street, Miami, Florida, 33156 (mailing address),
and Department of Zoology, University of Washington, Seattle,
Washington, 98105.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

Hermaphroditism in Mullet, Mugil cephalus Linnaeus
Martin A. Mog, JR.

THE striped or black mullet, Mugil cephalus Linnaeus, is found
throughout the world in tropical and temperate areas. Because
of its great commercial importance and abundance, this species
has been the subject of extensive research. Thompson (1963) has
provided a useful synoptic treatment of the world-wide biological
data that have been accumulated.

Despite wide interest in reproduction of striped mullet, only
two well authenticated observations of mullet spawning have
been recorded. The work of Anderson (1958) and the observa-
tions of Arnold and Thompson (1958) indicate that spawning
normally occurs well offshore in waters 20 to at least 750 fathoms
deep. When they have a total length of about 20 mm, the fry
enter inshore nursery areas. Stenger (1959) described the anatomy
and histology of the gonads of individuals ranging from 20 mm
total length to mature adults. He found that M. cephalus develops
through a juvenile, sexually-indifferent stage into either a male
or female. Normal, functional intersexuality does not exist in
M. cephalus, although oocyte-like cells are found in the undiffer-
entiated and male gonads.

HERMAPHRODITISM

In describing the commercial production of M. cephalus in
Australia, Kesteven (1942) stated that, “Hermaphrodite roe are not
very rare, one or two being found each season,” but he provided no
details. Stenger (1959) discovered an ovotestis when an apparently
normal ovary from a 250 mm SL individual was sectioned. Islets
of testicular tissue were scattered throughout both lobes of the
predominately ovarian gland. Male and female elements were
not separated by mesenteries, and small oocytes in various stages
of growth were scattered throughout the male tissues. The organ
was encased by a muscular tunica characteristic of a maturing
ovary. According to Atz (1964), these are the only recorded in-
stances of hermaphroditism in adult M. cephalus.

Orlandi (1902) described an ovotestis found in a specimen of
Mugil chelo Cuvier. Only the gonad was obtained from a fisher-
man. Orlandi’s description shows the gland to be primarily an
ovary with secondary development of testicular tissue in most of

112 QuaRTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

the left lobe. In addition, a small, superficially lobular appendix
filled with testicular tissue was laterally attached to the left side
of the common trunk.

Two hermaphroditic striped mullet, M. cephalus, have been
brought to the Florida Board of Conservation Marine Laboratory
in recent years. In both instances, commercial fishermen saved
the gonads as a curiosity and gave them to the Marine Laboratory
after the fish had been processed for market.

On October 28, 1959, an officer of the Florida Conservation
Patrol obtained an ovotestis of M. cephalus from a commercial
fisherman in Sarasota County. The gland was delivered to Dr.
V. G. Springer at the Marine Laboratory where it was preserved
in 10 per cent formalin. The ovotestis is small, about 30 mm in
length, and is predominantly testicular. Ovarian tissue forms the
posterior half of the right lobe and occurs on the left lobe as a
small superficial patch on the posterior end. A portion of the
patch of ovarian tissue on the left lobe is missing. Male and
female elements are separated by mesenteries, and there is ap-
parently no integration of male and female tissue. The ova aver-
age 0.3 mm in diameter (eggs of normal, ripe females are about
0.7 to 1.0 mm in diameter). The right lobe measures 30 mm in
length and 6.5 mm at its greatest width, and the left lobe measures
20 mm in length and 8.0 mm in width. The center of the right
lobe is constricted to 2.0 mm at the point where the male and
female elements join. The ducts and common trunk were de-
stroyed when the gonad was removed from the fish, and the pres-
ence of a vas deferens or an oviduct could not be determined.

In September 1963, a hermaphroditic M. cephalus was found
in the commercial catch at Saunders Fish House at Dunedin,
Florida. The gland was taken from the fish, eventually frozen,
and delivered to the Marine Laboratory. It was then refrozen and
came into my possession several days later. The posterior tip of
the right lobe was preserved in Bouin's fluid and the rest of the
gonad was preserved in 10 per cent formalin. This gonad consists
of two separate lobes joined posteriorly into a common trunk. The
ducts were not removed with the organ and their structure is un-
known. The membranes of the common trunk, however, have
the appearance of an ovarian muscular tunica, and the testicular
elements are separated from the lumen of the common trunk by
serosal membranes. Fig. 1, illustrates the general form of the

Mor: Hermaphroditism in Mullet 113
gonad. The left lobe is 125 mm long and 30 mm at its greatest

width, and the right lobe is 98 mm long and 35 mm wide. Testic-

supporting
mesogonium

Right
lobe

male tissue ze
female tissue

VENTRAL VIEW

1. Ovotestis of hermaphroditic mullet (Mugil cephalus). Maximum
aa a left lobe 125 mm.

114. QvuaARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

lar tissue fills both lobes of the gonad. Ovarian tissue is most
abundant in the anterior third of the left lobe, but even in this
area it does not completely replace the underlying testicular tissue.
Ovarian tissue occurs elsewhere as small superficial patches on the
testicular tissue. A patch of ovarian tissue on the posterior end
of the left lobe fills the common trunk. Male and female tissues
are completely separated by mesenteries.

The tip of the right lobe containing both male and female
elements was sectioned at eight microns and stained with Harris’
hematoxylin and eosin. The tissue was in poor condition because
of post-mortem histolysis and the effects of freezing; nevertheless,
the resulting preparations showed the major features of internal
structure. A section of the gland containing both male and fe-
male elements and the mesenteries separating them is shown in
Fig. 2. The gonad is encased by a complex mesenteric capsule
that varies in thickness. This capsule is thickest between the male
and female tissues and is generally thicker around the female ele-
ments than the male. The capsule appears to be an ovarian mus-

Fig. 2. Section through right lobe of ovotestis of hermaphroditic mullet
(Mugil cephalus). Haematoxylin and eosin 37X.

Moe: FHermaphroditism in Mullet nS)

cular tunica in the stained section and over most of the gonad.
The testicular tissue is encased in thin serosal membranes which
are exposed in areas distant from ovarian tissue and within the
common trunk.

The internal structure of the testicular elements in the stained
sections appears abnormal. ‘There is an extensive proliferation of
stromal tissue that has disrupted normal tubular structure, but
spermatogenesis appears normal. The ovarian tissue contains
advanced oocytes and shows a normal lamellar structure within
the stained sections. Both male and female elements are ripen-
ing and appear to be at the same developmental stage as normal
gonads of mullet destined to spawn within two or three months.
The diameter of the ova averaged 0.4 mm.

Both gonads described above appear to be basically female
with extensive secondary development of testicular tissue. In the
case of the larger gonad, this view is supported by the similarity
of the capsule to the muscular tunica of a normal ovary and the
disorganized structure of the testicular tissue.

The possibility of self-fertilization in abnormally hermaphro-
ditic fish has been the basis of much conjecture. Johnson (1932) dis-
cussed the problem and advanced the following three conditions
necessary for self-fertilization to take place: “1, simultaneous mat-
uration of ova and sperm in the one individual; 2, a complete sys-
tem of gonadial ducts which will permit union of the ripe germ
cells; 3, the sperm and ova must be of a physiological constitution
which permits of mutual activation resulting in fertilization.” To
these conditions should be added: 4, the individual must be
capable of spawning behavior characteristic of the species or be
capable of abnormal release of gametes. Seltf-fertilization may
occur either internally before the release of gametes, or externally
after the spawning act. Stenger (personal communication) has
suggested that the hermaphroditic mullet he described in 1959
would probably have spawned as a female in December, and if
the sperm were then present and viable, self-fertilization would
seem to be possible. The larger gonad described above (taken
in September 1963) might have spawned in October or November,
and if male and female gametes were released simultaneously, self-
fertilization would probably have occurred.

I do not believe macroscopically detectable hermaphroditism in
Florida mullet is as common as Kesteven (1942) reports for Austra-

116 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

lian mullet. No other reports of hermaphroditism in M. cephalus
were obtained despite conversation and correspondence with many
ichthyologists and fish dealers in Florida. It is, however, possible
that the type of hermaphroditism discovered by Stenger (1959) is
relatively common since the gland must be sectioned before the
abnormal condition is evident.

Doctors James W. Atz and Albert H. Stenger graciously re-
viewed the manuscript and Richard P. Saunders, Florida Board
of Conservation Marine Laboratory, provided the photograph.

LITERATURE CITED

ANDERSON, W. W. 1958. Larval development, growth, and spawning of
striped mullet (Mugil cephalus) along the south Atlantic coast of the
United States. U. S. Fish and Wildl. Serv., Fish. Bull., vol. 58,
no. 144, pp. 501-519.

ARNOLD, FE. L., AND J. R. THompson. 1958. Offshore spawning of the
striped mullet, Mugil cephalus, in the Gulf of Mexico. Copeia, 1958,
pp. 130-1382.

Arz, J. W. 1964. Intersexuality in fishes. In: Intersexuality in vertebrates
including man (C. N. Armstrong and A. J. Marshall, eds.), pp. 1-479.
Academic Press, New York and London.

Jounson, H. H. 1932. The problem of self-fertilization in teratologically
hermaphroditic fishes. Copeia, 1932, pp. 21-26.

KESTEVEN, G. L. 1942. Studies in the biology of Australian mullet. 1-An
account of the fishery and preliminary statement of the biology of
Mugil dobula Gunther. Coun. for Sci. Industr. Res. Austral., Bull. no.
lov pp. L147.

Oruanpi, S. 1902. Sopra un caso di ermafroditismo nel Mugil chelo Cuyv.
Atti Soc. Ligust. Sci. Nat. Georgr. Genova, vol. 13, pp. 3-6 (informal
translation, Dr. James W. Atz)

STENGER, A. H. 1959. A study of the structure and development of certain
reproductive tissues of Mugil cephalus Linnaeus. Zoologica, vol. 44,
no. 3, pp. 53-70, 8 pls.

THomeson, J. M. 1963. Synopsis of biological data on the gray mullet
Mugil cephalus Linnaeus 1758. CSIRO Fish. and Ocean., Fish. Synop.
NOs ee Dp OU:

Florida Board of Conservation Marine Laboratory, St. Peters-
burg, Florida. Contribution No. 105.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

Review of the Lutjanus campechanus Complex of Red Snappers
Luis R. Rivas

THE taxonomic status of certain species of Lutjanus, commonly
known as red snapper, has remained confused in spite of several
reviews during the past 80 years (Jordon and Swain, 1885; Jordan
and Fesler, 1893; Jordan and Evermann, 1898; Hildebrand and
Ginsburg, 1925; Ginsburg, 1930).

As pointed out by Camber (1955) and Carpenter (1965) at least
10 species of snappers are marketed as red snapper. Commercial
fishermen, however, recognize each of these as a separate species
to which they refer by its individual common name. The desig-
nation of “red snapper” is given only to the species variously re-
ferred to in the literature as Lutjanus aya, L. blackfordi, and L.
campechanus, and sometimes also to L. vivanus better known as
“silk snapper.”

This study shows that, in addition to the silk snapper, there
are two species of what may be called true red snappers. One
of these, L. campechanus, appears to be restricted to the Gulf of
Mexico and the South Atlantic coast of the United States. It
perhaps also occurs in Bermuda, the Bahamas, and along the
north coast of Cuba. The other, L. purpureus, occurs in the Car-
ibbean Sea and its range extends southeastward along the coast
of the Guianas and probably to Brazil. Commercial fishermen
call L. campechanus the “Gulf red snapper” and L. purpureus the
“Caribbean red snapper.”

Although Hildebrand and Ginsburg (1925) and Ginsburg (1930)
had previously recognized two species of red snappers their con-
clusions were open to question. Because of the paucity of speci-
mens available to them it was thought that the apparent specific
differences might be due to intraspecific variation. In addition
these authors misinterpreted the nomenclature as discussed below
under the species headings.

This study also shows that L. vivanus, the silk snapper, is close-
ly related to L. campechanus and L. purpureus, especially to the lat-
ter with which it has been confused. These three species, herein
referred to as the L. campechanus complex, form a group well dis-
tinguished from the other members of the genus Lutjanus. Gins-
burg (1930) pointed out the close relationship among these three
species.

118 QuaARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MATERIAL AND ACKNOWLEDGMENTS

Positive identification of the Gulf and the Caribbean red snap-
pers was initially effected by the study of fresh caught specimens
generously supplied by Clark Seafood, Inc., Pascagoula, Mississippi,
through the cooperation of Harvey R. Bullis, Jr. Later pertinent
material was studied at the United States National Museum
(usNM) and the University of Miami Ichthyological Museum
(umIM). The Museum of Comparative Zoology (mcz) and the
Academy of Natural Sciences of Philadelphia (ansp) made critical
specimens available for study. The cooperation of Ernest A. Lach-
ner (USNM), Mrs. Mywanwy M. Dick (mcz), and James Tyler (ANspP)
is sincerely appreciated.

A total of 188 specimens were examined. The number (in pa-
rentheses) following the catalogue number indicates the number
of specimens in the lot.

MeETHODS

Measurements and counts were made according to methods
already described by the author (Rivas, 1960) with the following
modifications and additions.

The mandible length is measured from the anterior tip of the
dentary to the posterior tip of the articular.

Lateral scales were counted as the number of oblique rows
(inclined forward) above the lateral line between the posttemporal
(scale bone) and the middle of the caudal base. The scales above
lateral line were counted downward and backward from the dor-
sal fin origin to, but not including, the lateral line. The scales
below lateral line were counted upward and forward from the
anal fin origin to, but not including, the lateral line. Opercular
scales were counted as the number of rows parallel with the mar-
gin of the subopercle; the uppermost row may comprise only one
scale. The scales above opercle were counted as the number of
oblique rows (inclined forward) above the opercle.

In the tables (2-4) discrepancies between the number of speci-
mens included and the total number of specimens available for
scale counts are due to partial or nearly total loss of scales as a
result of poor preservation. This occurred most frequently in the
L. campechanus specimens.

The gillraker counts presented a problem. In juveniles and
young the anterior rakers of the lower limb (first arch), although

Rivas: Revision of Red Snappers 119

short, may be made out easily. In the adults, however, these
rakers are reduced to tubercles which are difficult to distinguish
from the inter-raker tubercles. Also, the young show no distinc-
tion between developed and undeveloped (rudiments) gillrakers,
in contrast with the adult. On the upper arch all gillrakers whether
developed or rudimentary can be made out in both young and
adult. For this reason rudiments are not included in the counts
for the lower limb and, for this character, only specimens 100 mm
in standard length or larger are included in the key and Table 5.

Because of the long-standing confusion it has been difficult to
untangle the synonymies and references. Many references could
not be verified through lack of adequate descriptions, figures, or
records of available specimens. These references are not listed.

NONAPPLICABILITY OF NAMES PROPOSED BY BLOCH

Since its original proposal the specific name aya has been fre-
quently applied to the red snapper. However, a study of the orig-
inal description of Bodianus aya Bloch (1790, p. 45) reveals that
the name almost certainly does not apply to a species of Lutjanus.

Bloch states that a spine occurs at the posterior tip of the
opercle but this is not so in Lutjanus. The number of branchio-
stegals in lutjanids is 7, not 5 as stated by Bloch. His figure clearly
shows 9 dorsal spines followed by 19 rays and one anal spine fol-
lowed by 8 rays. In the text the number of dorsal spines is given
as 9 and the total number of dorsal] elements as 27. The total
number of anal elements is given as 9 of which one is a spine. In
western Atlantic Lutjanus there are 10 dorsal spines, not 9, and
12-14 dorsal rays, not 18 or 19, with a total of 22-24, not 27 or 28
dorsal elements. There are 3 anal spines and 8 or 9 anal rays,
a total of 11 or 12, not 9 anal elements. The anal fin is described
(and figured) as rounded whereas in the red snappers it is conspicu-
ously pointed. Also the anal fin is placed too far back and the
pectoral and pelvic fins are too short. Although not described in
the text the figure clearly shows the occurrence of scales on the
interorbital, top of snout, preorbital, and suborbital. These areas
are devoid of scales in Lutjanus. Finally, the habitat of aya, as
given by Bloch (lakes of Brazil) is certainly not that of the red
snappers.

Bloch’s figure is reminiscent of a Sciaenops-like sciaenid except

120 QuARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

for the shape of the dorsal and caudal fins. Although a red color
is not common among sciaenids it does occur in Sciaenops ocellata,
the channel bass, also commonly known in Florida as “redfish.”
The profile and squamation of the head, as shown in Bloch’s fig-
ure, are sciaenid-like. The emarginate caudal fin, as described
and figured, is not a sciaenid feature, but in the adult Sciaenops the
caudal fin is truncate or even somewhate emarginate. The single
anal spine is another sciaenid feature which occurs in several spe-
cies of Cynoscion and in Menticirrhus. With respect to the habitat
given by Bloch, “Landseen von Brasilien,’ it may be said that it
would apply to a sciaenid rather than to a red snapper. It is pos-
sible that the name aya will eventually be found to apply to an
as yet unrecognized sciaenid from the little known coastal lagoons
of Brazil.

At best, some sort of perciform fish is recognizable from Bloch’s
description and figure but certainly not a lutjanid. There is no
specimen available whereupon the identity of aya could be veri-
fied since the name was based on a pre-Linnaean description by
Marcgrave and a drawing presumably by Prince Maurice. AI-
though Cuvier and Valenciennes (1828, p. 346) claim that the
original figure was altered in Bloch’s copy, Prince Maurice's draw-
ing is equally unidentifiable.

The name Bodianus ruber Bloch and Schneider (1801, p. 330)
was based on a condensed version of the original description of
Bodianus aya Bloch.

THE Lutjanus campechanus COMPLEX

The western Atlantic species of Lutjanus may be subdivided
into three well defined species groups as characterized in the fol-
lowing key.

la. Scales above opercle in 2 or 3 rows. Lateral scales 40 to 48, usually

41 to 47. Jaws subequal or upper jaw projecting beyond lower. <Ac-
cessory lateral lines on caudal fin absent, or rarely present only on lower
half. Lateral spot absent. Coloration not predominantly red in life.

1. L. griseus species group

lb. Scales above opercle in 4 to 7 rows. Lateral scales 46 to 53, usually
47 to 51. Lower jaw slightly to strongly projecting beyond upper. Ac-
cessory lateral lines on caudal fin present. Lateral spot present or ab-
sent. Coloration predominantly red in life.

2a. Dorsal rays 12, rarely 13. Two vertical rows of scales between

Rivas: Revision of Red Snappers 121

posterior margin of orbit and upper end of preopercular margin.
Lower jaw strongly projecting beyond upper. Lateral spot present.
2. L. synagris species group

2b. Dorsal rays 14, rarely 13. Three or four vertical rows of scales be-
tween posterior margin of orbit and upper end of preopercular mar-
gin. Lower jaw slightly projecting beyond upper. Lateral spot
present or absent.

3. L. analis species group

The L. campechanus complex (L. campechanus, L. purpureus,

L. vivanus) belongs to the L. analis species group, which also in-
cludes L. buccanella. The relationships among these species are
analyzed in the following key, which also provides a means for
identification.

kay

lb.

Pectoral rays 15 to 17, usually 16. Géillrakers 7 or 8; only one developed
on upper limb. Lingual teeth absent. Vomerine patch of teeth crescent-
shaped, without a median backward extension. Suborbital width 10 to
12 percent of standard length. Lateral spot present in young and adult.

Ie Gs analis

Pectoral rays 16 to 18, usually 17. Gillrakers 8 to 12, usually 9 to 11;
one to three, usually two developed on upper limb. Lingual teeth pres-
ent. Vomerine patch of teeth anchor-shaped, with a median backward
extension. Suborbital width 6 to 9 percent of standard length. Lateral
spot always absent or present only in young.

2a. Scales above opercle in 4 to 6, usually 5 rows. Lingual teeth in
a single patch. Posterior margin of anal fin rounded, the middle
rays not exserted, Anal fin length 59 to 64 percent of head length.

A conspicuous jet-black, comma-shaped mark on base of pectoral fin.
Tips of middle caudal rays not black. Lateral spot always absent.

2. L. buccanella

2b. Scales above opercle in 6 or 7 rows. Lingual teeth in two patches,
the anterior one much smaller. Posterior margin of anal fin angu-
late or pointed, the middle rays exserted. Anal fin length 65 to 81
percent of head length. No jet-black, comma-shaped mark on base
of pectoral fin. Tips of middle caudal rays black, the fin sometimes
entirely margined with black. Lateral spot present in young, diffuse
or absent in adult. (L. campechanus complex).

3a. Anal rays 9, rarely 8. Lateral scales 46 to 50, usually 47 io 49.
Scales above lateral line 7 to 10, usually 8 or 9. Scales below
lateral line 15 to 19, usually 16 or 17. Géillrakers 8 to II,
usually 10. Scales on anterior side of body, below lateral line,
conspicuously larger than those on posterior side. Suborbital
width 8 or 9 percent of standard length.

3. L. campechanus

122 QuaRTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

3b. Anal rays 8, rarely 9. Lateral scales 49 to 53, usually 50 or 51.
Scales above lateral line 9 to 12, usually 10 to 12. Scales below
lateral line 16 to 24, usually 17 to 23. Géillrakers 9 to 12,
usually 10 or 11. Scales on anterior side of body, below lateral
line, not conspicuously larger than those on posterior side. Sub-
orbital width 6 or 7 percent of standard length.

4a. Scales below lateral line 16 to 19, usually 18. Scales above

lateral line 9 to 11, usually 10. Cheek scales in 6, rarely

5 or 7 rows. Scales above lateral line, on anterior side of

body, smaller than those below. Pelvic fin length 53 to

62 percent of body depth. Lateral spot, when present

(young), about equal to, or larger than eye. Iris red in
live and freshly preserved specimens.

4. L. purpureus

4b. Scales below lateral line 20 to 24, usually 21 to 23. Scales
above lateral line 10 to 12, usually 11 or 12. Cheek scales
in 7, rarely 8 rows. Scales above lateral line, on anterior
side of body, about equal to those below. Pelvic fin length
63 to 76 percent of body depth. Lateral spot, when pres-
ent (young), smaller than eye. Iris yellow in live and fresh-
ly preserved specimens.
5. L. vivanus

For more rapid and positive identification, with the campecha-
nus complex, the sum of lateral scales and scales above and below
lateral line may be used. This is 69-75 in campechanus; 77-81,
rarely 76 or 82 in purpureus; and 82-87, rarely 81 or 88 in vivanus.

Lutjanus campechanus (Poey)

Gulf red snapper

Mesoprion campechanus Poey, 1860, p. 149 (original description; no specific
locality designated); 1861, p. 365 (listed; common name; Campeche,
from hearsay); 1868, p. 294 (eye color; weight; Campeche; Key West;
Cuba, from hearsay).

Lutjanus campechianus, Poey, 1875, p. 29 (references; yomerine teeth; com-
parisons; Key West; Campeche); 1962, p. 86 (description; comparisons;
history; Key West; Campeche; Habana), pls. 70 C-J, 71 A-C.

Lutjanus blackfordii Goode and Bean, 1879, p. 176 (original description; com-
parison; Pensacola; Savannah).

Lutjanus_ blackfordii, Hildebrand and Ginsburg, 1925, p. 80 (description;
comparison; Pensacola; Rebecca Shoals; Key West), fig. 1. Ginsburg,
1930, p. 269 (characters; commercial importance; biology; nomencla-
ture; synonymy).

Rivas: Revision of Red Snappers 123

Lutjanus vivanus (not of Cuvier and Valenciennes), Jordan and Swain, 1885,
p. 453 (comments; synonymy in part; Key West).

Lutjanus aya (not of Bloch), Jordan and Fesler, 1893, p. 447 (common names;
synonymy in part; habitat in part; occurrence in part; specific name
doubted; comments on types of campechanus and blackfordi), pl. 30.
Carpenter, 1965, pp. 1-35 (review of fishery; Gulf of Mexico), fig.
(cover photograph).

Neomaenis aya (not of Bloch), Jordan and Evermann, 1898, p. 1264 (common
names in part; description; comments on type of campechanus; synon-
ymy in part; Key West), pl. 197, fig. 516.

As the names aya Bloch (1790) and ruber Bloch and Schneider
(1801) do not apply to a snapper, campechanus Poey (1860) is the
oldest name available for the Gulf red snapper.

The most recent reviewers (Hildebrand and Ginsburg, 1925,
p. 82; Ginsburg, 1930, p. 372) applied the name campechanus to the
Caribbean red snapper. The evidence presented below, however,
indicates that the name campechanus refers to the Gulf red snap-
per and that the frequently accepted name blackfordi is synony-
mous with it.

In the first paragraph of the original description Poey states
that the fish is so named (campechanus) “parce quon le péche
également sur le banc de Campéche ...” Although no specific
locality is given in the original description Poey subsequently
states (1868, 1875) that his campechanus is taken in Campeche Bank
and in Key West. In his last (posthumous) publication Poey (1962)
added Havana to Campeche Bank and Key West. Poey never
gave any indication that campechanus occurred outside the Gulf
of Mexico. As already indicated in the introduction the Caribbean
red snapper ( purpureus) is not known to occur in the Gulf of
Mexico and the Gulf red snapper is not known to occur in the
Caribbean.

Since critical diagnostic characters were not given by Poey,
the original description of campechanus could apply to either the
Gulf or the Caribbean red snapper. In the light of the above dis-
cussion, however, and since the type almost certainly came from
Key West (see below), the name campechanus is here accepted
as the valid one for the Gulf red snapper.

Jordan and Evermann (1898) stated that the type of campech-
anus “ ... is a stuffed skin of a young fish... . ” without any indi-
cation of length or locality. Subsequently Howell-Rivero (1938,

124 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

p. 196) stated that the types of campechanus comprise two speci-
mens (mMcz 9982) the largest of which is the “holotype.” A study
of these specimens, however, shows that they actually are Car-
ibbean red snapper (purpureus) and neither one could have been
Poey’s type of campechanus for the following reasons.

The original description of campechanus was based on a single
specimen 370 mm. in length (total, as it was customary with Poey).
The largest specimen, considered by Howell-Rivero as the “holo-
type, is 355 mm in total length, 15 mm short of the required
length. The caudal fin is undamaged.

Among the specimens examined at the United States National
Museum a Gulf red snapper (USNM 25235) 373 mm in total length
(285 mm standard length) is believed to be the holotype of cam-
pechanus. The length is almost in perfect agreement and, accord-
ing to the records, the specimen came from Key West (to Havana)
and was sent by Poey. Furthermore, this specimen has only 8
anal rays (as given in the original description) instead of 9 which
is the usual number for campechanus (Table 5). This specimen,
here recognized as the holotype, is described as follows.

Dorsal spines, 10. Dorsal rays, 14. Anal spines, 3. Anal rays,
8. Pectoral rays, 17. Lateral scales, 47. Scales above lateral line,
7; below, 17. Cheek scales in 6 rows. Gillrakers, 9 (plus 5 rudi-
ments); one (plus 5 rudiments), on upper limb. Predorsal length,
418. Preanal length, 710. Head length, 400. Snout length, 153.
Suborbital width, 88. Maxillary length, 151. Mandible length,
193. Orbit diameter, 67. Interorbital width, 84. Body depth, 386.
Caudal peduncle depth, 121. Dorsal base length, 505. Anal base
length, 154. Pectoral fin length, 323. Pelvic fin length, 235. Anal
fin length, 272. Middle caudal rays length, 203. Scales on ante-
rior side of body, below lateral line, conspicuously larger than
those on posterior side. Scales above lateral line, on anterior side
of body, smaller than those below. Posterior margin of anal fin
pointed, the middle rays exserted. Lingual teeth in two patches,
the anterior one much smaller. Vomerine patch of teeth anchor-
shaped, with a median backward extension. General coloration
yellowish-brown after more than 100 years in preservation. Tips
of middle caudal rays black.

The holotype of L. blackfordi from Pensacola, Florida, 544 mm
in standard length (usnM 21330) has been examined and found to
be conspecific with campechanus. Two specimens of campech-

Rivas: Revision of Red Snappers 125

anus, 585 and 606 mm in standard length (usnmM 87823, 87824),
reported by Hildebrand and Ginsburg (1925, p. 81) as blackfordii
have also been examined.

Camber (1955, p. 16) discussed the occurrence of two “types”
of red snapper in the Gulf of Mexico (Campeche Bank). Type B
consisted of 16 specimens which were less humped, more slender,
and with smaller scales than 619 specimens of Type A. Camber
also stated that radiographs showed osteological differences be-
tween the two types but he did not say what those differences
were. The more slender smaller-scaled specimens of Type B are
suggestive of the Caribbean red snapper (purpureus) but the
material examined in this study does not show any species differ-
ences correlated with Camber’s types A and B. Of the specimens
examined from Campeche Bank, including some used by Camber
in his study (ummm 4848), some were slender, some were humped,
and some were intermediate. In fact, there was gradual intergrad-
ation between the extremes corresponding to Camber’s_ types.
There was no correlation between body depth and relative size
(number) of scales or any other meristic, proportional, or color
character. All these specimens are typical campechanus. The
possibility, however, that purpureus may occur in the Gulf of Mex-
ico cannot be dismissed.

This species differs from purpureus and vivanus in the higher
number of anal rays (Table 5), the fewer scales and gillrakers
(Tables 2-5), the longer head, snout, maxillary, mandible, and anal
fin, the deeper body, and the wider suborbital (Table 1). The
enlarged scales on the anterior side of the body are a good field
character to distinguish campechanus from purpureus and vivanus.

The distribution of campechanus appears to be restricted to the
shelves bordering the Gulf of Mexico and the South Atlantic coast
of the United States northward to Cape Hatteras. No verifiable
records from the Bahamas, the North coast of Cuba, or the Car-
ibbean Sea are available to the author. In South Florida and Cam-
peche Bank, at least, campechanus occurs syntopically (Rivas, 1964)
with vivanus.

The absence of a shelf along the Caribbean coast of Yucatan
and the Caribbean coast of extreme western Cuba, and on both
sides of the Windward Passage may be of significance in the allo-
patric distribution of campechanus and purpureus. The collecting
records available and other sources (Camber, 1955, p. 23) indicate

126 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

that these two species usually occur at depths of less than 80
fathoms.

Good illustrations of campechanus are given by Hildebrand and
Ginsburg (1925, fig. 1, as Lutianus blackfordii) and by Carpenter
(1965, cover photograph, as L. aya).

The commercial fishery for this species has been recently re-
viewed in detail by Carpenter.

Material examined. 129 specimens from the following 36 lo-
calities. Campeche Bank: Triangle lighthouse, usnm 196785 (1);
28 n. mi. ESE of Arcas Cays, USNM 158426 (8); 12 n. mi. NE of Arcas
Cays, ummm 4839 (1); Arenas Cays, umm 6107 (12); 65 n. mi. WNW
of Campeche, Mexico, umm 4837 (1); 130 n. mi. Nw of Campeche,
Mexico, umm 4840 (2); 58 n. mi. NW of Campeche, Mexico, UmMiM
4836 (1); 75 n. mi. N of Carmen, Mexico, umm 2425 (2); Gulf of
Campeche, umim 1226 (1); ummm 4842 (2); ummm 4848 (15). Off
Texas: 80 n. mi. S of Galveston, usnM 126763 (1); 83 n. mi. S of
Galveston, UsNM 185539 (1); 19 n. mi. E of Brazos Santiago, UMIM
2419 (11); 115 n. mi. ESE of St. Josephs Island, ummm 2420 (15);
98 n. mi. E of Corpus Christi, ummm 4883 (14). Off Louisiana: 25
n. mi. SE of Barataria Bay, umim 4841 (14). Off Mississippi: S ot
Mississippi Delta, usnm 155381 (1); usnm 155382 (1); S of Horn
Island, umm 6061 (1). Off Alabama: 35 n. mi. SW of Mobile,
UMIM 2422 (6). Off Florida: S of Pensacola, usnM 21330 (1); usnM
158625 (1); usnm 80682 (1); usnm 31918 (1); usnm 21463 (1); 22
n. mi. NNW of Loggerhead Key, Dry Tortugas, umim 4843 (1); off
Rebecca Shoals, usnm 87824 (1); 5 n. mi. N of Rebecca Shoals,
umiM 2374 (4); off Key West, usnM 25235 (1); off Miami, umm
672 (4); off Port Everglades, umm 4368 (1); 44 n. mi. sE of Cape
Canaveral, umim 6071 (3); 33 n. mi. ENE of St. Augustine, USNM
188515 (1). Off North Carolina: © of Cape Hatteras, usnm 133966
(1).

Lutjanus purpureus Poey
Caribbean red snapper

Mesoprion aya (not of Bloch), Cuvier and Valenciennes, 1828, p. 346 (descrip-
tion; size; comments; Haiti). Poey, 1866, p. 267 (compared with
profundus = vivanus; name purpureus attributed to Cuvier and Valen-
ciennes; Santo Domingo).

Lutjanus purpureus Poey, 1867, p. 157 (compared with profundus = vivanus);
1875, p. 28 (name attributed to Cuvier and Valenciennes; compared
with profundus = vivanus), p. 29 (original designation of name pur-

Rivas: Revision of Red Snappers PAT

pureus; synonymy in part; eye color; Batabano, Cuba). Jordan and
Fesler, 1893, p. 446 (comments on validity and name).

PNeomaenis aya (not of Bloch), Evermann and Marsh, 1900, p. 174 (common
names; description; life color; range in part; commercial value; habits;
angling value; synonymy excepted; Puerto Rico), pl. 20.

Lutianus campechanus (not of Poey), Hildebrand and Ginsburg, 1925, p. 82
(description; comparison; off Honduras). Ginsberg, 1906, p. 268
(characters in key), p. 273 (comments; comparison; eye color; nomen-
clature; synonymy in part; off Honduras). Howell-Rivero, 1938, p. 196
(specimens only: erroneously designated as types of L. campechanus).

Lutjanus aya (not of Bloch), Poey, 1962, p. 85 (synonymy in part; coloration;
compared with profundus — vivanus; comments; history; Batabano,
Buba; Santo Domingo; Puerto Rico), pl. 70 B.

The use of the name purpureus for the Caribbean red snapper
appears to be justified on the basis of the following discussion.

Poey (1866, p. 267) stated that Cuvier and Valenciennes (1828,
p. 346) had changed the name aya to purpureus in a subsequent
page of the same publication. At the same time Poey compared
his profundus (= vivanus) with the aya of Cuvier and Valenciennes
and commented that it (aya) could be confused with profundus.
Poey also stated that he had seen a specimen from “Santo-Domin-
go’ which he believed to be the same as Cuvier and Valenciennes’
aya (purpureus) but different from his profundus. The name pur-
pureus was never mentioned by Cuvier and Valenciennes and there
is no explanation as to why Poey erroneously attributed the name
to them. The fact remains, however, that Poey mentioned pur-
pureus in his paper and that he recognized it as representing a red
snapper closely related to, but different from profundus = vivanus.
It is also significant that Poey does not mention campechanus,
the other close relative described by him six years previously and
which he probably considered distinct enough not to be confused
with purpureus and vivanus. Subsequently Poey (1867, p. 157)
stated that the main difference between purpureus and profundus
(= vivanus) is the location of the small scales on either side of
the nape. This difference, although real, is of very minor impor-
tance in distinguishing these two species. Later Poey (1875, p.
28) again erroneously attributed the name purpureus to Cuvier
and Valenciennes. He also discussed his specimen from Santo
Domingo (previously referred to by him, 1866, p. 267) as an indi-
vidual 300 to 350 mm long, very similar to profundus (= vivanus)

128 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

but different and believed by him to be the true purpureus. In
this paper, on the next page, Poey (1875, p. 29) formally listed
“Lutjanus purpureus’ as a species heading (between L. profundus
and L. campechianus) with the names aya Bloch, ruber Bloch and
Schneider, aya Cuvier and Valenciennes, and purpureus Cuvier
and Valenciennes as synonyms. In addition, Poey stated that pur-
pureus is also found in Santo Domingo and that he had seen it
only once from Cuba (Batabano, south coast). Finally, in his last
(posthumous) publication, Poey (1962, p. 85) listed purpureus as a
synonym of aya which he considered distinct from campechanus
and profundus (= vivanus). The specimen figured in this publi-
cation is said by Poey to have come from Batabano.

It may be concluded from the above discussion that the name
purpureus refers to the Caribbean red snapper and that it should
be attributed to Poey, not to Cuvier and Valenciennes. Also,
although the name was mentioned by Poey in 1866 and 1867 his
designation in 1875: 29 may be accepted as the original. Although
no formal description or definition of purpureus was given by Poey
in 1866, 1867, and 1875 the name may not be declared a nomen
nudum according to the now current Rules since it was proposed
before 1931 and there are sufficient “indications.” In fact, the
species was actually described and figured by Poey (1962, p. 85,
pl. 70 B), as “Lutjanus aya” in his last publication.

Only one specimen was definitely referred to by Poey (1875, p.
28) as the true purpureus. He stated that the fish, from Santo
Domingo, was “.. . de 300 a 350 milimetros de largo . . . ” (total)
and that it had been sent by him to Agassiz. There is only one
specimen at the Museum of Comparative Zoology which could
possibly be this specimen. It is the specimen (mcz 9982), largest
of two by 110 mm, erroneously designated as the holotype of
campechanus by Howell-Rivero (1938, p. 196). This specimen, 355
mm in total length (273 mm. standard length), a typical purpureus,
is here recognized as the holotype of the species and described
below.

Dorsal spines, 10. Dorsal rays, 14. Anal spins, 3. Anal rays,
8. Pectoral rays, 17. Lateral scales, 50. Scales above lateral line,
10: below, 19. Cheek scales in 6 rows. Géillrakers, 11 (plus 5 rudi-
ments); 2 (plus 5 rudiments) on upper limb. Predorsal length, 393.
Preanal length, 698. Head length, 359. Snout length, 125. Sub-
orbital width, 68. Maxillary length, 135. Mandible length, 166.

Rivas: Revision of Red Snappers 129

Orbit diameter, 77. Interorbital width, 84. Body depth, 352.
Caudal peduncle depth, 111. Dorsal base length, 498. Anal base
length, 139. Pectoral fin length not measured (tip broken off).
Pelvic fin length, 206. Anal fin length, 249. Middle caudal rays
length, 179. Scales on anterior side of body, below lateral line, not
conspicuously larger than those on posterior side. Scales above
lateral line, on anterior side of body, smaller than those below.
Posterior margin of anal fin pointed, the middle rays exserted.
Lingual teeth in two patches, the anterior one much smaller. Vom-
erine patch of teeth anchor-shaped, with a median backward ex-
tension. General coloration yellowish-brown after about 100 years
in preservation. Tips of middle caudal rays black.

A specimen of purpureus, 540 mm. in standard length (UsNM
87822) reported by Hildebrand and Ginsburg (1925, p. 82) as
Lutianus campechanus has been examined.

This species differs from campechanus in the characters already
indicated under that species. It is more closely related to vivanus,
from which it differs mainly in the fewer scales (Tables 2-4), the
shorter pectoral and pelvic fins (Table 1), the larger lateral spot,
and the red eye (yellow in vivanus). The eye color combined with
the number of anal rays and the relative size of the scales (see key)
constitute good field characters to distinguish purpureus trom
campechanus and vivanus.

The collecting data of the specimens studied and the tew veri-
fiable records from the literature indicate that purpureus occurs
on the shelves bordering the Caribbean Sea and that its range
extends southeastward along the coast of the Guianas probably
to Brazil. It is sympatric with vivanus with which it is also known
to occur syntopically (see Rivas, 1964). As already indicated under
campechanus the absence of a shelf on both sides of the Yucatan
Channel and the Winward Passage may be of significance in the
allopatric distribution of purpureus and campechanus.

A good illustration of purpureus (as Lutianus campechanus) is
given by Hildebrand and Ginsburg (1925, fig. 2).

Material examined. 41 specimens from the following 17 lo-
calities. Off Honduras: no specific locality, usnM 87822 (1); umm
6112 (6). Off Panama: 3 n. mi. N of Cabo Tiburon, umm 6094 (2).
Off Colombia: 28 n. mi. wsw of Cabo La Vela, umm 6111 (1). Off
Venezuela: 22 n. mi. NE of Cabo La Vela (Colombia), umrm 6086
(2); 12 n. mi. NNw of Punta Manzanillo, ummm 6093 (2). Off Aruba:

130 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

5 n. mi. sw of w end, umm 6110 (1). Off British Guiana: 70 n. mi.
N of Georgetown, ummm 2424 (10). Off Surinam: 65 n. mi. NE of
Paramaribo, USNM 185195 (1); 60 n. mi. NNW of Paramaribo, USNM
185328 (1). Off French Guiana: 90 n. mi. Nw of Cayenne, USNM
185047 (1); 80 n. mi. NNW of Cayenne, UsNM 185307 (2), UMim 2423
(2); 45 n. mi. NNE of Cayenne,usNM 185379 (3). Off Lesser Antilles:
St. Lucia, usnm 41281 (1); Martinique, usnmM 178626 (1). Off
Haiti: Port-au-Prince, usnmM 132545 (1), usnm 183695 (1). Off His-
paniola?: mcz 9982 (2).

Lutjanus vivanus (Cuvier and Valenciennes)
Silk snapper

Mesoprion vivanus Cuvier and Valenciennes, 1828, p. 343 (original descrip-
tion; depth of capture; size; common names; Martinique). Jordan,
1887, p. 534 (comments on types).

Mesoprion profundus Poey, 1860, p. 150 (original description; Cuba); 1861,
p. 365 (listed; common name; Cienfuegos, Cuba); 1866, p. 267 (com-
pared with aya = purpureus); 1867: 157 (compared with purpureus;
nuchal scales); 1868, p. 294 (common name; eye color; opercle; food
value; weight). Howell-Rivero, 1938, p. 196 (synonymy; type speci-
mens).

Lutjanus torridus Cope, 1871, p. 469 (original description; St. Kitts), fig. 5.
Hildebrand and Ginsburg, 1925, p. 77 (species not identifiable from
original description).

Lutjanus profundus, Poey, 1875, p. 28 (comments; compared with purpureus);

1962, p. 85 (compared with aya = purpureus; Cuba), pl. 70 A.

Lutjanus vivanus, Jordan and Fesler, 1893, p. 445 (common names; synonymy
in part; eye color; comments). Hildebrand and Ginsburg, 1925, p. 78
(comments on identity). Ginsburg, 1930, p. 265 (common names; de-
scription; variation; economic importance; relationship; characters in
key; nomenclature; synonymy; range; Campeche Bank), fig. 1.

Neomaenis vivanus, Jordan and Evermann, 1898: 1262 (common names; de-
scription; synonymy in part; Cuba). Evermann and Marsh, 1900: 175
(common names; description; synonymy; Puerto Rico).

Although lacking in most diagnostic characters the original de-
scription of vivanus obviously refers to one of the three species of
the complex reviewed in this study. The Gulf red snapper, cam-
pechanus, may be eliminated on the basis of locality since vivanus
was described from Martinique. The original locality, however,
is of no value in deciding whether the name vivanus applies to the

Rivas: Revision of Red Snappers 13]

silk snapper or to the Caribbean red snapper since both of these
species are sympatric. The conspicuous black margin of the caudal
fin and the depth of capture (90-100 fathoms) given by Cuvier and
Valenciennes for vivanus may be good clues. In the silk snapper
the black margin of the caudal fin is much more conspicuous than
in the Gulf and the Caribbean red snappers. Also the silk snapper
is usually taken along the edge of the shelf at depths of 80-120
fathoms whereas the other two species are usually taken at depths
of less than 80 fathoms (Camber, 1955, p. 23, and personal obser-
vations).

Because of the statements in reference to depth of capture and
the black margin of the caudal fin given in the original description
there is the strong possibility that the name vivanus refers to the
silk snapper rather than to the Caribbean red snapper, purpureus
(see also discussion under the latter species). Furthermore, the
name vivanus has been currently applied to the silk snapper and
for the sake of stability it is advisable to retain it for the silk
snapper.

Contrary to statements by Jordan and Swain (1885, p. 453),
Jordan (1887, p. 534), and others, no specimen that could be con-
sidered as the type of vivanus was mentioned by Cuvier and Va-
lenciennes in the original description. Therefore, there is no
specimen in existence whereupon the name vivanus could be posi-
tively verified or a type designated.

In view of the absence of original type material a specimen
from Mayaguez, Puerto Rico (closest available to type locality),
270 mm in standard length (usnmM 164632) is here designated as
the neotype of vivanus and described as follows.

Dorsal spines, 10. Dorsal rays, 14. Anal spines, 3. Anal rays,
8. Pectoral rays, 17. Lateral scales, 51. Scales above lateral line,
12; below, 22. Cheek scales in 7 rows. Gillrakers, 11 (plus 5 rudi-
ments); 2 (plus 5 rudiments), on upper limb. Predorsal length,
400. Preanal length, 726. Head length, 374. Snout length, 140.
Suborbital width, 66. Maxillary length, 145. Orbit diameter, 79.
Interorbital width, 82. Body depth, 342. Dorsal base length, 495.
Anal base length, 148. Pectoral fin length, 330. Pelvic fin length,
241. Anal fin length not measured (tip broken off). Middle cau-
dal rays length, 189. Scales on anterior side of body, below lateral
line, not conspicuously larger than those on posterior side. Scales
above lateral line, on anterior side of body, about equal to those

132 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

below. Lingual teeth in two patches, the anterior one much
smaller. Vomerine patch of teeth anchor-shaped, with a median
backward extension. General coloration yellowish-brown after
about 67 years of preservation. Tips of middle caudal rays black.

The original description of profundus (Poey, 1860, p. 150) refers
to this species, but the two specimens recorded as “cotypes” by
Howell-Rivero (1938, p. 196) are not the types of profundus. Poey
stated that his description was based on a single specimen 260 mm
(total length) but the specimens recorded by Howell-Rivero (mMcz
9966, 9990) have total lengths of 340 and 415 mm, respectively.
One of two specimens sent by Poey to the United States National
Museum and identified by him as profundus is almost certainly
the original type. This specimen (usNM 24796), here recognized as
the holotype of profundus, measures about 255 mm in total length
(upper caudal fin lobe frayed), 198 mm in standard length, and
bears Poey’s original cloth tag with his species No. 28.

The holotype of Lutjanus torridus Cope (1871, p. 469) from St.
Kitts, Lesser Antilles, 229 mm in standard length (ANSP 13225)
has been examined and found to be conspecific with L. vivanus.

This species differs from campechanus in the characters already
indicated under that species. As already discussed vivanus is more
closely relate dt opurpureus from which it differs in the characters
already discussed under the latter. The yellow eye is a good field
character to distinguish vivanus from campechanus and purpureus.

This species is known to occur along the edge of the shelves
bordering the southern Gulf of Mexico, the Straits of Florida, and
the Caribbean Sea. North of the Yucatan Channel vivanus occurs
sympatrically with campechanus. In the Caribbean Sea it occurs
sympatrically with purpureus.

As already discussed under campechanus and purpureus ab-
sence of a shelf on both sides of the Yucatan Channel and of the
Windward Passage could be interpreted as a barrier to the south-
ward dispersal of campechanus and to the northward dispersal of
purpureus. This need not be so in the case of vivanus since avail-
able collecting data and records from the literature (Camber, 1955:
23) indicate that this species usually occurs at depths of 80 to 100
fathoms or more.

Material examined. 18 specimens from the following 12 lo-
calities. West Indies: no specific locality, usnM 33264 (1). Off
Jamaica: no specific locality, usnmM 37730 (1). Off Puerto Rico:

Rivas: Revision of Red Snappers 133
Mayaguez, usnM 164632 (1). Off Lesser Antilles: St. Kitts, ansp
13225 (1). Off Panama: 18 n. mi. ENE of Punta Manzanillo, umim
6100 (1); 4 n. mi. N of Cabo Tiburon, umm 6097 (1). Off Cuba: no
specific locality, usnm 12557 (2); Bahia Honda, usnm 82436 (1);
Havana, usnM 24782 (1), usnm 24796 (1), usnM 25010 (1). Off Flor-
ida: Marathon, ummm 6010 (1); Miami ummm 6011 (1); Port Ever-
glades, ummm 2626 (3), umm 2712 (1).

TABLE 1

Comparison of similar length specimens of Jutjanus campechanus complex

L. campechanus L. purpureus L. vivanus

(No. = 21) (No. = 10) (No. = 9)
Character! Range Mean Range Mean Range Mean
Standard length 190-418 296 160-435 SL5 198-420 266
Head length 376-411 392 338-397 361 356-377 369
Snout length 146-161 151 121-147 131 127-140 135
Suborbital width 80-89 85 65-70 68 64-73 67
Maxillary length 145-158 152 129-150 138 123-146 138
Mandible length 180-195 187 160-186 2 170-177 174
Body depth 350-405 376 312-381 354 320-354 337
Pectoral fin length 316-343 329 287-313 301 310-339 32]
Pelvic fin length 215-250 233 189-224 205 222-246 233
Anal fin length 268-319 288 241-288 262 243-284 256

1 Standard length in millimeters; other characters expressed in thousandths
of the standard length.

TABLE 2

Frequency distribution of lateral scales in the Lutjanus campechanus complex

Lateral scales

Species INOw Oe 47 ee aor 49) 50S 510) 52). 53 Mean
L. campechanus 76 Ce la 26 ne 22 8 48.2
L. purpureus 38 Cy Ses iG 3 50.4
L. vivanus 18 See all 2 Dy ASMP

134 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 3

Frequency distribution of scales above lateral line and cheek scale rows
in Lutjanus campechanus complex

Scales above lateral line Cheek scale rows
Species No. 7 8 91011 12 Mean No. 5S) )655/GeMicam
L. campechanus 76 4 42 25 5 8.4 G2 SO 2iaan 5.9
L. purpureus 38 Ge2i el 10.1 3112} Slee 6.1
L. vivanus 18 295 ee) Males 18 17 lee
TABLE 4

Frequency distribution of scales below lateral line in
Lutjanus campechanus complex

Scales below lateral line

Species No. 15. 16 17 18 19 20 21. 222324 ies

L. campechanus CS S20, 425 lO) Gea

L. purpureus 37 Saal hOgess 17.9

L. vivanus 7 2 3 Gy) Ano
TABICn 5

Frequency distribution of anal rays and of gill rakers on lower limb of
first arch in Lutjanus campechanus complex

Anal rays Gillrakers

Species No. 8 9 Mean No. 8 9 10). 02s Mean

Eycampechanus, 126 “147 AD 819 80 2 28° 48 “222 RoiG
L. purpureus AN Be) OF oe? 35 L 14 TO SiesOG
L. vivanus Ses 8.0 18 l 4 2 aieeslOes

Ul

Rivas: Revision of Red Snappers 13

LITERATURE CITED

Biocu, M. E. 1790. Naturgeschichte der auslandischen Fische. Pt. 4, 128
DPM Oley —2'52.,

Biocu, M. E., AND J. G. ScHNEmeER. 1801. M. E. Blochii . . . systema
ichthyologiae iconibus ex illustratum, 584 pp., 10 pls. Bibliopolio San-
deriano comm. Barolini.

Camber, C. I. 1955. A survey of the red snapper fishery of the Gulf of
Mexico, with special reference to the Campeche Banks. Florida State
Dept. Conserv., Techn. Ser., no. 12, 63 pp., 14 figs.

CARPENTER, J. S. 1965. <A review of the Gulf of Mexico red snapper fishery.
Fish and Wildl. Serv., Bur. Comm. Fish., Circ. no. 208, 35 pp., 26 figs.

Corr, E. D. 1871. Contribution to the ichthyology of the Lesser Antilles.
Trans. American Philos. Soc., vol. 14, new ser., pt. 3, pp. 445-483,
10. figs.

Cuvier, G., AND A. VALENCIENNES. 1828. Histoire naturelle des poissons,
VOle2, S71 pp., pls. 9-40. Paris.

EVERMANN, B. W., ann M. C. Marsw. 1900. The fishes of Porto Rico.
Bull. U. S. Fish Comm., pp. 51-350, 49 pls., 3 maps.

GinssurG, I. 1930. Commercial snappers (Lutianidae) of Gulf of Mexico.
BullU. Ss; Bur. Fish., vol. 46, pp. 265-276, 2 figs.

Goope, G. B., ANp T. H. BEAN. 1879. Descriptions of two new species of
fishes, Lutjanus blackfordii and Lutjanus stearnsii, from the coast of
londaweeroc, U.S. Nat. Mus., vol. 1, pp. 176-177.

HILDEBRAND, S. F., AND I. GinsBpurc. 1925. Distinguishing characters of two
species of red snappers of the Atlantic coast of North America. Bull.
UneSmebureiaish.., vol, 42, pp. 77-85, 3 figs.

Howe.i-Rivero, L. 1938. List of the fishes, types of Poey, in the Museum
of Comparative Zoology. Bull. Mus. Comp. Zool., vol. 82, no. 3,
pp. 169-227.

Jorpan, D. S. 1887. Notes on typical specimens of fishes described by
Cuvier and Valenciennes and preserved in the Musee d'Histoire Nat-
urelle in Paris. Proc. U. S. Nat. Mus., vol. 9, pp. 525-546.

JorpAN, D. S., AND B. W. EverMANN. 1898. The fishes of North and Mid-
dle America. Bull. U. S. Nat. Mus., no. 47, pt. 2, pp. 1241-2183.

Jorpan, D. S., ANd B. Fesuter. 1893. A review of the sparoid fishes of
America and Europe. Rep. U. S. Comm. Fish and Fish. (1891), pp.
421-544, pls. 28-62.

136 QuvaARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Jorpan, D. S., AND J. Swarn. 1885. A review of the species of Lutjaninae
and Haplopagrinae found in American waters. Proc. U. S. Nat. Mus.,
vol. 7, pp. 427-474.

Pory, F. 1860. Poissons de Cuba, especes nouvelles. Mem. Hist. Nat.
isla de Guba, vols 2-art. 49, pp. 115-356) plshl2=n9)

1861. Conspectus Piscium Cubensium. Mem. Hist. Nat. Isla de
Cuba, vol. 2; art. 50, pp. 357-404.

—. 1866. Revista de los tipos Cuvierianos y Valenciennianos correspon-
diendo a los peces de la Isla de Cuba. Rep. Fisico-nat. Isla de Cuba,
V.Olnl ep pailia=2o:

1867. Revista de los peces descritos por Poey. Rep. Fisico-nat.
Isla de Cuba, vol. 2, pp. 153-174.

—. 1868. Synopsis Piscium Cubensium. Rep. Fisico-nat. Isla de Cuba,
vol. 2, pp. 279-484.

——. 1875. Enumeratio Piscium Cubensium. Anal. Soc. Espanola Hist.
Nat., vol. 4, 87 pp., 3 pls.

1962. Ictiologia cubana, vol. 1, 106 pp., 87 pls. Acad. Sci. Re-
publica de Cuba, Habana.

Rivas, L. R. 1960. The fishes of the genus Pomacentrus in Florida and the
western Bahamas. Quart. Jour. Florida Acad. Sci., vol. 23, no. 2, pp.
130-162, 9 figs.

——. 1964. A reinterpretation of the concepts “sympatric” and “allo-
patric” with proposal of the additional terms “syntopic” and “allotopic.”
Syst. Zool., vol. 18, no. 1, pp. 42-48.

Department of Biology, University of Miami, Coral Gables,
Florida. Contribution No. 57 from the Ichthyological Laboratory

and Museum.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

Centrarchid Spawning in the Florida Everglades
James P. CLucsron

KNOWLEDGE of the spawning characteristics and requirements
of freshwater fishes is important to present day fish management.
Predictions of future fishing success are sometimes made possible
by knowledge of spawning activities. In some instances fish pop-
ulations have been regulated by encouraging or discouraging the
spawning of certain species. Water levels kept at the proper height
during the spawning season of the shore-spawning species will
permit a normal hatch while a drawdown can be used to suppress
a population when it has become overcrowded. Treatment of the
margins of a lake with a fish toxicant during spawning periods
can also be used as an aid in reducing the numbers of an undesir-
able species.

Peninsular Florida is almost unique compared to the rest of the
continental United States because much of it is subtropical and
has little seasonal fluctuation in temperature. Old residents tell
of year-round spawning of bass and the smaller sunfishes, but
little information appears in the literature to establish the actual
spawning periods of these species. Available information on cen-
trarchid spawning in Florida is from the central portion of the
state; an area approximately 250 miles north of the study area
described in this paper.

THE Stupy AREA

In 1955 the Florida Game and Fresh Water Fish Commission
undertook a number of studies to learn more about the fish popu-
lations and other aquatic life within three Conservation Areas in
southern Florida located approximately 20 miles west of the
Miami-Fort Lauderdale-West Palm Beach region. These three
areas, totaling approximately 1,345 square miles, are a portion of
the original Florida Everglades that is now enclosed by a vast
network of levees and canals. The Conservation Areas were cre-
ated to retain surplus water during the hurricane and rainy season,
to regain irrigation water supplies in time of drought, to serve as
potential reservoirs to maintain urban ground water tables, and to
provide for fish and wildlife. These marshes and their bordering
canals are the primary freshwater fishing areas of south Florida.

138 QuvuaRTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The majority of the investigations by the Game and Fresh
Water Fish Commission were carried out in Conservation Area
2, a tract of 137,000 acres. The ground elevation slopes from
about 13 feet to between 7-10 feet above mean sea level from
north to south. Because of the gradual slope over a 15-mile stretch,
a change of one foot in water level can either inundate or dry
vast areas of marsh. The water depths in the southern portion of
Area 2 range between 2-5.5 feet. The water level stages are es-
tablished and controlled by the Central and Southern Florida Flood
Control District.

SPAWNING SURVEY

Observations of spawning were made between December 1960
and July 1964 in Conservation Area 2. The levees around this
area provided a berm or shelf of rock and clay on certain parts of
which centrarchid nests were easily visible. While driving slowly
along the levees, a biologist wearing polaroid glasses could look
down without leaving the car and observe fish on the levee berm.
The species observed spawning were: largemouth bass (Micropterus
salmoides), bluegill (Lepomis macrochirus), redear sunfish (Lepo-
mis microlophus), and spotted sunfish (Lepomis punctatus). Other
centrarchids that were present in the Conservation Areas, but not
observed spawning on the levee berm were: warmouth (Chaeno-
bryttus gulosus), dollar sunfish (Lepomis marginatus), bluespotted
sunfish (Enneacanthus gloriosus), black crappie (Pomoxis nigro-
maculatus), and the Everglades pygmy sunfish (Elassoma ever-
gladei).

During the period between December 1960 and September
1961, counts of bass nests were made over a 10-mile section of
levee berm (Table 1). The marsh extended directly to this levee
and because of the thick vegetation, only limited sections of the
berm were visible. Counts were made in the same sections each
time. The completion of an additional levee in September caused
the area where the counts were made to become dry, consequently
losing its fishery value. Thereafter, observations were made along
a new interior levee to the north. The new levee had a 125-foot
wide canal adjacent to it which made observations possible along
its entire length. However, actual counts were impractical because
of wind action on the water surface and the great numbers of
spawning fish at certain times of the year. Therefore, only gen-

CLucsTon: Spawning of Centrarchids 139

eral notes regarding the relative abundance of each species were
made.
TABLE 1

Counts of bass nests along 10 miles of levee in Conservation Area 2,
Broward and Palm Beach counties, Florida. Water temperature
was recorded 12 inches below the surface at mid-day

Date Nests with bass Nests without bass Water temperature
1960
Dec. 4 0 0 een Dec. yal) >
Dec. 14 0 0 G2 (Deckse19)™ *
1961
Jan. 26 Numerous GORE (iante23) 4
Feb. 10 56 63 Omi (Kel. 37)" ?
Feb. 16 104 33 65 F
Feb. 28 50 102 74 F
Mar. 3 20 35 16 Es
Mar. 10 28 35 64 F
Mar. 16 67 3 70 F
Mar. 23 13 14 TS a
April 5 26 i 78 F
April 13 25 = (Ae
April 18 12) ‘ 80 F
May 3 82 F
through 0 0 to
Aug. 31 92 F

*It was not possible to discern between vacant bass nests and_ nests
vacated by other centrarchids on these dates.

**Frish, Harry M. 1962. Annual water quality variations at certain
points in the Everglades. Florida Game and Fresh Water Fish Comm., Talla-
hassee, 14 pp. (mimeographed report).

SPAWNING

Largemouth Bass. The largemouth bass is the first of the cen-
trarchids to spawn each year in south Florida. Spawning usually
commenced when the water cooled to the low 60s F. During
the period of the investigation, these temperatures occurred be-
tween mid-December and mid-January. Clugston (1964) reported
successful spawning of largemouth bass between November 15 and
December 15, 1960, in a 1.2-acre pond located approximately 20
miles east of the Conservation Areas. February was the peak
spawning month during every year of the study. Most bass spawn-

140) QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ing occurred when the water temperature neared 70 F, and all
spawning appeared to stop when the water temperature rose above
80 F. The bass ceased spawning along the levees by April Ist
in 1960, 1962, and 1964, and by May Ist in 1961 and 1963. The
spawning season ranged between 2.5-4.0 months in lengths. The
general spawning pattern observed each year corresponded closely
with the 1960-1961 season shown in Table 1. The location of
nests at least 10 feet from each other showed the characteristic
territorial behavior of the largemouth bass. Most of the bass nests
were observed in water 12-30 inches deep.

Bluegill and Redear Sunfish (Shellcracker). The spawning re-
quirements and habits of these two species are quite similar.
Therefore, it was not always possible to distinguish which of the
two species was being observed. Generally the bluegill and red-
ear sunfish started spawning in late February or early March when
the water temperature approached 70 F. Nests were most abun-
dant when the water temperature was between 75-80 F, but on
a number of occasions, they were observed spawning when the
water temperature was 90 F. Spawning along the levee appeared
to cease by October 1. The spawning season normally lasted 6-7
months, although during this period, there might often be times
when they ceased spawning for 1-3 weeks.

Counts were made on three occasions in 1964 to illustrate the
great numbers of the smaller sunfishes spawning along the in-
terior levee. The total number of nests observed were: 1,284 on
April 10; 1,373 on April 24; and 724 on June 9. These counts in-
cluded bluegill, redear sunfish, and spotted sunfish. In April the
mid-day water temperature was 75-77 F. By June 9, the tempera-
ture had risen to 86 F.

The bluegill and redear sunfish are community spawners. Fre-
quently, when the nests were only a few inches from each other,
the area appeared as one large nest 6 or more feet in diameter.
The nests were usually found in water 18 to 36 inches deep.

Spotted Sunfish (Stumpknocker). The spawning season of the
spotted sunfish in the Everglades is difficult to define. This species
spawned intermittently throughout the spring and summer and
until November in 1963. They were found spawning in small
numbers when the mid-day water temperature was 64 F. More
commonly, they started to spawn in fair numbers when the water
temperature neared 75 F in March or April. The preferred spawn-

Ciucston: Spawning of Centrarchids 141

ing temperature range appeared to be 80-85 F. Spotted sunfish
were observed spawning in great numbers on August 31, 1961,
when the water temperature was 92 F. However, during the same
year, no spotted sunfish could be found spawning from mid-June
until the end of July when the mid-day water temperature ranged
between 86-89 F.

This species generally fanned out its nests within one foot of
the shore in water that was only 6 inches deep. They were the
most pugnacious of the centrarchids observed. Bass, bluegill,
and redear sunfish would usually flee the nest and wait in the
shadows until the observer left the area before returning to the
bed. The spotted sunfish would dart away from the nest, but
return almost at once to stay over the eggs. On a number of oc-
casions, they returned while the observer was holding a _ther-
mometer in the nest.

DIscussroN

Great numbers of largemouth bass, bluegill, redear sunfish, and
spotted sunfish were observed spawning along the levees of Con-
servation Area 2. The observed maximum spawning seasons varied
from 4 months for the largemouth bass to 8 months for the spotted
sunfish. November was the only month in which no centrarchid
spawning was observed.

It is known that both light and temperature play an important
part in controlling the reproductive cycle of many fishes. Cen-
trarchids normally are predisposed to spawn during a lengthening
photoperiod and increasing temperature. Many have observed that
bass spawning occurs first in the spring when the water tempera-
ture warms to 60-65 F. Bass spawning may start in mid-May in
Wisconsin (Mraz, Kmiotek, and Frankenberger, 1961) and in mid-
April in Alabama (Swingle and Smith, 1950). In central Florida
(latitude 28-29° N) it appears that bass spawning occurs in March
and April (Carr, 1942; Horel, 1951; McLane, 1955) although they
have been observed spawning “as early as January’ in the St.
Johns River (McLane, op. cit.). In south Florida (latitude 26°
N.) bass spawning commenced in the coolest period of the year,
i.e., during December or January. The water temperature appar-
ently must drop to nearly 60 F, or approximately the same tem-
perature that the water must warm to in the cooler latitudes, be-
fore spawning begins. This is also the period of the least day-

142 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

light hours. However, bass spawning increases as the daylight
hours become longer and until the water temperature increases
to around 70 F.

Aronson (1957), in reviewing the work of other investigators,
discusses many ecological factors which may affect the reproduc-
tive habits of fishes. He breaks these into three major headings:
meteorological, habitat, and social. The more irregular spawning
described for the smaller sunfishes suggests that factors in addi-
tion to photoperiod and temperature contribute to their reproduc-
tive pattern although no specific items were detected in this study.

The depth of the water in most of the marsh is not too deep
for centrarchid spawning anytime throughout the year. Because
of the dark bottom (peat) of the marsh and the thick vegetation,
it was not possible to determine the degree to which the many
centrarchid species used the marsh for spawning. All of the
species observed spawning on the berm were also seen spawning
on the peat or roots of the vegetation in the marsh. The great
numbers of fry of all centrarchids that were found many miles
from the levees while taking population estimates (Clugston and
Dineen, unpublished data) indicate that spawning occurred over
the entire area.

Lowering of the water level occurs during much of the spawn-
ing season, but it occurs at such a slow rate that it does not appear
to affect the spawning adversely in any way. During May and
June, when the water level was at its lowest, all species except
the largemouth bass were seen spawning. However, from April
through August, bass fry 1-1.5 inches long were observed in the
marsh a number of times. Perhaps, because of the greater water
depths and the shade afforded by the dense vegetation, the water
may remain sufficiently cool in the marsh to encourage bass spawn-
ing at a date later than actually observed.

ACKNOWLEDGMENTS

The author wishes to acknowledge the assistance of the fol-
lowing individuals who participated in the spawning survey:
J. Walter Dineen, Walter M. Tatum, Forrest Ware, Loren Flagg,
and William Lindall. |

This paper is a contribution from Dingell-Johnson Project
F-16-R, Florida.

CLucston: Spawning of Centrarchids 143

LITERATURE CITED

Aronson, L. R. 1957. Reproductive and parental behavior. In: The physi-
ology of fishes. Vol. 2, pp. 271-303. Academic Press, New York.

Carr, Marjorie H. 1942. The breeding habits, embryology and _ larval
development of the largemouth black bass in Florida. Proc. New
England Zool. Club, vol. 20, pp. 43-77.

CiucsTon, JAMEs P. 1964. Growth of the Florida largemouth bass, Microp-
terus salmoides floridanus (LeSueur), and the northern largemouth bass,
M. s. salmoides (Lacépéde), in subtropical Florida. Trans. Am. Fish.
Soc., vol. 93, no. 2, pp. 146-154.

Horet, Georce J. 1951. The major bedding areas of largemouth black
bass in Lake George, Florida. Florida Game and Fresh Water Fish
Comm., Tallahassee, 10 pp., (mimeographed report).

McLane, Wituiam M. 1955. The fishes of the St. Johns River system.
Unpublished doctoral thesis, University of Florida, 361 pp.

Mraz, D., S. KmI1oTEK, AND L. FRANKENBERGER. 1961. The largemouth
bass; its life history, ecology and management. Wis. Cons. Dept.
Publication 232, 15 pp.

SwWINGLE, H. S., AND E. V. SmitH. 1950. Factors affecting the reproduction
of bluegill bream and largemouth black bass in ponds. Ag. Exp. Sta.

Ala. Poly. Inst. Bull., vol. 87, pp. 1-8.
Game and Freshwater Fish Commission, Leesburg, Florida
(present address: U. S. Bureau of Commercial Fisheries Biological

Laboratory, Galveston, Texas).

Quart. Jour. Florida Acad. Sci. 29(2) 1966

The Exotic Herpetofauna of Southeast Florida
WAYNE KING AND THOMAS KRAKAUER

In 1910 Barbour reported the first successful invasion of south
Florida by an exotic species of amphibian, Eleutherodactylus plan-
irostris, and in 1922 Stejneger reported the first exotic reptiles,
Hemidactylus turcicus and Sphaerodactylus cinereus, to be estab-
lished in the state (see also Fowler, 1915). By 1940 the list of
exotics had increased to eight species (Carr, 1940), and by 1957 to
twelve (Duellman and Schwartz, 1958). The introductions come
from three sources, accidental importation on produce shipped
into the state, escape from wholesale animal dealer-importers, and
the intentional release of animals. The last two sources account
for the rapid increase in the rate of herpetofaunal introductions
since 1945. The success of many of the introductions results from
south Florida's semitropical climate and flora and depauperate
fauna.

This paper lists the exotic species that have been introduced
into south Florida, delimits their present local ranges, and records
how each was introduced, when known. The animals are treated
in three groups, species established and breeding, species not known
to breed locally, and species unreported since their release.

ESTABLISHED BREEDING SPECIES

The giant toad, Bufo marinus, was first reported in southeast
Florida by Neill (1957). Duellman and Schwartz (1958) listed
B. marinus in the “ ... western part of Miami, Dade County, 17
May 1955.” The present population is not the result of an intro-
duction near Pennsuco prior to 1958 (see Duellman and Schwartz,
1958; Riemer, 1959), but results from the accidental release of
approximately 100 specimens by an importer formerly located at
Miami International Airport, whence the species has spread. Other
animal dealers deliberately released the species in 1963 at Pem-
broke Park, Broward County, and in 1964 at Kendall, Dade County.
The Kendall animals came from Surinam; all others were from
Colombia. The present distribution of B. marinus in Florida
extends from Homestead in southern Dade County, north to Pem-
broke Park and Hollywood in Broward County, and from the
Intracoastal Waterway west to the Everglades.

Kinc AND Krakauer: Introduced Herpetofauna 145

The greenhouse frog, Eleutherodactylus planirostris planiros-
tris, from Cuba, has been recorded throughout most of south Flor-
ida (Barbour, 1910; Deckert, 1921; Carr, 1940; Goin, 1947; Neill,
1957; Duellman and Schwartz, 1958). It is abundant throughout
the Florida Keys and the eastern half of Dade, Broward, and
Palm Beach counties. Its introduction was probably an accidental
consequence of ship commerce.

The giant treefrog, Hyla septentrionalis, from Cuba and His-
paniola, occurs from Key West in southern Monroe County, where
it was first recorded, northward along the eastern half of Dade and
Broward counties to Dania (Barbour, 1931; Trapido, 1947; Wright
and Wright, 1949; Peterson, Garrett, and Lantz, 1952; Schwartz,
1952; Allen and Neill, 1953; Duellman and Schwartz, 1958; King,
1960). Eggs and tadpoles of this frog are found in many outdoor
fishponds and swimming pools in southeast Florida. Its initial
introduction seems to be an accident of shipping.

The yellow-headed gecko, Gonatodes albogularis fuscus, was
established in Key West prior to 1939 (Carr, 1939, 1940; Duellman
and Schwartz, 1958) and is still abundant there today. By 1965
a reptile fancier had secondarily introduced it into Dade County,
in the vicinity of Day Avenue and Matilda Street, Coconut Grove.
This species probably was introduced to Key West on shipments
from Cuba or Jamaica.

The Indo-Pacific gecko, Hemidactylus garnoti, was introduced
into Dade County prior to 1964. This species is established in two
widely separated locations, in the immediate vicinity of 3811 Wood
Avenue in Coconut Grove, and 19 Terrace between 70 and 73
Courts SW, Miami. As both localities are homes of University of
Miami Institute of Marine Science personnel, the introduction
may have resulted from the International Indian Ocean Expedition
(1960-1963).

The Mediterranean gecko, Hemidactylus turcicus turcicus, has
been reported primarily from Key West and Big Pine Key, Monroe
County (Fowler, 1915; Stejneger, 1922; Barbour, 1936; Duellman
and Schwartz, 1958; King, 1959). In the Miami area it occurs
in several localities, 22 Street and 3 Avenue NE, and along NW
South River Drive between 42 and 27 Avenues NW. Whether
these are isolated populations, or part of a single widespread pop-
ulation, is not known to us. A reptile fancier established one dis-
junct colony in the vicinity of 9885 SW 80 Drive, Sunset Park.

146 QvaRTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The mainland populations are undoubtedly secondary introduc-
tions from the Key West colony, and the latter probably was an
accident of shipping.

The ocellated gecko, Sphaerodactylus argus argus, was re-
ported from Key West, Monroe County, by Savage (1954). Duell-
man and Schwartz (1958) were unable to find the colony in Key
West, and therefore reported it no longer extant. The colony,
however, was located on the western end of the island near the
aquarium as recently as 1964. These animals could have been
introduced in shipments from Jamaica or North Bimini, Bahamas.

The ashy gecko, Sphaerodactylus cinereus, a Cuban and His-
paniolan species, was first reported from Key West by Stejneger
(1922). Duellman and Schwartz (1958) report that the colony is
thriving on Key West, and that S. cinereus also occurs on Boca
Chica Key, Monroe County. These are the only populations known
to us. While S. cinereus is restricted in distribution, Sphaerodac-
tylus notatus is widespread both in the Florida Keys and on the
mainland. These distributions led Duellman and Schwartz (1958)
to believe that S. notatus is an indigenous member of the Florida
fauna, a conclusion with which we agree.

The Bahamian bark anole, Anolis distichus distichus, was in-
itially discovered in Brickell Park, Miami, in 1946 (Smith and Mc-
Cauley, 1948; Duellman and Schwartz, 1958). This lizard is abun-
dant in the Miami area from the Miami River south to Kendall,
and from the Atlantic shore to West Miami. In addition, it occurs
on Key Biscayne in the area of Crandon Park Zoo. It is most
abundant in Coconut Grove and Coral Gables. The first popula-
tion almost certainly was a deliberate release. In Coconut Grove
the species has spread without human aid, but virtually all other
populations result from secondary introductions by reptile fanciers.
In life, this race has an ash-gray dorsal groundcolor and a pale
yellow dewlap.

The green bark anole, Anolis distichus dominicensis, a Hispan-
iolan race, occurs in a small colony in Miami, along the Tamiami
Canal near 32 Avenue and 24 Street-Road NW. It was most prob-
ably accidentally introduced on a freight boat that trades be-
tween the Miami River and Hispaniola. In life, this race has a
pea-green or gray-green dorsal groundcolor and a pale orange
dewlap.

The knight anole, Anolis equestris equestris, from Cuba, was

KiNG AND KRAKAUER: Introduced Herpetofauna 147

reported in south Florida by Neill (1957). The main colony, and
area of first introduction, is centered in a twenty city block area
of Dade County, Coral Gables, from Coral Way in the north to
Bird Road in the south, and from Le Jeune Road on the east to
Segovia Avenue on the west. Secondary releases occurred in
Coconut Grove in the vicinity of Day Avenue and Matilda Street;
in Coral Gables on Old Cutler Road near Matheson Hammock;
in Sunset Park in the vicinity of 9885 SW 80 Drive; and in Peters
at US Highway 1 and Eureka Drive. It is also reputed to be on
Key Biscayne; in Brickell Hammock on the grounds of the Viscaya
Museum; and in Broward County, in the vicinity of Davie, but
this has not been confirmed by us. The original introduction was
made in 1952 by a student in the University of Miami Department
of Biology, whose buildings are in the center of the main popula-
tion. The subsequent spread was accomplished by reptile fan-
ciers, who feared that the species would become “extinct” in
Miami.

The Bahamian ground anole, Anolis sagrei ordinatus, was re-
ported to occur in a six block area adjacent to Shenandoah Ele-
mentary School at 1023 SW 21 Avenue (Bell, 1953). This locality
now supports a colony of Anolis sagrei sagrei. Duellman and
Schwartz (1958) do not list A. s. ordinatus from the Miami area,
and we suggest that if A. s. ordinatus occurred at Shenandoah
School, it was never abundant there. Oliver (1948; see also Carr
and Goin, 1955) reported A. s. ordinatus from Lake Worth city,
Palm Beach County. A small colony also exists in South Miami,
in the vicinity of US Highway 1 and 61 and 62 Avenues SW.
Although the Lake Worth introduction was deliberate, the South
Miami population seems to have been derived from accidental
introduction on Bahamian shells and corals sold at that location.
In life, males of this race have a pale brown to ash-gray dorsal
groundcolor and a dewlap with a mustard-yellow groundcolor.

The Cuban ground anole, Anolis sagrei sagrei, has been col-
lected from Key West, Miami, Tampa, St. Petersburg, and West
Palm Beach (Barbour, 193la; Oliver, 1950; Duellman and Schwartz,
1958; King, 1960; see also Ruibal, 1964, for the synonymy of
A. sagrei stejnegeri with A. s. sagrei). This species abounds through-
out most of metropolitan Dade, Broward, and Palm Beach coun-
ties, and in Key West and Cudjoe Key, Monroe County. Its pres-
ent distribution is the result of at least three separate accidental

148 QvuaARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

introductions into Florida ports—Key West prior to 1931; Port
of Palm Beach, 1960; Port Everglades (Broward County), 1964.
From these initial populations, the species has spread by secondary
releases and by its own abilities to compete and increase its
numbers until today it is one of the most successful reptiles in
south Florida. In life, males of this race have a rust-brown to
ash-gray dorsal ground color and a dewlap with a red ground
color.

The curly-tail lizard, Leiocephalus carinatus armouri, from the
islands of the Little Bahama Bank, was first reported in Palm Beach
County (Dueliman and Schwartz, 1958; King, 19€0). Another race,
L. c. virescens, occurred in Miami prior to 1940 (Barbour, 1936;
Carr, 1940), but died out shortly thereafter. Colonies of L. c.
armouri occur in Dade County on the grounds of iMami Seaquar-
ium on Virginia Key, and Crandon Park Zoo on Key Biscayne.
Both colonies were produced by deliberate releases by the per-
sonnel of these institutions. In addition, L. carinatus coryi from
Bimini, Bahamas, is believed to be present at Crandon Park Zoo,
although we have been unable to collect any. The Palm Beach
population is still extant.

The Colombian ground lizard, Ameiva ameiva petersi, was first
reported by Duellman and Schwartz (1958; see also Neill, 1957)
from 34 Avenue and 79 Street NW, Miami. This colony began
when animals escaped prior to 1954 from an animal importer for-
merly located at that locality. They are believed to have come
from Colombia. Since that time, the colony has occupied an
area of about 25 city blocks, from 79 Street NW, Miami (E 25 St.,
Hialeah), south to 76 Street NW, Miami (FE 22 St., Hialeah), and
from 36 Avenue NW, Miami, west to E 8 Avenue, Hialeah (Le
Jeune Rd.). This is an area of weed-filled vacant lots, railroad
right-of-ways, and residential lawns and gardens.

ESTABLISHED NON-BREEDING SPECIES

The second group includes established species for which we
have no evidence of breeding at the time of this writing. These
colonies may exist only as long as the adult animals live, or evi-
dence of breeding might be found at some later date.

The red-eared turtle, Pseudemys scripta elegans, from the
northern United States, is concentrated in a small canal in Miami

KinG AND KRAKAUER: Introduced Herpetofauna 149

bordering the Little River Canal, located at 107 Street at 18 Avenue
NW. Adult specimens were collected for us by children from
this locality. Another colony exists in the canals in the area of
Crandon Park Zoo, Key Biscayne. Both colonies resulted from
deliberate releases.

The spectacled caiman, Caiman sclerops, is found as a feral
pet in the various canal systems in and adjacent to cities in south
Florida. It has been found as far north as Palm Beach County.
It appears to successfully overwinter, but nests have not been
found. This species is imported in large numbers from Colombia,
and is a common item in many tourist shops throughout Florida.

The Tokay gecko, Gekko gecko, from southeast asia, is present
in the immediate vicinity of 3310 NW South River Drive, Miami.
At least three specimens of unknown sex were released prior to
1965 by an animal importer at this locality in an attempt to control
roaches in the buildings. In addition, four or five specimens were
released by a reptile fancier in the vicinity of Day Avenue and Ma-
tilda Street, Coconut Grove, although we have been unable to
discover the species at this locality.

This species has also been introduced into north Florida. In
August 1963, a professor in the University of Florida released
two adult females and one adult male Gekko gecko on his house
in Gainesville, Alachua County, Florida. Two more of unknown
sex were released October 1964. They survived two winters with
a low temperature of —11C. As of April 1965, at least one male
and one female were alive, but there was no evidence of young.

The rhinoceros iguana, Cyclura cornuta cornuta, from Hispani-
ola, was introduced, as part of an exhibit, to several “islands” at
Miami Seaquarium, Virginia Key, Dade County. Several of these
animals have escaped from this exhibit and are frequently seen
running around the grounds of the Sequarium. Additional speci-
mens reportedly have been seen on nearby Key Biscayne.

The green iguana, Iguana iguana iguana, is a frequent escapee
from animal dealers and reptile fanciers. The species appears
to survive and grow quite well, but the occasional winter frost
probably eliminates most of the population. Areas of concentra-
tion in the Miami area are near W 27 Street and E 7 Avenue,
Hialeah; in the vicinity fo Caballero Boulevard and Hardee Road,
Coral Gables; in the southwest corner of Miami International Air-
port; and on Key Biscayne. At the first of these localities over

15Q QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

300 individuals were released by one animal dealer between May
and September 1964. Most of these animals are imported from
Colombia.

The Texas horned lizard, Phrynosoma cornutum, from the
southwestern United States, appears occasionally in south Florida.
The species has been recorded from Dade, Duval, Escambia, In-
dian River, Lake, Marion, Orange, Polk, and Putnam counties
(DeSola, 1934; Goff, 1985; Carr, 1940; Carr and Goin, 1955; Allen
and Neill, 1955). The species was successfully breeding in north
Florida on Fort George Island, Duval County, as of September
1953. Several individuals were released by a reptile fancier prior
to 1964 in Dade County, in the vicinity of 9885 SW 80 Drive, Sun-
set Park, and over 100 specimens were released between May and
September 1964 by an animal dealer in Hialeah, near W 27 Street
and E 7 Avenue. Finally, a small colony near NW South River
Drive and 33 Avenue, Miami, resulted when animals escaped
from an animal dealer at that address. One specimen was also
collected in southwest West Palm Beach, Palm Beach County.

The South American ground lizard, Ameiva ameiva ameiva,
is established in Dade County, Kendall, in the immediate vicinity
of 78 Avenue and 125 Street SW. Approximately eight adults
were imported from Surinam by an animal dealer and released
near his house prior to 1964. Numerous reports place this species
on Elliott and Little Arsenicker Keys, Dade County, but we have
repeatedly been unable to obtain specimens from these islands.
Six specimens were also released in 1964 by an animal dealer in
Hialeah near W 27 Street and E 7 Avenue.

The boa constrictor, Constrictor constrictor, is collected infre-
quently in cities in south Florida. These snakes are undoubtedly
feral pets, although one reptile fancier released juveniles along
Loop Road (State Highway 94) in an attempt to establish it in the
Everglades. Nowhere, to our knowledge, is there a center of con-
centration for this species. Several races are imported by the
animal dealers in south Florida.

SPECIES UNREPORTED SINCE RELEASE

The third group consists of species which were released in
sufficient numbers to have become established, but which have
not been found by us since. As the list of species is long and the

KING AND KraAkAvuErR: Introduced Herpetofauna 151

number of localities is small, we will list these by locality. The
approximate number of individuals released is given preceding
each species name.

In the vicinity of W 27 Street and E 7 Avenue, Hialeah, includ-
ing the adjacent Hialeah (Red Road) Canal, an animal importer
at that address released the following species between May and
September 1964: saLaAMANDER:—200 Notophthalmus viridescens
viridescens, 200 Triturus pyrrhogaster; rRoG:—1000 Hymenochirus
boettgeri, 200 Xenopus laevis, 10 Agalychnis dacnicolro; ruRTLES—
25 Kinosternon scorpioides, 25 Chrysemys picta belli, 25 Chrysemys
picta dorsalis, 100 Graptemys pseudogeographica kohni, 20 Pseu-
demys dorbigni, 20 Pseudemys scripta callirostris, 150 Pseudemys
scripta ornata, 10 Podocnemis lewyana, 10 Podocnemis sexituber-
culata, 250 Podocnemis unifilis; L1zARDs—10 Cordylus cordylus ni-
ger, 6 Sceloporus magister, 60 Sceloporus poinsetti, and 40 Cnemi-
dophorus picturatus. Most of these animals were released rather
than destroyed, when they became sick and unsuitable for sale.
A few were released rather than to glut the market for a com-
mercially rare species. During 1946, reptile fanciers released 48
juvenile Pseudemys malonei, from Great Inagua, Bahamas, into
this canal (Hodsdon and Pearson, 1946), but the exact site of
release is not known.

In the vicinity of Pembroke Road and US Interstate Highway
95, Pembroke Park, Broward County, prior to 1964, an animal deal-
er released the following species: rroc—6 Bufo blombergi; Tur-
tLES—oChelus fimbriatus, 20 Pseudemys scripta ornata; and Liz-
ARpsS—4 Basiliscus basiliscus.

At 9550 SW 67 Avenue, Kendall, Dade County, the owners of a
plant nursery released an unknown number of the Panamanian
frog, Atelopus zeteki. This species was last seen at this locality
three years ago.

In the vicinity of 84 Street and 100 Avenue SW, Sunset Park,
Dade County, prior to 1965, a herpetologist released a small num-
ber of West Indian species: FRoc—Eleutherodactylus portoricensis;
LIzARDS—Hemidactylus brooki, Sphaerodactylus macrolepis, Anolis
conspersus conspersus, Anolis cybotes cybotes, and Anolis distichus
ignigularis.

Discussion

If such wholesale introductions continue, they will have a det-
rimental effect on the native fauna of Florida, for when the intro-

152. QuaARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

duced animal competes for a niche already filled by a native spe-
cies, the result can be the elimination of the latter. Indeed there
is evidence that the native species of Bufo and Anolis are being
adversely affected by competition from Bufo marinus and the
many introduced anoles.

It is our sincere hope that such introductions will be stopped
before an exotic amphibian or reptile undergoes a population
explosion, as has occurred with certain mammals ands birds. It
is to prevent just such an occurrence that the State of Florida
prohibits unauthorized introductions (Florida Administrative Code,
1962, Chap. 165-16), and yet none of the above introductions were
made under State permit. If introductions are to occur, we hope
they will be made by responsible biologists only after extensive
studies have determined that the introduced species will not com-
pete severely with the native fauna.

ACKNOWLEDGMENTS

We wish to acknowledge the aid of Neil Chernoff, Richard
Bartlett, Ralph Curtis, Glen and Ted Greenwald, Craig Hutche-
son, Phil Mueller, Dennis Paulson, Albert Schwartz, and C. Rhea
Warren in collecting specimens, or for revealing sites of introduc-
tions. For help with the manuscript, we wish to thank Albert
Schwartz and Fred Thompson. Specimens of the animals listed
in the first two groups have been deposited in the University of
Florida Collections.

LITERATURE CITED

ALLEN, E. R., AND W. T. Nem. 1953. The treefrog, Hyla septentrionalis, in
Florida. Copeia, 1953, no. 2; pp. 127-128.

1955. Establishment of the Texas horned toad, Phrynosoma cornutum,
in Florida. Copeia, 1955, no. 1, pp. 63-64.

Barsour, T. 1910. Eleutherodactylus ricordii in Florida. Proc. Biol. Soc.
Washington, vol. 23, p. 100.

———. 1931. Another introduced frog in North America. Copeia, 19381,
no. 3, v. 140.

———, 198la. A new North American lizard. Copeia, 1931,>nos=3)%pp:
87-89.

——. 1986. Two introduced lizards in Miami, Florida. Copeia, 1936,
NO, POs JUS.

Kinc AND Krakauer: Introduced Herpetofauna 153

Bett, L. N. 1953. Notes on three subspecies of the lizard Anolis sagrei
in southern Florida. Copeia, 1953, no. 1, p. 68.

Carr, A. F., Jr. 1939. A geckonid lizard new to the fauna of the United
States. Copeia, 1939, no. 4, n. 232.

—. 1940. A contribution to the herpetology of Florida. Univ. Florida
ube Biol, oct. Ser., vol. 3, no. 1; pp. 1-118.

Carr, A. F., Jr., AND C. J. Gorn. 1955. Guide to the reptiles, amphibians,
and fresh-water fishes of Florida. Univ. Florida Press, pp. i-ix +
1-341, figs. 1-30, pls. 1-67.

Deckert, R. F. 1921. Amphibian notes from Dade County, Florida. Co-
peia, no. 92, pp. 20-23.

DeESota, C. R. 1934. Phrynosoma from Florida. Copeia, 1934, no. 4,
p. 190.

DuELLMAN, W. E., anp A. ScHwartz. 1958. Amphibians and reptiles of
southern Florida. Bull. Florida State Mus., vol. 3, no. 5, pp. 181-324.

FLormA ADMINISTRATIVE CopE. 1962. The official compilation of rules and
regulations of regulatory state agencies. Secretary of State, Tallahas-
see. Vol. II, chaps. 130-255.

Fow.er, H. W. 1915. Cold-blooded vertebrates from Florida, the West
Indies, Costa Rica, and eastern Brazil. Proc. Acad. Nat. Sci. Phila-
delphia, vol. 67, pp. 244-269.

Gorr, C. C. 1935. An additional note on Phrynosoma cornutum in Florida.
Copeia, 1935, no. 1, p. 45.

Gorn, C. J. 1947. Studies on the life history of Eleutherodactylus ricordi
planirostris (Cope) in Florida. Univ. Florida Studies, Biol. Sci. Ser.,
VOMAw now. pp. ix + 1-67, 7 fies.,, 6 pls.

Hopspon, L. A., AND J. F. W. Pearson. 1948. Notes on the discovery and
biology of two Bahaman fresh-water turtles of the genus Pseudemys.
Proc. Florida Acad. Sci., vol. 6, no. 2, pp. 17-28.

Kinc, W. 1959. Observations on the ecology of a new population of the
Mediterranean gecko, Hemidactylus turcicus, in Florida. Quart. Jour.
Florida Acad. Sci. (1958), vol. 21, no. 4; pp. 317-318.

1960. New populations of West Indian reptiles and amphibians
in southeastern Florida. Quart. Jour. Florida Acad. Sci., vol. 23,
no. 1, pp. 71-78.

Net, W. T. 1957. Historical biogeography of present-day Florida. Bull.
Florida State Mus., vol. 2, no. 7, pp. 176-220.

Ouiver, J. A. 1948. The anoline lizards of Bimini, Bahamas. Amer. Mus.
Novitates, no. 1383, pp. 1-36.

154. QuARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

————,' 1950, Anolis sagrei in Florida. Copeia, 1950) no, Jeeppmo5-sG8

PETERSON, H. W., R. Garrett, AND J. P. Lantz. 1952. The mating period
of the giant tree frog Hyla dominicensis. Herpetologica, vol. 8, pt. 3,
De (OS,

RieMErR, W. J. 1959. Giant toads of Florida. Quart. Jour. Florida Acad.
Sci. (1958), vol. 21, no. 3, pp. 209-211.

Rurpat, R. 1964. An anotated checklist and key to the anoline lizards of
Cuba. Bull. Mus. Comp. Zool., vol. 130, no. 8, pp. 473-520.

SAvAGE, J. M. 1954. Notulae herpetologicae 1-7. Trans. Kansas Acad. Sci.,
vol. 57, no. 3, pp. 326-334.

ScHwarTtz, A. 1952. Hyla septentrionalis Dumeril and Bibron on the Flor-
ida mainland. Copeia, 1952, no. 2, pp. 117-118.

SmirH, H. M., anp R. H. McCaurery. 1948. Another new anole from south
Florida. Proc. Biol. Soc. Washington, vol. 61, pp. 159-166.

STEJNEGER, L. 1922. Two geckos new to the fauna of the United States.
Copeia, no. 108, p. 56.

Trapiwwo, H. 1947. Range extension of Hyla septentrionalis in Florida.
Herpetologica, vol. 3, no. 6, p. 190.

Wricut, A. H., anp A. A. Wricutr. 1949. Handbook of frogs and toads.
Ithaca: Comstock, pp. i-xii + 1-640, 7 figs., 126 pls., 37 maps.

Department of Biology, University of Miami, Coral Gables,
Florida.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

The Vertebral Musculature of Chersydrus (Serpentes)
WALTER AUFFENBERG

ALTHOUGH earlier studies have shown that epaxial myology of
snakes may be extremely useful in determining relationships, much
work remains to be done. Mosauer’s tripartie classification (1935)
of myological patterns (boid, crotalid, colubrid) appears to be
sound, though too simplified to be useful in detailed evolutionary
studies of trunk musculature. Muscle arrangements intermediate
between typical colubrid and boid snakes have been reported
(Auffenberg, 1958), and a more detailed scheme of myological evo-
lution in haenophidian snakes has been proposed (Auffenberg,
1961).

The present study was stimulated by the recent suggestion
that the genera Acrochordus and Chersydrus are not colubrids, but
comprise a distinctive family of haenophidians, probably close to
the Aniliidae (Hoffstetter and Gayard, 1964). Two adult specimens
of Chersydrus granulatus from Bombay, India, were dissected as
the basis for the following myological descriptions. Unless stated
otherwise, all remarks pertain to the muscles of the middle trunk
region.

M. spinalis et semispinalis
(Fig. 1, sp.ssp.)

As in the Boidae, this muscle cannot be separated into spinalis
and semispinalis portions. It arises from a tendinous arch that
stretches between the neural spine and the postzygapophysis of
the same vertebra, with the concave edge of the arch directed
caudally. From this arch the muscle extends as a flat ribbon,
directed forward, medially and dorsally. It is inserted by a long
tendon into the caudodorsal edge of the neural spine of a more
anterior vertebra. In the middle of the body the muscle spans 8
segments. The medial portion of the muscle seems to represent
the spinalis and the lateral portion the semispinalis. Both are
fused throughout their entire lengths and are inserted by means
of a common terminal tendon.

M. interarticularis superior
(Fig. 1, ras)

In Chersydrus this muscle is very similar to that found in all
boids examined, except Sanzinia (Auffenberg, 1958). It arises

156 QuARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

from the lateral portion of the tendinous arch of the spinalis-semi-
spinalis and is directed caudoventrad to be inserted on the medial
tendon of the longissimus immediately before the insertion of
the latter in the posterior lateral border of the postzygapophysis
of the vertebra next craniad. A few fibers are also interlaced
with both more anterior and posterior interarticularis muscles.
Near its origin some of the fibers are interlaced with those of
the multifidus and these are directed under the preceeding tendi-
nous arch towards the insertion of the multifidus. Still deeper
fibers of the interarticularis superior run from one postzygapophy-
sis to the next postzygapophysis anteriorly.

M. multifidus
(Fig. 1, m)

As in most boid snakes, this muscle originates on the ventral
surface of the tendinous arch in Chersydrus. It extends cranially
for two to three segments and is then inserted into the caudal edge
of the lamina of a preceding vertebra. At its lateral surface it is
joined by a few fibers of the interarticularis superior.

M. longissimus dorsi
(Ete leate)

This muscle is similar to that in Constrictor, with a tendinous
origin at the craniolateral portion of a prezygapophysis. At the
level of about the seventh vertebra cranial to its origin the muscle
becomes blended into an aponeurosis forming the lateral fascial
tunnel. The aponeurosis is slightly but regularly thickened into a
series of posterior directed V’s. Each arm of the V-shaped aponeu-
rosis becomes more tendinous and better defined further along its
length. The more ventral tendon serves as part of the origin of the
medially belly of the retractor costae. The dorsal tendon inserts on
a vertebra, where it is joined by tendons of the interarticularis.
The longissimus and its tendon span 10 to 12 vertebrae.

M. retractor costae
(Fig. 1, RCM, RCL)

The two lateral retractor costae muscles are differentiated from
one another only by an indistinct tendinous area. The medial

AUFFENBERG: Musculature of Chersydrus ALI5S7/

SP, SSP

2 :
g : AK
wu ie

mma

- oy

Fig. 1. Vertebral musculature of Chersydrus. Dorsal (A), lateral (B),
and ventral (C) views. CCS, costocutaneus superior; CVD, costovertebrocosta-
lis, dorsal member; CVV, costovertebrocostalis, ventral member; IAI, inter-
articularis inferior; JAS, interarticularis superior; L, longissimus dorsi; LC,
levator costae; M, multifidus; RCL, retractor costae, lateral belly; RCM,
retractor costae, medial belly; SCD, supracostalis dorsalis; SP, spinalis; SSP,
semispinalis; TA, tendinous arch.

158 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

belly (RcM) originates on the aponeurosis of the medial tendon of
the longissimus, as well as by means of a slender tendon that runs
dorsocaudally to the upper portion of a rib. The insertion of the
lateral belly (RcL) is into the dorsal side of a rib, where it is closely
associated with the origins and insertions of the costocutaneus
superior and the supracostalis dorsalis, respectively. These two
muscles are identical to those found in the genus Constrictor.

M. costovertebrocostalis

(Fig. 1, cvp, cvv)

This muscle is represented by two weakly defined heads. The
shorter-fibered, dorsal member (cvp) originates on the costal tu-
bercle and the lateroventral surface of a centrum and is inserted
on the transverse process and the anterior portion of the head
of the succeeding rib.

The longer-fibered, ventral member (cvv) arises on the cranial
border of a diapophysis and the adjacent rib head and inserts by
means of a tendon on the caudolateroventral border of the hypa-
pophysis of a vertebra four segments anteriorly. Some fibers from
the deeper head are interlaced with those of the more superficial
head. Mosauer (1935) considered the fibers of the longer head as
representing a distinct muscle, the transversohypapophysis. How-
ever, in a previous study of vertebral musculature in snakes the
writer (1958) suggested that the transversohypapophysis is a well
developed portion of the costovertebrocostalis found in all snakes
possessing hypapophyses.

M. interarticularis inferior
(Fig. 1, raz)

This muscle arises from the cranial circumference of an ac-
cessory process, passing anteriorly to insert into the caudal cir-
cumference of the accessory process of the next anterior vertebra,
but mostly into the tendons of the levator costae. Mosauer (1935)
believed that the association of this muscle with the levator costae
is typical of colubrid snakes, but the arrangement is also found
in the crotalids (Mosauer, 1935).

AUFFENBERG: Musculature of Chersydrus 159

M. levator costae
(Eiecs ene)

This muscle arises by means of a small tendon from the caudal
circumference of the tip of the accessory process. At this point
the fibers of the interarticularis inferior are inserted into its sur-
face. The fibers of the levator costae arising from this tendon
spread laterally, caudally and ventrally to insert into the cranial
border of the rib of the succeeding caudal vertebral element.

Other Muscles

Most of the remaining vertebral muscles have been shown to
be quite stable when different groups of snakes are compared.
In Chersydrus granulatus the Mm. intercostalis quadrangularis and
tuberculocostalis muscles are present and very similar to those
described for colubrids (Mosauer, 1935) and some boids (Auffen-
berg, 1958). The Mm. intervertebralis, supracostalis dorsalis, su-
pracostalis lateralis superior and inferior, antercostalis proprius,
intercostaloginosus, costalis inferior, transversus abdominus, obli-
quus abdominus internis, costocutaneus superior and inferior are
also very similar to those described for other snakes.

DISCUSSION

The musculature of Chersydrus is unquestionably more like
that of the haenophidians than the caenophidians. This statement
is based on (1) the presence of a well developed tendinous arch,
(2) giving rise to an inseparable spinalis and semispinalis, as well
as (3) the multifidus. Among the haenophidians the arrangement
of the spinalis and interarticularis superior complexes are rather
distinctive for each major group. The simplest arrangement is
found in the genera Python, Boa, and Constrictor, and is only
slightly modified in Xenopeltis. The spinalis and semispinalis are
not divided. The interarticularis superior is composed of fibers
running from the anterior edge of one postzygapophysis to the pos-
terior edge of the postzygaphysis of a more anterior vertebra. In
Sanzinia and Calabaria the interarticularis superior has similar
attachments, but it also has a mass of fibers running to the semi-
spinalis portion of the tendinous arch. This last arrangement has
been considered as representing a primitive diagastricus (Auffen-

160 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

berg, 1958, 1961), and is the pattern found in Chersydrus. In the
remaining haenophidians the spinalis-semispinalis complex is sep-
arated into two distinct muscles, one of which has migrated off of
the tendinous arch (Auffenberg, 1961).

In all haenophidians in which the vertebral musculature is
known, except Python and Anilius, the longissimus runs from an
accessory process of one vertebra to the neural spine of a more

CONSTRICTOR & XENOPELTIS PYTHON

ERYX COLUBER

Fig. 2. Generalized vertebral musculature in haenophidian snakes, com-
pared with that of a caenophidian, Coluber. IAS, interarticularis superior;
LO, longissimus dorsi; NS, neural spine; POZ, postzygapophysis; PRZ, prezy-
gapophysis; RCM, retractor costae, medial belly; SP, spinalis; SSP, semispin-
alis. The longissimus inserts on a neural spine in genera in the left column,
on a postzygapophysis in genera in the right column.

AUFFENBERG: Musculature of Chersydrus 161

anterior vertebra. In these two genera, as well as in Chersydrus,
the muscle runs between the process and a postzygapophysis.

In Sanzinia and Calabaria the multifidus originates on the neu-
ral arch, not on the tendinous arch, where it originates in all other
haenophidians and in Chersydrus.

The remaining vertebral muscles of Chersydrus are so similar
to those of other snakes that a comparison is unnecessary. <A dia-
gramatic comparison of the most important vertebral muscles of
Chersydrus and other known haenophidians is provided in figure 2.

CONCLUSIONS

1. Chersydrus (and probably the closely related Acrochordus)
is properly placed with the haenophidians on the basis of its verte-
bral musculature, principally because of the presence of a well
developed tendinous arch.

2. Certain peculiarities of its vertebral musculature separate
it from all other haenophidian muscle arrangements described so
far. These differences are equivalent to those characterizing the
familial level within the haenophidians. Conclusions 1 and 2 sup-
port those of Hoffstetter and Gayard (1964) and Underwood (in
litt.) based on other morphological characters.

3. The vertebral musculature of Chersydrus (and probably
Acrochordus) exhibits certain similarities to the Boyleriinae (ar-
rangement of the spinalis-semispinalis and interarticularis complex-
es) on the one hand, and to the Aniliidae on the other (insertion of
longissimus on postzygapophysis). On the basis of its vertebral
musculature Chersydrus seems closer to the Boidae than to the
Aniliidae, and more distantly related to the Xenopeltidae.

LITERATURE CITED

AUFFENBERG, W. 1958. The trunk musculature of Sanzina and its bearing

on certain aspects of the myological evolution of snakes. Breviora,
MO. WA Foye). ISI

—. 1961. Additional remarks on the evolution of trunk musculature
in snakes. Amer. Midl. Naturalist, 65(1): 1-16.

HorFstTETTer, R., AND Y. Gayarp. 1964. Observations sur l’osteologie et la
classification des Acrochordidae (Serpentes). Bull. Mus. Nat. d’Hist.
Nat., 36(5): 677-696.

162 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MosavuErR, W. 1935. The myology of the trunk region of snakes and its sig-
nificance for ophidian taxonomy and phylogeny. Univ. Calif. Los
Angeles, Publ. Biol. Sci., vol. 1, pp. 81-120.

Florida State Museum, Gainesville, Florida.

Quart. Jour. Florida Acad. Sci. 29(2) 1966

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1966

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Presbyterian College
Florida Southern College
Florida State University
Jacksonville University
Marymount College
Miami-Dade Junior College
Mound Park Hospital Foundation
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida

University of Tampa

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1966

President: MARGARET GILBERT
Department of Biology, Florida Southern College
Lakeland, Florida

President Elect: Jackson P. SICKELS
Department of Chemistry, University of Miami
Coral Gables, Florida

Secretary: JouNn D. KiLsy
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: JAMES B. FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: Prerce BropkKoRB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges

should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

>
C?

1

Fé | ER
: ‘ Quarterly Journal

of the
Florida Academy of Sciences

Vol. 29 September, 1966 No. 3

CONTENTS

Variscite from the Hawthorn Formation
Frank N. Blanchard and Stephen A. Denahan 163

An investigation of ostracode preservation Mervin Kontrovitz 171
Florida Academy of Sciences award for 1967 178

Revision of the selenodont artiodactyls from Thomas Farm
Thomas H. Patton 179

Distribution of Euglenida in North Florida John J. McCoy 191
Two new subfamilies of monogenetic trematodes C. E. Price 199
Mayfly nymphs from northwestern Florida Robert F. Schneider 202

Caribbean recruitment of Florida’s spiny lobster population
Harold W. Sims, Jr., and Robert M. Ingle 207

Mailed October 20, 1967

QUARTERLY JOURNAL OF THE FLORA ACADEMY OF SCIENCES
Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Makrcins should be 1% inches all around.

Tirtes should not exceed 42 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and
AppREss in paragraph form following Literature Cited.

LrreraTurE Crrep follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLES are charged to authors at $16.70 per page or fraction. Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

Lecenps for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($15.80 per page, $14.00 per half
page for zinc etchings; $14.50 per page, $13.25 per half page for halftones).
Drawincs should be in India ink, on good board or drafting paper, and let-
tered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. PHoTocrapus should be of good contrast, on glossy paper. Do not
write heavily on the backs of photographs.

ProoF must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

Vol. 29 September, 1966 No. 3

Variscite from the Hawthorn Formation
FRANK N. BLANCHARD AND STEPHEN A. DENAHAN

THE purpose of this article is to report the occurrence of micro-
crystalline variscite, (Al, Fe) PO,:2H,O, as an abundant constitu-
ent of weathered phosphatic sandstones in an exposure of the
Hawthorn Formation north of Ocala, Marion County, Florida.
Weathered portions of the Bone Valley Formation in central Florida
and weathered phosphatic beds of the Hawthorn Formation (farther
north) contain secondary phosphate minerals (the aluminum phos-
phate zone), derived, in part, from original grains and pellets of
carbonate-fluorapatite. Investigations of the mineralogy of the
aluminum phosphate zone in the Bone Valley Formation have dis-
closed the presence of secondary phosphate minerals, including
metastrengite (Hill et al., 1950), crandallite (Altschuler et al.,
1956), wavellite (Bergendahl, 1955), millisite (Owens et al., 1960),
and vivianite. Although variscite is a mineral that might be ex-
pected, as far as we know it has not been reported previously.

OcCURRENCE

Samples containing variscite were collected at a road cut in the
Hawthorn Formation on the east side of new U. S. 441, 8.1 miles
north of the intersection (in Ocala) of U. S. 441 and State Road
40. The cut exposed a maximum thickness of about 35 feet of clays
and phosphatic sands and sandstones of the Hawthorn Formation.
Within this section, variscite is confined to the uppermost sand-
stone beds, from the soil down to a vertical depth of about 6 to 9
feet. The variscite-containing rocks are poorly to moderately well
indurated, gray to yellowish brown, porous, fine-grained sandstones
with an argillaceous appearance. Five samples from the uppermost
sandstone beds are dealt with in this report and are numbered

164 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

(2.5, 5, 6, 9, 11.5) to indicate the approximate vertical depth (in
feet) from which the sample was taken, measured from the ground
surface. In addition to the location just north of Ocala, we have
found that variscite occurs in very small amounts intimately mixed
with the wavellite at several locations farther north in Alachua
County.

IDENTIFICATION

Variscite was first noted as an unidentified peak on a differential
thermal analysis and later as several unidentified lines on an X-ray
diffraction pattern. Further collection of specimens from the same
location provided material containing large enough concentrations
of variscite so that it could be identified by X-ray, optical, and
thermal techniques. Nearly all the quartz sand and silt was easily
separated from the other minerals in the variscite-bearing rocks by
gentle crushing and sieving (200 mesh); however, attempts to
separate variscite (by specific gravity) from other minerals in the
minus-200 mesh fraction failed. Therefore, X-ray diffraction patterns
and DTA curves consist of the superimposed patterns of several
minerals.

X-ray diffractometer patterns were made on a General Electric
XRD-5 unit with a copper tube operated at 35 kv. and 15 ma. Using
highest amplifier gain, a time constant of 4 seconds, and a pulse
height analyzer to reduce background, each sample was scanned
from 32° to 6° 26 at a rate of 2° 26 per minute. The resulting pat-
terns (Fig. 1.) show various mixtures of variscite, wavellite, apatite,
kaolinite, and quartz. The d-spacings for the variscite, computed
from the 24 angles through the range scanned, fall between the
values recorded for variscite and ferrian variscite on A.S.T.M.
cards no. 15-281 and no. 7-69, indicating the presence of a small
amount of iron that has substituted for aluminum (Table 1).

The presence of iron is also indicated by optical properties. Be-
cause of small crystal size, no interference figures were obtainable,
however (statistical) minimum and maximum refractive indices
were measured—X = 1.566 and Z=1.602. Palache et al. (1951) give
X—1.563, Y—1.588, Z=1.594 for the pure variscite end member
(AIPO,:2H,.O) of the variscite-strengite series, and X=1.707,
Y=1.719, Z=1.741 for the pure strengite (FePO,:2H.O) end
member. The difference in refractive indices between pure variscite
and Florida variscite is most logically accounted for by assuming

ee a ee a a a |
ied
= > ae PLoS o> > >
na #~@ =.0—mUC—<‘<‘<;2xH Liles ei Ga Gs
Poets ea as RNS Ug
n
Cy |]
oy Wiehe
oF. 3° 3
> >
zB x >> ££ £ B> z -
-_—_—
=
wo WW
uJ
a
=——
a
x
z z Pa
a
uJ
(an)
ve)
aD
a
=
<
n
°
o
Ls)
=> zz s =

10 20 30
DEGREES 26——-

Fig. 1. X-ray diffractometer patterns (tracings) for samples 2.5, 5, 6, 9,
11.5. V, Variscite; W, Wavellite; A, Apatite; K, Kaolinite; and Q, Quartz.
For sample 2.5 approximate 29 angles are given for each variscite line.

166 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

the presence of a small amount of substitutional iron, an interpreta-
tion which is consistent with the X-ray data.

TABLE 1

Comparison of 24 Angles and d-spacings for Variscites

Variscite Variscite Fe-Variscite
(ADS; DM. No. 15-281 ) (Florida ) (A. S. T. M. No. 7-69)
26 d 26 d 20 d
16.51 5.365 16.46 5.38 16.35 5.42
18.42 4.82 18.37 4.83 18.26 4.86
20.84 4.257 20.76 4,28 20.69 4,29
22.86 3.887 22.64 3.92 22.54 3.94
24.56 3.625 24,32 3.66 24.31 3.66
26.48 3.366 26.66 3.34 26.52 3.36
27.81 3.204 OAL TC! SPA OAT) 3.24
29.39 3.039 29.14 3.07 29.08 3.07

Differential thermal analyses of each of the minus-200 mesh
fractions of the samples were carried to 800° C at a rate of 10° per
minute using a sensitivity of 25 per cent on a Deltatherm unit. On
these curves (Fig. 2) endothermic reactions at 175° C are attrib-
uted to variscite, endothermic reactions at 250° C and, in some
samples, at 275° (weak) are caused by wavellite, and endothermic
reactions (weak due to small percentage at 575° to 600° C are those
of kaolinite. Endothermic reactions below 150° C may represent
escape of hygroscopic water or reactions involving clays. For refer-
ence, in Fig. 2, DTA curves of wavellite (Montgomery County,
Arkansas ) and variscite (Fairfield, Utah) are included. Apparently
both variscite and wavellite produce pronounced and diagnostic
DTA curves.

PETROGRAPHIC CHARACTERISTICS

The variscite occurring in at least some of the rocks studied
may be readily seen and distinguished from other minerals in thin
section. The birefringence is moderately high, so that even with
very thin crystals second order interference colors can be seen.
Some of the variscite is found in silt- to sand-sized aggregates of
small equant or slightly elongate particles, somewhat like chert but
with moderate positive relief. Under very high magnification it can

BLANCHARD AND DENAHAN: Hawthorn Variscite 167
be seen that the particles are mostly aggregates of several elongate

radiating crystals. In other places variscite occurs as secondary
cement formed by both replacement and precipitation.

200 400 600

2.5

6
SAMPLE DEPTH — FEET

mina Tai fie a aT EP Oe ea

200 400 600

TEMPERATURE — °C.

Fig. 2. Differential thermal analysis curves (tracings) for samples 2.5,
5, 6, 9, 11.5. V, variscite; W, wavellite; and K, kaolinite. Variscite (Fairfield,
Utah) and Wavellite (Montgomery County, Arkansas) included for compari-
son. Endothermic reactions are downward.

168 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The relationships between variscite and other minerals in the
rocks show clearly that variscite is a late phase formed by replace-
ment of pre-existing minerals and by filling of cracks and cavities.
Figs. 3 and 4 are photomicrographs typical of the appearance of
sample no. 2.5. In Fig. 3 the rounded, relatively large masses of
variscite are thought to be former apatite pellets replaced (at least
ultimately ) by variscite; matrix variscite is also present. In Fig. 4
the matrix of the sandstone consists almost entirely of variscite; in
places pores can be seen with chalcedonic-like linings of variscite.
There are many indications that clay and possibly wavellite have
been replaced by variscite, although no single clear-cut textural
feature can be cited as proof.

1 Bs

Fig. 3. Photomicrograph of sample 2.5 showing quartz grains, pellets of
aggregated variscite, fine-grained matrix containing variscite. Crossed polars.
Scale equals 1 mm.

ORIGIN

Using the X-ray patterns and DTA curves as a rough guide to
the relative proportions (in the minus-200 mesh fractions) of the
minerals variscite, wavellite, and apatite, at different levels in the
section from the ground surface to minus 11.5 feet, and from obser-

BLANCHARD AND DENAHAN: Hawthorn Variscite 169

Fig. 4. Photomicrograph of sample 2.5 showing quartz grains, pore spaces,
and matrix of almost pure variscite. Crossed polars. Scale equals 1 mm.

vations of textures seen in thin sections, we gain some insight into
the mode of origin of these minerals. From the X-ray patterns,
apatite (line at 32.0° 26) is absent (or nearly so) in sample no. 2.5,
and increases in percentage downward, reaching a maximum in the
lowest sample (no. 11.5). From the X-ray and DTA patterns, vari-
scite (X-ray line at 16.5° 20) is greatest in the upper three samples
and does not appear in the lowest sample. Wavellite (X-ray line at
10.4° 26) is practically absent at the top of the section, is abundant
in the middle samples and decreases downward. These variations
(which are confirmed by observations of the thin sections ) are con-
sistent with a weathering origin for variscite, where variscite and
wavellite are formed at the expense of apatite (and probably clay
minerals ), and where some variscite may be formed at the expense
of wavellite. Possibly variscite has formed in the nearer-surface
more acidic environment while wavellite is the stable phase in the
slightly deeper acidic environment. The development of wavellite
and other minerals in the aluminum phosphate zone of the Bone
Valley Formations has been described by Altschuler et al. (1956)

170 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

as the result of lateritic weathering of porous sediments, containing
carbonate-fluorapatite, by downward moving acidic solutions. Vari-
scite originates in a manner similar to that of wavellite, except per-
haps in a more acidic environment.

ACKNOWLEDGMENTS

This research was supported in part by NSF Grant No. GY-330
and in part by the Graduate School of the University of Florida
through a summer research appointment (1966) granted to the
senior author. We are grateful to Dr. R. W. Gould and the Depart-
ment of Metallurgy, University of Florida for use of the X-ray
diffraction equipment and to the Department of Soils for the differ-
ential thermal analyses.

LITERATURE CITED

ALTSCHULER, Z. S., E. B. JAFFEE AND FRANK CurttitTA. 1956. The aluminum
phosphate zone of the Bone Valley Formation, Florida and its uranium
deposits. U.S. Geol. Survey Prof. Paper 300, pp. 495-504.

BERGENDAHL, M. H. 1955. Wavellite spherulites in the Bone Valley Formation
of entral Florida. Am. Mineral., vol. 40, pp. 497-504.

Hitt, W. L., W. H. Armicer, AND S. D. Goocu. 1950. Some properties of
pseudowavellite from Florida. Mining Eng., vol. 187, pp. 699-702.

Ownes, J. B., Z. S. ALTSCHULER, AND R. BERMAN. 1960. Millisite in phosphorite
from Florida. Am. Mineral., vol. 45, pp. 547-561.

PALACHE, C., H. BERMAN, AND C. FRONDEL. 1951. Dana’s system of mineral-
ogy., vol. 2, John Wiley and Sons, New York, New York, 1124 pp.

Department of Geology, University of Florida, Gainesville,
Florida.

Quart. Jour. Florida Acad. Sci. 29(3) 1966 (1967)

An Investigation of Ostracode Preservation
MERVIN KONTROVITZ

Many authors have considered the problem related to fossil
preservation as it affects interpretation in paleoecology. If a fossil as-
semblage is used to reconstruct an ecological situation, knowledge
of the composition of the original population is necessary. Ager
(1963) emphasizes that preservation is selective for various groups
and within one group.

Ostracodes are abundant in many geologic formations and as
fossils they have been used as indicators of habitat (Benson and
Coleman, 1961). A factor that must be considered in the paleo-
ecology of ostracodes (and other groups) is precisely what kinds of
specimens are preserved in relation to the proportions in the once
living assemblage.

To initiate this study, four ostracode species were selected from
Recent bottom samples collected in Florida Bay, near Vaca Key.
Bairdia victrix Brady, Cushmanidea elongata Brady, Loxocorni-
culum postdorsalatum (Puri), and Mutilus confragosa (Edwards )
were chosen because they are quite different in size, valve thick-
ness, and ornamentation. Bairdia victrix Brady is a large smooth
species in contrast to Cushmanidea elongata (Brady) which is
small and thin-shelled. Moderate ornamentation is found on Loxo-
corniculum postdorsalatum (Puri), while Mutilus congragosa (Ed-
wards ) is heavily ornamented.

PROCEDURES AND RESULTS

Using the species just mentioned, 40 adult valves of each were
selected. Each valve was examined under magnification to insure
that there were no incipient fractures.

The base of the test device used consisted of a wooden box filled
with concrete into which four test tube holes were formed. While
the concrete was wet the plastic test tubes were coated with silicon
lubricant and thrust into the concrete, and left until it had hardened.
The concrete thus supported the plastic test tubes, but the tubes
could be removed at will. See Figure 1.

A small portion of the sediment from the sample was placed
in a plastic test tube which was to be encased in concrete (Fig. 1).
Ostracodes of each test species were placed on the wet matrix and

172 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

covered with more sediment from the sample. The entire column of
sediment and ostracodes was then wetted, either with fresh or sea
water. The water was allowed to soak into the sample and at no
time during testing was the material dried out.

weight

cam

Figs. 1-2. Fig. 1 (upper). Slow crushing apparatus. Fig. 2 (lower). Sedi-
ment agitator.

A wooden dowel was tipped with the rubber portion of a medi-
cine dropper. This plunger assembly was then inserted into the
tube with ostracodes and matrix. A silicon lubricant was spread on

Kontrovitz: Ostracode Preservation 173

the rubber plunger head to prevent water evaporation. During the
insertion, however, excess water was forced out past the seal, but
then the remaining water could not evaporate.

The tops of the wooden dowels (plungers) were covered with
a board so that the load would be evenly distributed. A plastic con-
tained was placed on the board as shown in Fig. 1. Slowly, over a
period of a week, water was added to the container until the entire
load equalled 20 pounds. Using the cross-sectional area of the
plungers it was found that the pressure on the top of the matrix
was about 15 pounds per square inch.

Two trials were made, each with two tubes wetted with fresh
water, and two wetted with sea water. Each tube contained five
valves of each test species; therefore, each trial tested 20 valves
of each species.

After the load was removed, each test tube was carefully washed
and all matrix material was recovered in a 200 mesh screen. Under
magnification all particles were examined and ostracode valves and
fragments were recovered.

The results of the slow crush tests show that there is a difference
in resistance of ostracode valves to loading. Each of the four
species displays a different percentage of recovery.

Based on 40 valves of each species, it was found that the great-
est recovery was for Bairdia victrix, with 26 valves and two trag-
ments. Mutilus confragosa was second in abundance after the crush
tests. It yielded 18 complete valves and one recognizable fragment.
Loxocorniculum postdorsalatum showed more effect of the loading
giving only six valves and two fragments. Only one valve of the
original 40 was recovered of Cushmanidea elongata.

Another mechanical test was performed using the four test
species. Again, 40 valves of each were selected and these were also
examined to insure that they were not cracked or broken.

The 40 ostracode valves were placed in a small glass vial with a
thin layer of wetted sediment from the sample. The vial was closed
and placed in a metal cylinder so arranged that the vial rested on
an eccentric wheel. The eccentric was driven by a small electric
motor which rotated at 100 revolutions per minute (Fig. 2).

The maximum lift of the eccentric was 0.8 inches. Eccentric
configuration is shown in Fig. 2. Several shapes were tried until
one was selected that provided moderate agitation and smooth
operation.

174. QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The vial was allowed to remain in the cylinder with the motor
operating for a week. When examined the ostracodes showed only
a polish; no further damage could be seen.

After two weeks of total operation the vial was again removed
and the contents examined. The smooth Bairdia victrix displayed
only a high polish. Cushmanidea elongata displayed a high polish
with some fractures visible. Ornamentation was slightly worn on
both Loxocorniculum postdorsalatum and Mutilus confragosa.

Finally, after three weeks of agitation, the process was stopped
and the ostracodes were removed from the sediment. It should be
noted that all examinations were carried out using a specially con-
structed screen made of standard 200 mesh. Each time, the entire
contents of the vial were examined under a microscope to insure
complete recovery of all fragments.

At the end of the three week test the most abundant ostracode
found was Bairdia victrix. Of the 40 originally put into the vial, 36
of this species were found complete and one had been broken.
Mutilus confragosa was second in numbers recovered, with 24
whole valves and two fragments. Loxocorniculum postdorsalatum
remained as 20 valves and five fragments. The least number of
valves seen were those of Cushmanidea elongata; 14 valves and two
fragments were recovered.

In both tests the species showed the same order of resistance.
The tests were not intended to reproduce geologic time in any way,
but simply to demonstrate the ability of the various ostracode
valves to withstand crushing and abrasion.

An attempt was made to test, directly, the crush strength of
single valves by applying weight to a single valve.

A rod was attached to a light plastic container. This assembly
was free to move through a rigidly mounted vertical cylinder. Two
such assemblies were fabricated, one weighed 11.25 grams and the
other 30.04 grams, dry weight. Water was slowly added to the
cylinder thereby increasing the weight ( Fig. 3).

The crushing tip of the apparatus was gently placed on the
outside surface of an ostracode valve. Water was allowed to flow
from a burette into the plastic contained. In this way it was hoped
that the weight needed to crush a valve could be measured.

All valves were wetted with tap water before crushing. This
is important to note because the strength of the valves is different
if dry. Once wetted the valves are less strong and this was selected

Kontrovitz: Ostracode Preservation 175

burette

—— waler container

wooden dowel
—— cylinder

metal tip

7 ostracade

Fig. 3. Rapid crushing apparatus.

176 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

as the standard. Certainly in bottom sediment the valves are wet,
and it is a likely condition with ground water also.

It was hoped that weights determined for crushing could be
offered only as a means of comparing the four species. The values
were not to be construed as absolute crushing strengths.

In practice it was found that Cushmanidea elongata and Loxo-
corniculum postdorsalatum were crushed with the lighter of the two
units even if no water was added. The lighter unit weighed 11.25
grams as mentioned above.

Bairdia victrix and Mutilus confragosa were not crushed by the
lighter unit filled with water; the weight was 26.88 grams.

When the heavier unit was applied all Mutilus confragosa were
crushed without the addition of water. Bairdia victrix all required
some water to be added to cause crushing.

It is obvious that new crushing units wili have to be con-
structed which span the entire range of ostracode valves. Many
more trials will be required before the crushers can be made of
the proper weights. It will be interesting to see if fast crushing
can be correlated to the abrasion and slow crushing tests.

The abrasion and slow crush tests do indicate that ostracode
valves probably are not preserved in proportion to the species in
the living assemblage. If this premise is indeed probable, it is in
order to use a sample as an example of what could result. Ecological
data are derived from Benson and Coleman (1963), Puri and Hul-
ings (1957), Hulings and Puri (1964), and Van Morkhoven (1963).

In the sample used in this study, Cushmanidea elongata equaled
41.8 per cent of the recovered ostracodes. If this species is not pre-
served in direct proportion to the living population, then the fossil
assemblage would reflect neither the true population nor the
ecology. This error would exist even if there were no disturbance
by factors such as currents.

In the sample, Bairdia victrix is 6.7 per cent of the total popula-
tion. If this ostracode is preserved in greater proportions than Cush-
manidea elongata an erroneous paleoecologic situation would be
reconstructed. Instead of a very shallow water, low salinity (less
than 30 parts per thousand) environment, the investigation might
conclude that the water was greater than 65 feet deep with a
salinity of 35 to 40 parts per thousand.

It is believed that much work should be done to approximate
the probable ratios of preservation of ostracodes and other micro-

Kontrovitz: Ostracode Preservation Lee

fossils. In this manner the paleoecology can be more validly used.
If no relationship can be established between living and fossil
population, the usefulness of the preserved populations in paleo-
ecology is doubtful.

SUMMARY

Even if the ecology of modern groups were perfectly known,
the paleoecologists would be confronted with the question of
differential preservation. This is not a new idea, but a means is
needed to clearly demonstrate what groups are preserved or
destroyed.

Testing by mechanical means indicates that various ostracode
taxa are probably not preserved in proportion to the representation
in the living fauna. Extension of such testing will be useful to pre-
dict what proportions are preserved, and in reconstructing the
fossil populations of ostracodes and other groups.

ACKNOWLEDGMENTS

Dr. Richard A. Edwards and Dr. Caspar Rappenecker of the
Geology Department of the University of Florida made the com-
pletion of this study possible.

LITERATURE CITED
AcER, D. V. 1963. Principles of Paleoecology. New York, McGraw-Hill Book
Commnes Silepp., 8 pls., 147 figs.

BENSON, R. H. and G. L. CoLEMAN. 1963. Recent Marine ostracodes from
the eastern Gulf of Mexico. Kansas Univ., Paleont. Contr. 31, Arthro-
poda, art. 2, 52 pp., 8 pls., 33 figs.

Huuincs, N. C., anp H. S. Purr. 1964. The ecology of shallow water ostra-
cods of the West Coast of Florida. Pubbl. staz. zool., Suppl., Vol. 33,
pp. 308-344, 17 figs.

Purr, H. S., anp N. C. Hutines. 1957. Recent ostracode facies from Panama
City to Florida Bay area. Trans. Gulf Coast Assoc. Geol. Soc., vol. 7,
pp. 167-190, 11 figs.

VaN MorkHOovEN, F. P. C. M. 1963, Post-Paleozoic Ostracoda. New York,
American Elsevier Publ. Co., vol. 2, 478 pp., 763 figs.

Department of Geology, University of Florida, Gainesville,
Florida (present address: Department of Geology, Tulane Uni-
versity, New Orleans, Louisiana).

Quart. Jour. Florida Acad. Sci. 29(3) 1966 (1967)

Florida Academy of Sciences Award for 1967

THE recipient of the Florida Academy of Sciences honors award
for 1967 was Dr. Alfred H. Lawton. The medal and citation were
presented by Dr. C. C. Clark at the 3lst annual meeting of the
Academy in Tampa, March 9-11.

Dr. Lawton holds the M.D. and Ph.D. degrees from North-
western University and received an honorary Sc.D. degree from
Simpson College. During World War II he was an officer with the
U. S. Public Health Service. Afterwards he served as chief of the
Research Division for Medicine and Surgery of the Veterans Ad-
ministration in Washington, director of medical research for the
United States Air Force, and more recently associate chief of staft
with the Veterans Administration Center at Bay Pines, Florida, and
director of research on aging for the U. S. Public Health Service at
St. Petersburg, Florida. He joined the staff of the University of
South Florida in 1965, where he is assistant dean of academic affairs.

His publications include two books and some 90 papers dealing
with chronic diseases and aging. His research includes experi-
mental studies on the toxicity of trinitrotoluene, antagonism of the
vasopressor action of nicotine by potassium compounds, aortic
stenosis and sudden death, lactic acid metabolism, and biochemical
and physiological measurements of aged populations.

Dr. Lawton addressed the Academy on the subject “A Lifelong
Plunge into the Universe of Ideas.”

Quart. Jour. Florida Acad. Sci. 29(3) 1966(1967)

Revision of the Selenodont Artiodactyls from Thomas Farm
THomas H. Patron

THE unusual and fascinating nature of the Miocene artiodacty]
fauna from the Thomas Farm quarry in north central Florida was
first brought to light through a series of articles by T. E. White
(1940, 1941, 1942, 1947). Although Simpson (1932) had earlier
described Oxydactylus floridanus (=Nothokemas floridanus) and
an indeterminate cervid from the Thomas Farm, it was only after
the extensive Harvard University excavations from 1939 to 1947
that the extent of the fauna was realized.

Several Miocene and Pliocene artiodactyls occur both in Florida
on the Texas Gulf Coastal Plain. Comparison of material from
the two regions led to a reévaluation of several artiodactyls from
Texas and a suprageneric reallocation of some of the Thomas Farm
species (Patton, 1967 and Ms). The author considers the Thomas
Farm Fauna to be of Early Hemingfordian (early Middle Miocene )
age.

The Miocene vertebrate faunas from the Gulf Coastal Plain of
Texas and Florida are becoming sufficiently well known that a
revision of at least a portion of the artiodactyl fauna would be
timely and helpful to those workers not familiar with this fossil
province. Earlier listings of the Thomas Farm artiodactyls are pro-
vided in Romer (1947), Ray (1957), Olsen (1962), and Patton
(1964). Apparently C. Ray and B. Patterson, in an unpublished list
of these forms, visualized some of the changes made below.

I should like to thank Bryan Patterson of the Museum of
Comparative Zoology, Stanley Olsen of the Florida State Geologi-
cal Survey, and Beryl Taylor of the Frick laboratory, American
Museum of Natural History, for the generous loan of most of the
fossil material discussed herein.

Order ARTIODACTYLA Owen, 1848
Suborder TyLopopa Illiger, 1811
Family Camelidae Gray, 1821
Subfamily Camelinae Gray, 1821

Nothokemadinae White, 1947

180 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Genus Nothokemas White, 1947
Nothokemas floridanus (Simpson), 1932

Oxydactylus floridanus Simpson, 1932, p. 35, figs. 20-21.
Paratylopus grandis White, 1940, p. 33, pl. 5, figs. A, B.
Nothokemas grandis (White ); White, 1947, p. 508, figs. 5-6, in part.

Type. FGS V-5247, part of a right maxilla with P?-M?. Para-
type, FGS V-5238, a right ramus with P;-Ms3.

The genus Nothokemas was originally described by White
(1947) on the basis of characters exhibited in a crushed skull, a
right maxilla with P’-M*, and several mandibles with partially pre-
served dentitions, all recovered from the Thomas Farm quarry, but
not definitely associated. The type species of Nothokemas was des-
ignated by White (1947) as Nothokemas grandis, first described by
him (1940) as Paratylopus grandis, also from the Thomas Farm.

On the basis of comparisons of specimens from the Thomas
Farm and from the Garvin Gully Fauna of Texas, it is the writer's
opinion that Nothokemas grandis White (1947) and Oxydactylus
floridanus Simpson (1932) are synonyms. A comparison of the lower
jaws of the two genera follows (all after Patton, Ms):

In the unreduced condition of the premolars and the configuration of
both molars and premolars, the lower jaw of Nothokemas is remarkably
similar to Oxydactylus. As in Oxydactylus, the body of the mandible is deep
and thin, but the diastema anterior to P., is considerably longer. The mandible
in this region is much more attenuated than in Oxydactylus. Both genera are
brachyodont and both display a relative elongation of the molars. In a specimen
(MCZ 4323) referred by White (1947) to N. grandis, just anterior to the
mental foramen there is what appears to be an alveolus for a large canine-like
tooth. In a smaller and slightly dissimilar specimen (UTBEG 40067-10),
assigned to Nothokemas minimus n. sp., a large caniniform tooth is present.
Because of its position well anterior to the anterior mental foramen and the
posterior border of the symphysis, I believe this tooth to be the true canine.
Loss of P,, therefore, should be included in the generic diagnosis of
Nothokemas.

The lower molars of Nothokemas differ from Oxydactylus in the presence
of an intercolumnar tubercle between the protoconid and hypoconid. The
posterior edge of the entoconid of M, overlaps the hypoconulid, giving the
appearance of the beginning of an “outer lobe” on the talonid (White, 1947,
Dar ole)

The anterolingual fold of P, and P, is distinctly more pronounced in
Nothokemas than in Oxydactylus. The posterolingual cusp (entoconid?)
of P, and P, is rather strongly expressed in Nothokemas; In Oxydactylus it
is distinct on three of the four specimens with P, intact, but it has virtually
disappeared from P,, the posterolabial cusp (hypoconid?) being the only
posterior cusp remaining.

Patton: Miocene Selenodonts 181

Because the genus Oxydactylus is in need of revision and pres-
ently lacks taxonomic unity, this and all subsequent comparisons
in this paper will be made with the following species only: O.
longipes Peterson (type species), O. brachyodontus Peterson, O.
campestris Cook, and O. benedentatus (Hay ).

The molars of the upper jaw of Nothokemas differ considerably
from those of Oxydactylus. Strongly developed stylar cusps are
present on the paraselene and metaselene of Nothokemas, and a
prominent rib extends up each selene from the base of the crown
to the tip of both paracone and metacone. These features are not
as well expressed in Oxydactylus. In addition, intercolumnar
tubercles, or accessory tubercles, between the protoselene and
hyposelene, completely absent in Oxydactylus, are a consistent
character of the molars of Nothokemas. Prominent cingula are
present among the anterior margin of the anterior crescents of
M'-M*. On M! of the type( FSGS V-5247 ) of Nothokemas floridanus,
this cingulum extends posteriorly around the anterior crescent, and
although part of the tooth is broken off in this region, it probably
connected with the intercolumnar tubercle.

The upper premolars of Nothokemas are unusually large and
robust. P? and P* are both proportionally longer and wider than
those teeth in Oxydactylus. P? of Nothokemas aiso differs from that
of Oxydactylus in its relatively greater size and in the greater de-
velopment of the anterolabial flexus and the internal cingulum.

In summary, the long diastema anterior to P., loss of P,, the
posterior extension of the entoconid on Ms, the presence of inter-
columnar tubercles on the upper and lower molars, the greater de-
velopment of the posterolingual cusp on P,; and P;, and the more
pronounced anterolabial flexus and internal cingulum on P?, all
serve to separate Nothokemas from Oxydactylus. The systematic
affinities of Nothokemas are discussed below in the section on
Floridatragulus.

A partial left maxilla with dP?-*-M! (MCZ 4328), referred by
White (1947) to Nothokemas grandis, is placed here in Prosynthe-
toceras texanus. I have not examined two additional specimens.
These are a partial skull (MCZ 4329) and a partial right maxilla
(MCZ 4322).

182 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Subfamily Floridatragulinae new subfamily
Genus Floridatragulus White, 1940
Floridatragulus dolichanthereus White, 1940

Floridatragulus dolichanthereus White, 1940, p. 35.
Hypermekops olseni White, 1942, pp. 11-13, pl. 9, figs. 1-3.

Type. MCZ 3635, a partial mandible with right and left M,-M3.

White described the genus Floridatragulus from material
recovered from the Thomas Farm deposits and designated Florida-
tragulus dolichanthereus as the type species. His generic diagnosis
was given as follows:

A large brachyodont hypertragulid with a very long mandibular symphysis,
diastema between P, and P, equal to two thirds of that between P, and P.,,
basal pillars of molars low and elongate antero-posteriorly, ‘heel’ of M, divided
so that it forms two grinding crescents.

In 1942, White described Hypermekops on the basis of a skull
containing [’-?, P? and P*, and M?*° of the right side, and I? and P*
to M® on the left side. Hypermekops olseni was designated the type
species. White’s generic diagnosis of Hypermekops was given as
follows:

A large brachyodont hypertragulid with three incisors in the premaxillary,
fourth premolar and molars similar in form to those of Leptomeryx, P? three
rooted and probably with a median spur, P? double rooted, elongate antero-
posteriorly and without median spur, I’ to P! caniniform and slightly recurved,
I largest.

Although White (1942, P. 13) was aware that Hypermekops
and Floridatragulus were closely related (he inferred on the basis
of snout length that Floridatragulus was derived from Hyper-
mekops ), the lack of a comparable material available at that time
precluded closer comparison of the two genera. On the basis of a
skull and jaw of Floridatragulus collected from the Thomas Farm
in 1964 by Stanley Olsen (now in the Frick Collection of the
AMNH), it now is possible to state that the skull assigned by
White (1942) to Floridatragulus clearly belongs to the same
species. Hypermekops olseni is thus a synonym of Floridatragulus
dolichanthereus. A similar conclusion has been reached by Clayton
Ray and by Bryan Patterson in unpublished studies of these forms
(McKenna, 1966).

The skull and jaw collected by Olsen were not actually articu-
lated but were recovered from the same part of the Thomas Farm
quarry (S. J. Olsen, personal communication ). The size, preserva-

Patron: Miocene Selenodonts 183

tion, degree of tooth wear, extreme length of snout, and corre-
spondence of occluding surfaces of these two specimens indicate
that they belong to the same individual. No other Thomas Farm
artiodactyl shows such remarkable morphological similarity to these
specimens. P*? in MCZ 3711 is represented only by alveoli. P? in
both specimens is slender, distinctly tricuspate, and has a weak
internal cingulum. More compelling evidence for the identity of
these two taxa is seen in the long, attenuated snout of both. In the
better preserved specimen (MCZ 3711) the elongated muzzle
possesses four caniniform teeth or alveoli. White (1942, p. 11-12)
interpreted these as representing I'-I*, C/, and P'. The largest
caniniform tooth in this series, according to White's interpretation,
would be I’. In the Frick specimen alveoli for the only two canini-
form teeth occur between P* and the largest caniniform. What
appears to be the maxillo-premaxillary suture in this specimen is
located just anterior to the large caniniform tooth. If the maxillo-
premaxillary suture can be used to separate I* and C/, the anterior-
most tooth in this series would be I’, the second (just posterior to
the maxillo-premaxillary suture) would be the upper canine, and
the subsequent three (on MCZ 3711) alveoli probably would repre-
sent a deciduous canine, P', and deciduous P'. The presence of
only two alveoli posterior to C/ in the Frick specimen indicates
that the retention of the more posterior deciduous teeth is some-
what variable. In any event, it is clear that the two skulls are too
similar to regard as representing separate taxa.

Floridatragulus barbouri White, 1947
Floridatragulus barbouri White, 1947, p. 505, fig. 4.

Two species of Floridatragulus are recognized in the Thomas
Farm Fauna, F. dolichanthereus White and F. barbouri White.
Floridatragulus dolichanthereus differs from F.. barbouri in its larger
size and in having a longer diastema between P, and P;. Remains
of Floridatragulus are not abundant in the Thomas Farm deposits,
and the sample available is small, but the differences appear to be
constant. Perhaps separation of the two species is justified best by
their stratigraphic occurrence.

It is uncertain whether F. dolichanthereus and F. barbouri lived
contemporaneously, or whether they represent an actual ancestor-
descendent lineage. The nature of the Thomas Farm deposit and

184 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

the history of its excavation (White, 1942; Bader, 1956) suggest
that those forms as they are known so far were separated tem-
porally. The largest species, F. dolichanthereus, was recovered from
the uppermost layers of the deposit and has not been found in any
of the deeper sediments (White, 1942, p. 30). The smaller less
advanced F. barbouri was recovered from a deeper portion of the
quarry. Hence, there is reason to accept the second alternative; that
F. barbouri is a representative of an earlier group which evolved
(there is no evidence to suggest replacement) into F. doli-
chanthereus.

Other evidence, based on associated fossil horses, also suggests
that the Thomas Farm fauna is heterochronic (Bader, 1956; Patton,
1964 ).

Other floridatragulines are present in Miocene deposits in the
Texas Coastal Plain (Patton, Ms). An as yet unstudied specimen
from the Garvin Gully Fauna appears to be similar to Florida-
tragulus barbouri, and F. dolichanthereus corresponds closely to
F. texanus from the Burkeville Fauna, differing from that species
only in having smaller premolars and a more invaginated talonid
on M,. A still larger form, F. hesperus, occurs in the Texas Cold
Spring Fauna; it has no known counterpart in Florida.

Discussion. Systematic placement of the Floridatragulinae is
uncertain. To date only relatively fragmentary and unassociated
material from Miocene deposits in Texas and Florida has been
studied and, expectedly, has revealed little in the way of phyletic
clues. Fortunately, skull and jaw material from the Thomas Farm
is now available, and it is hoped that study of this material will
help in understanding the puzzling systematics of this group. A few
preliminary remarks are in order, however.

Floridatragulus bears considerable resemblance to members of
two families of artiodactyls, the Hypertragulidae and the Cameli-
dae. Floridatragulus resembles members of the Hypertragulidae in
the presence of the intercolumnar pillars, a double enamel loop on
the heel of M;, rather prominent cingula, and in the occurrence
of a diastema between P, and P,. In the hypertragulids the de-
tached P., tends to be unicuspid; in Floridatragulus this tooth is
bicuspid, though weakly so. The anterior lower premolars of Flori-
datragulus are laterally compressed as in camels, and P,, although
more foreshortened than in most camels, retains the general features
of that group. In the skull and jaws of Floridatragulus from the

Patton: Miocene Selenodonts 185

Thomas Farm, now in the Frick Laboratory, the extremely elon-
gated muzzle has four caniniform teeth (or alveoli) on each side,
probably representing modifications of the incisors, C/, and P'.
This is a characteristic feature of the Tylopoda, in which the upper
incisors and anterior premolars may undergo reduction, but in which
there is never a complete loss of the upper incisors. In contrast,
the hypertragulids are marked by either extreme reduction or com-
plete suppression of the upper incisors, although they may be re-
tained in some of the very primitive types.

Because of its similarity to the camels, but also because of its
distinctness within that group, I have referred Floridatragulus to
the Camelidae but as a separate subfamily. Certainly those char-
acters which distinguish it from the Camelinae are equivalent in
taxonomic weight to those of other camel subfamilies, even, for
example, the Stenomylinae. Perhaps for parallel reasons, White
(1947) tentatively placed the genus Nothokemas (as the type of
a new family) in the Hypertragulonidea. Of these two problem-
atical genera, Nothokemas more closely approximates the features
of the Camelidae. In the unreduced condition of the premolars and
the configuration of both molars and premolars, the lower jaw of
Nothokemas is remarkably similar to Oxydactylus. As in Oxy-
dactylus, the body of the mandible is deep and thin, but the dias-
tema anterior to P, is considerably longer. The mandible in this
region is much more attenuated than in Oxydactylus. Both genera
are brachyodont and both display a relative elongation of the
molars.

In addition, if the skull (MCZ 4329) assigned to Nothokemas
by White (1947) does in fact belong to that genus, it can be seen
to possess a closed orbit, a more advanced feature not shared by
Floridatragulus. Although certain primitive characters possessed by
both Nothokemas and Floridatragulus are shared by members of
the Hypertraguloidea, e.g., intercolumnar tubercles, they are not
restricted to that group but may be regarded as archaic traits re-
tained by several disparate artiodactyl groups, including the Cam-
elidae. The increase in similarity between the camels and hyper-
tragulids as we look further back in geologic is well documented
(Scott, 1940; Matthew, 1905; Colbert, 1941; Simpson, 1945). The
two divergent groups are supposedly recognizable as early as the
Late Eocene, but in the Middle Miocene of the Gulf Coastal Plain
we have two genera which share few features in common with any

186 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

of the contemporary representatives of those basal stocks. Nothoke-
mas conceivably could be derived from some early Oxydactylus,
but that genus seems too advanced to have provided an ancestor
for Floridatragulus. Whether or not the morphological similarity
between either one or both of the Gulf Coast genera and some
members of the Camelidae is the result of recent common descent,
parallelism, or convergence is too fine a distinction to make with
the information available. However, the possibility remains that
both genera are descendants of an early and distinct branch off the
tylopod lineage which became isolated in the Gulf Coastal Plain
and occupied an adaptive zone in this region similar to that occu-
pied by some camels in the Great Plains. With the invasion of the
Gulf Coast by several diversified camel genera from the Great
Plains during the Early Pliocene or slightly earlier, the autoch-
thonous Gulf Coast genera became extinct.

Suborder RuMINANTIA Scopoli, 1777

Infraorder TRAGULINA Flower, 1883

Family Protoceratidae Marsh, 1891

Genus Prosynthetoceras Frick, 1937

Prosynthetoceras texanus (Hay), 1924

Dromomeryx texanus Hay; Hay, 1924, p. 15-16, pl. 11, figs. 8-12.
Dromomeryx angustidens Hay; Hay, 1924, p. 16, pl. 11, figs. 6-7.
Merycodus grandis Hay; Hay, 1924, p. 17-18, pl. 111, figs. 9-11.
Protolabis francisi Hay; Hay, 1924, p. 14, pl. 111, figs. 5, 6. (in part).
?Cranioceras texanus (Hay ); Frick, 1937, p. 82, 97.
?Synthetoceras rileyi Frick; Frick, 1937, p. 603, 605, fig. 66.
Blastomeryx texanus (Hay); Wood and Wood, 1937, p. 137, pl. 1, figs. 5, 6.
PSyndyoceras texanus (Hay); Hesse, 1942, p. 163 (?Syndeoceras texanus, p.

167, lapsus).

Syndyoceras australis White, 1941, p. 97, pl. XV, figs. 1, la, 2, 2a.
Synthetoceras (Prosynthetoceras ) douglasi White, 1947, p. 504, fig. 3a.
cf. Miolabis sp. indet., Simpson, 1932, p. 37.

cf. Miolabis tenuis, Ray, 1957, p. 18.

Nothokemas grandis White, 1947, p. 508. (in part).

Type. TAMU 2387, a right M® from the Garvin Gully Fauna
of Texas.

Two genera of synthetocerines, Syndyoceras and Synthetoceras
(Prosynthetoceras), have been recognized from fossil material re-
covered from the Thomas Farm. In 1941, white described Syndyo-
ceras australis on the basis of a right lower jaw with P, and P,-M;
(MCZ 3654, type), and a left maxilla with P*-M? (MCZ 3642, para-

Patron: Miocene Selenodonts 187

type). He also referred to this species three right mandibles and a
core of a postorbital horn. In 1947, White described Synthetoceras
(Prosynthetoceras) douglasi from a badly crushed palate with
P*-M? of both sides.

A comparison of the Thomas Farm specimens with material
from the Garvin Gully and Burkeville faunas of Texas shows that
the specimens assigned to S. australis White and P. douglasi White
are inseparable from P. texanus (Hay) and should be included in
the synonymy of that species. In cusp morphology as well as in
dimensions of the teeth and jaws, there appear to be no significant
differences between and among the specimens from both areas.
Inasmuch as I have not seen the horn core White (1942) referred
to Syndyoceras australis (it was not illustrated by White), no
comparisons could be made with the Texas material. However,
the horns, especially the rostral horn, of Prosynthetoceras texanus
are considerably advanced over those of Syndyoceras cooki from
the Great Plains. The evolution of the rostral horn of the synthe-
tocerines has involved primarily the lengthening of the main shaft,
resulting in an increase in the distance from the base of the rostral
horn to the point of bifurcation. In Syndyoceras the point of bifur-
cation is very close to the base of the horn; in fact the horns flare
away from each other at the point of union just above the nasal
passage. The rostral horn of Prosynthetoceras, on the other hand,
has lengthened to the degree where there is a definite shaft between
the maxillary union and the point of bifurcation.

In 1932, Simpson referred several teeth (FSGS V-4970) from
Midway, Florida, to the genus Miolabis but considered the species
indeterminate. He regarded these specimens as bearing consider-
able resemblance to Miolabis tenuis, but declined to designate them
as such. Ray (1957) lists this reference as “cf.Miolabis tenuis.”
A comparison of the original specimens from Midway with ma-
terial from Thomas Farm and Garvin Gully and Burkeville faunas
reveals no significant difference in size or morphology between the
teeth referred to Miolabis and those confidently assigned to
Prosynthetoceras texanus. Although isolated molars of Prosynthe-
toceras and Floridatragulus are very similar in morphology, those
of Floridatragulus are considerably larger than those of Prosynthe-
toceras in beds of the same age. For this reason, I have included
White (1947) to Nothokemas grandis is assigned to P. texanus.

188 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

A partial left maxilla with dP?-*-M! (MCZ 4328) referred by
White (1947) to Nothokemas grandis is assigned to P. texanus.
This immature dentition is far too small to belong to Nothokemas
but compares very closely to specimens of P. texanus from Texas
deposits.

For reasons I have given elsewhere (Patton, in press) I believe
that the elevation of Prosynthetoceras to generic rank more accu-
rately reflects its relative position in the taxonomic hierarchy of the
Synthetoceratinae. Thus, the synthetocerine material from the Gar-
vin Gully and Burkeville faunas of Texas and that from the Thomas
Farm of Florida are considered by the writer to belong to a single
species, which on the basis of priority, is designated Prosyn-
thetoceras texanus.

Infraorder Pecora Linnaeus, 1758
Family Cervidae Gray, 1821
Genus Blastomeryx Cope, 1877

Blastomeryx Parablastomeryx floridanus (White), 1940, p. 34.

The larger deer specimens from the Thomas Farm, including
not only White’s type (MCZ 3626) but several other specimens in
the collections of the University of Florida and the Florida State
Geological Survey, are all characterized by a relatively short post-
symphysial diastema. If Fricks (1937) criteria for separating
genera in his Division Blastomerycini are applied, the Thomas
Farm specimens would be assigned to the genus Parablastomeryx,
which White (1940) did. The systematics of this group, however,
is clearly in need of revision and, accordingly, it is often difficult
to decide on supraspecific placement. Not only is the validity of
certain morphological criteria in question, but the relative weight
of Frick’s supraspecific categories is sometimes indeterminable and
unmanageable. Until the confusion is removed, it seems advisable
to defer judgment on this assignment.

Genus Machaeromeryx Matthew, 1926
Machaeromeryx gilchristensis White, 1941

Machaeromeryx gilchristensis White, 1941, p. 97, pl. 14, fig. 5

Generic assignment for this tiny ruminant appears to be valid.
However, because of the scanty and fragmentary nature of the

Patton: Miocene Selenodonts 189

Thomas Farm specimens specific comparisons between it and speci-
mens assigned to the type species from the Upper Harrison beds of
Nebraska are not likely to be particularly useful. In view of the
great geographic separation between the two occurrences, specific
differentiation is probable.

In summary, the revised list of selenodont artiodactyls of the
Thomas Farm Fauna is as follows:

Family Camelidae

Subfamily Camelinae
Nothokemas floridanus (Simpson), 1932

Subfamily Floridatragulinae!

Floridatragulus dolichanthereus White, 1940
Floridatragulus barbouri White, 1947

Family Protoceratidae
Prosynthetoceras texanus ( Hay), 1924

Family Cervidae

Blastomeryx gilchristensis White, 1941
Machaeromeryx gilchristensis White, 1941

LITERATURE CITED

Baver, R. S. 1956. A quantitative study of the Equidae of the Thomas
Farm Miocene. Bull. Mus. Comp. Zool., vol. 115, no. 2, pp. 49-78.

CotsperT, E. H. 1941. The osteology and relationships of Archaeomeryx, an
ancestral ruminant. Amer. Mus. Novit., no. 1135, pp. 1-24.

Frick, C. 1937. Horned ruminants of North America. Bull. Amer. Mus.
Nat. Hist., vol. 69, pp. 1-669.

MatrHew, W. D. 1905. Notice of two new genera of mammals from the
Oligocene of South Dakota. Bull. Amer. Mus. Nat. Hist., vol. 21,
pp. 21-26.

McKenna, M. C. 1966. Synopsis of Whitneyan and Arikareean camelid
phylogeny. Amer. Mus. Novit., no. 2253, pp. 1-11.

1Through coincidence while this paper was passing through the press,
the subfamily Floridatragulinae was described as new by another author,
Vincent Joseph Maglio (A revision of the fossil selenodont artiodacytls from
the Middle Miocene Thomas Farm, Gilchrist County, Florida, Breviora, Museum
of Comparative Zoology, no. 255, pp. 1-27, figs. 1-4, “December 6, 1966,”
[postmarked January 20, 1967]).

190 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

OtsEN, S. J. 1962. The Thomas Farm fossil quarry. Quart. Jour. Florida
Acad. Sci., vol. 25, no. 2, pp. 142-146.

Patron, T. H. 1964. The Thomas Farm fossil vertebrate locality. Guide-
book for the 1964 Field Trip of the Society of Vertebrate Paleontology
in Central Florida, pp. 12-20. (mimeographed ).

. 1967. Reévaluation of Hay’s artiodactyl types from the Miocene
of the Texas Coastal Plain. Texas Jour. Sci., vol. 10, no. 1, pp. 35-40.

—. Ms. Miocene artiodactyls, Texas Coastal Plain. Unpublished Ph.D.
dissertation, University of Texas, 166 pp.

Ray, Ciaytron E. 1957. A list, bibliography, and index of the fossil vertebrates
of Florida. Florida Geol. Survey, Spec. Publ. no. 3, 175 pp.

Romer, A. S. 1948. The fossil mammals of Thomas Farm, Gilchrist County,
Florida. Quart. Jour. Florida Acad. Sci., vol. 10, no. 1, pp. 1-11.

Scott, W. B. 1940. The mammalian fauna of the White River Oligocene.
Pt. 4, Artiodactyla. Trans. Amer. Phil. Soc., new series, vol. 28, pt. 4,
pp. 363-746.

Simpson, G. G. 1932. Miocene land mammals of Florida. Bull. Amer. Mus.
Nat. Hist., vol. 59, art. 11, pp. 149-211.

. 1945. The principles of classification and a classification of mammals.
Bull. Amer. Mus. Nat. Hist., vol. 85, pp. 1-350.

Wuire, T. E. 1940. New Miocene vertebrates from Florida. Proc. New
England Zool. Club, vol. 18, pp. 31-38.

. 1941. Additions to the Miocene fauna of Florida. Proc. New England
Zool. Club, vol. 18, pp. 91-98.

. 1942. The lower Miocene mammal fauna of Florida. Bull. Mus.
Comp. Zool., vol. 92, no. 1, pp. 1-49.

. 1947. Additions to the Miocene fauna of north Florida. Bull. Mus.
Comp. Zool, vol. 99, no. 4, pp. 497-515.

Florida State Museum and Department of Zoology, University
of Florida, Gainesville, Florida.

Quart. Jour. Florida Acad. Sci. 29(3) 1966 (1967 )

Distribution of Euglenida in North Florida
Joun J. McCoy

THE great diversity of aquatic situations in the State of Florida
lends itself ideally to limnological and ecological studies of the
microfauna of various habitats. During the course of this study, a
number of these habitats in north and central Florida were peri-
odically sampled for determination of their protozoan fauna. This
paper represents a compilation of the data for the order Euglenida.

DESCRIPTION OF HABITATS

Large Streams. One river, the Santa Fe, was sampled peri-
odically during this study. From its source near Lake Santa Fe,
Bradford-Alachua County, it flows westerly and slightly to the north,
to a point near Branford, Suwannee County, where it empties into
the Suwannee River. The Santa Fe is a “black” river, colored by
the addition of various organic materials derived from pine flat-
woods and cypress swamps. Four sampling stations were used.
These were at O’Leno State Park in Columbia County, Worthington
Springs in Union County, four miles south of Starke on U.S. Route
301 in Bradford County, and at the rivers mouth in Suwannee
County. :

Small Streams. Hogtown Creek in Gainesville, Alachua Coun-
ty, was sampled during this study. This creek is a relatively shallow
stream with moderate flow. Both sandy and silty bottom conditions
occur. Samples were taken at a number of randomly picked points.

Sinks and Lakes. Sinks and lakes are conspicuous elements in
the aquatic situations of north and central Florida. Mainly of solu-
tion origin, these vary in size from a few feet in diameter to rela-
tively large bodies of water. Bottom conditions range from sand to
silt, and the amount of rooted and floating vegetation varies con-
siderably. Lake Alice, Lake Santa Fe, Lake Newnan, and Lake
Hampton in Alachua County and Lake Kingsley in Bradford Coun-
ty were periodically sampled. In addition, samples were taken from
several sinks found on the campus of the University of Florida.

Cypress Ponds. Cypress ponds are numerous in north and cen-
tral Florida. Fluctuating water levels create a variable habitat fre-
quently reflected by changes in the species composition of the
microfauna inhabiting these ponds. A number of ponds near Otter

192 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Creek in Levy County, and several smaller ponds in Alachua Coun-
ty were sampled. The latter were located within the Gainesville
city limits, and near the large sink known locally as the Devil’s
Millhopper.

Miscellaneous. In addition to the habitats listed above, a num-
ber of temporary or artificial aquatic situations were sampled. These
included numerous roadside ditches throughout north and central
Florida, settling lagoons for sewage disposal plants in Ocala and
Gainesville, and a small pond, hereafter referred to as Campus
Pond, behind the sewage disposal plant for the University of
Florida. Campus Pond is maintained by receiving final effluent from
the disposal plant.

METHODS

With the exception of roadside ditches, habitats were sampled
monthly. The temporary nature of such ditches prevented periodic
sampling. Samples were taken from the bottom, surface, and at
intermediate depths. Formaldehyde was added to these as a pre-
servative. Temperature and pH data were recorded at the time of
collecting. Samples were centrifuged and subsamples of the pre-
cipitate examined.

FAUNAL LIsT

Euglena acus Ehrenberg. Santa Fe River, Hogtown Creek, sinks, Lake Alice,
cypress ponds, roadside ditches, Campus Pond.

Euglena sciotensis Lackey. Campus Pond.

Euglena oxyuris Schmarda. Hogtown Creek, cypress ponds, roadside ditches,
Campus Pond.

Euglena spadix Gojdics. Sinks.

Euglena oblonga Schmitz. Sinks.

Euglena mutabilis Schmitz. Santa Fe River.

Euglena rostrifera Johnson. Hogtown Creek, settling lagoons, Cypress Pond.

Euglena gaumei Allorge and Lefevre. Cypress ponds, Campus Pond.

Euglena flava Dangeard. Sinks, Lake Santa Fe, Campus Pond.

Euglena agilis Carter. Settling lagoons.

Euglena fusca (Klebs) Lemmermann. Lake Newnan, cypress ponds, Campus
Pond.

Euglena helicoideus (Bernard) Lemmermann. Cypress ponds.

Euglena vivida Playfair. Roadside ditches.

Euglena convoluta Korshikov. Cypress Ponds.

Euglena allorgei Deflandre. Campus Pond.

Euglena chadefaudii Bourrelly. Sinks.

Euglena limnophila Lemmermann. Santa Fe River, Hogtown Creek, sinks,
cypress ponds, roadside ditches.

McCoy: Euglenida in Florida 193

Euglena spirogyra Ehrenberg. Hogtown Creek, cypress ponds, Campus Pond.

Euglena hemichromata Skuja. Sinks, settling lagoons.

Euglena mangini Lefevre. Santa Fe River, Campus Pond.

Euglena ehrenbergii Klebs. Lake Santa Fe, cypress ponds, roadside ditches,
Campus Pond.

Euglena grisoli Deflandre. Hogtown Creek.

Euglena velata Klebs. Santa Fe River, settling lagoons, Campus Pond.

Euglena splendens Dangeard. Cypress ponds.

Euglena chlamydophora Mainx. Cypress ponds.

Euglena mainixi Deflandre. Campus Pond.

Euglena deses Ehrenberg. Hogtown Creek.

Euglena proxima Dangeard. Santa Fe River, Lake Santa Fe, Campus Pond.

Euglena subehrenbergii Skuja. Campus Pond.

Euglena elenkinii Poljanski. Campus Pond.

Euglena tuberculata Drezepolski. Santa Fe River, cypress ponds, Campus
Pond.

Trachelomonas volvocina Ehrenberg. Hogtown Creek, sinks, Santa Fe River,
Lake Santa Fe, Lake Newnan, Lake Hampton, cypress ponds, roadside
ditches, Campus Pond.

Trachelomonas hispida (Perty) Stein. Santa Fe River, Hogtown Creek, sinks,
Lake Alice, Lake Santa Fe, Lake Hampton, cypress ponds, roadside
ditches, settling lagoons, Campus Pond.

Trachelomonas caudata (Kefferath) Conrad. Lake Alice.

Trachelomonas armata (Ehrenberg) Stein. Cypress ponds, roadside ditches.

Trachelomonas fusiformis Deflandre. Cypress ponds.

Trachelomonas urceolata Stokes. Sinks, Campus Pond.

Trachelomonas acanthostoma Stokes. Sinks, cypress ponds.

Trachelomonas rotunda Swirenko. Santa Fe River, cypress ponds, Campus
Pond.

Trachelomonas gibberosa Playfair. Cypress ponds, Campus Pond.

Trachelomonas oblonga Lemmermann. Santa Fe River, cypress ponds.

Trachelomonas abrupta Swirenko. Santa Fe River, Lake Hampton.

Trachelomonas superba Swirenko. Lake Hampton.

Trachelomonas pulcherrima Playfair. Cypress ponds, Campus Pond.

Trachelomonas polonica Drezepolski. Cypress ponds.

Trachelomonas bacillifera Playfair. Sinks, cypress ponds, Campus Road.

Trachelomonas zorensis Deflandre. Lake Hampton, Campus Pond.

Trachelomonas acuminata (Schmarda) Stein. Santa Fe River, sinks, Campus
Pond.

Trachelomonas rugulosa Stein. Lake Newnan.

Trachelomonas speciosa Deflandre. Cypress ponds.

Trachelomonas intermedia Dangeard. Cypress ponds.

Trachelomonas angustispina Deflandre. Cypress ponds, Santa Fe River.

Trachelomonas allia Drezepolski. Cypress ponds, roadside ditches.

Trachelomonas verrucosa Stokes. Campus Pond.

Trachelomonas perforata Awer. Cypress ponds, roadside ditches, Campus
Pond.

Trachelomonas allorgei Deflandre. Campus Pond.

194 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Trachelomonas subglobosa Skvortzov. Santa Fe River.

Trachelomonas tambowika Swirenko. Santa Fe River.

Trachelomonas dubia Swirenko. Cypress ponds.

Trachelomonas hexangulata Swirenko. Cypress ponds.

Trachelomonas lacustris Drezepolski. Cypress ponds.

Trachelomonas giardiana Playfair. Cypress ponds, Campus Pond.

Trachelomonas woycickii Koczwara. Lake Hampton.

Trachelomonas spectabilis Deflandre. Cypress ponds.

Trachelomonas globularis (Awer.) Lemmermann. Santa Fe River, cypress
ponds.

Trachelomonas magdaleniana Deflandre. Cypress ponds.

Trachelomonas bernardinensis Vischer. Cypress ponds.

Trachelomonas sarmatica Drezepolski. Cypress ponds.

Trachelomonas niklewskii Drezepolski. Cypress ponds.

Trachelomonas lackeyi McCoy. Campus Pond.

Lepocinclis ovum (Ehrenberg ) Lemmermann. Santa Fe River, Hogtown Creek,
sinks, Lake Alice, cypress ponds, Campus Pond.

Lepocinclis marssonii Lemmermann. Santa Fe River, cypress ponds.

Lepocinclis texta (Dujardin) Lemmermann. Hogtown Creek, sinks, settling
lagoons, Campus Pond.

Lepocinclis cymbiformis Playfair. Sinks, cypress ponds, Campus Pond.

Lepocinclis glabra Drezepolski. Cypress ponds, Campus Pond.

Lepocinclis globosa France. Campus Pond.

Phacus curvicauda Swirenko. Santa Fe River, sinks, Campus Pond.

Phacus alatus Klebs. Sinks, Campus Pond.

Phacus pusilus Lemmermann. Sinks.

Phacus horridus Pochmann. Cypress ponds.

Phacus glaber (Deflandre) Pochmann. Sinks, cypress ponds, Campus Pond.

Phacus inflexus Pochmann. Campus Pond.

Phacus pyrum (Ehrenberg) Stein. Sinks.

Phacus circumflexus Pochmann. Campus Pond.

Phacus hamatus Pochmann. Campus Pond.

Phacus corculum Lemmermann. Campus Pond.

Phacus caudatus Hubner. Lake Alice, cypress ponds.

Phacus onyx Pochmann. Cypress ponds, roadside ditches, settling lagoons,
Campus Pond.

Phacus undulatus Pochmann. Sinks, cypress ponds.

Phacus margiritatus Pochmann. Santa Fe River, sinks.

Phacus granum Drezepolski. Campus Pond.

Phacus thrombus Pochmann. Sinks, Campus Pond.

Phacus segreti Allorge and Lefevere. Cypress ponds.

Phacus ranula Pochmann. Sinks.

Phacus suecicus Lemmermann. Santa Fe River, sinks, cypress ponds, Campus
Pond.
Phacus tortus (Lemmermann) Skvortzov. Sinks, cypress ponds, Campus

Pond.
Phacus trypanon Pochmann. Cypress ponds, Campus Pond.
Phacus musculus Pochmann. Sinks, cypress ponds.

McCoy: Euglenida in Florida 195

Phacus atrakoides Pochmann. Cypress ponds, Campus Pond.

Phacus acuminatus Stokes. Sinks, cypress ponds, Campus Pond.

Phacus platalea Drezepolski. Cypress ponds, Campus Pond.

Phacus longicauda (Ehrenberg) Dujardin. Sinks, cypress ponds, roadside
ditches, Campus Pond.

Phacus orbicularis Hubner. Sinks.

Phacus wettsteini Drezepolski. Campus Pond.

Phacus makrostigma Pochmann. Sinks, cypress ponds.

Phacus carinatus Pochmann. Sinks, Campus Pond.

Phacus circulatus Pochmann. Cypress ponds.

Phacus pleuronectes (O. F. Muller) Dujardin. Sinks, cypress ponds, Campus
Pond.

Phacus raciborski Drezepolski. Cypress ponds.

Phacus aenigmaticus Drezepolski. Campus Pond.

Cryptoglena pigra Ehrenberg. Sinks, cypress ponds, Campus Pond.

Anisonema steini Stokes. Cypress ponds.

Anisonema grande Stein. Sinks

Entosiphon sulcatum (Dujardin) Stein. Sinks.

Entosiphon ovatum Stokes. Sinks.

Menoidium gracile Stokes. Sinks, cypress ponds, Campus Pond.

Menoidium tortuosum Stokes. Sinks.

Menoidium pelucidom Perty. Sinks.

Distigma proteus Ehrenberg. Sinks, cypress ponds.

Peranema tricophorum Ehrenberg. Campus Pond.

Euglenopsis vorax Klebs. Sinks.

DISCUSSION

Eleven genera representing 121 species were recorded during
the course of this study. Cypress ponds, Campus Pond, and sinks
were the habitats containing the greatest number of species. For
each habitat, the percentage of the total number of species found
is as follows: cypress ponds, 52 per cent; Campus Pond, 49 per
cent; sinks, 35 per cent; Santa Fe River, 18 per cent; lakes (com-
bined) 13 per cent; ditches, 9 per cent; Hogtown Creek, 8 per
cent; settling lagoons, 6 per cent. A given species was frequently
found in more than one habitat. For each genus, the largest
number of species was found in either cypress ponds, Campus
Pond, or sinks (Table 1).

Cypress ponds, sinks, and Campus Pond contained 109 of the
121 recorded species, 75 of which were only found here. A total
of 44 species was recorded from more than one of these habitats
with the following distribution: cypress ponds and sinks, 7 species;
cypress ponds and Campus Pond, 15 species; sinks and Campus
Pond, 8 species; cypress ponds, sinks, and Campus Pond, 14 species.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

196

6S L IT €9 9T GV OT 6G IGI s[P}0,L,
Bre C0 te 0 e208 eee ely Oe a ee
if 0 0 0 0 0 0 0 I DUWAaUDIAg
0 0 0 it 0 I 0 0 if DWS14S1C
T 0 0 I 0 € 0 0 3 uni p1oUua J
0 0 0 0 0 G 0 0 6 uoydisojuy
0 0 0 I 0 I 0 0 C pwauosiuy
I 0 0 I 0 I 0 0 I puajs0jdhuy
G I 0 V I S G G 9 syoumao0daT
1G I G 8I I SI 0 € VE snovud
el I V GG 6 G I OL 6E SDUOUWLO]OYIDL I,
LI V g GL q L L L T€ puajany
puog suoosey soyoyiq spuog soyr'T SyUIS Yeon IOAN papsloooyy snuosy
sndwey) suljyeg epispeoy ssaiddry UMO}JSOF] IY vYURG satoeds

vplugjsny Jo saloadg jo yeyqey
T @TaVv.L

McCoy: Euglenida in Florida 197

Environmental conditions in the habitats above were apparently
more favorable to euglenoids than those of other sample areas. The
number of shared species suggest important similarities between
these habitats. With the limited data available, such conditions
and similarities could not be determimned. The pH of these habi-
tats was variable. Sinks were alkaline with a range of 7.1 to 7.8
and a mean of 7.3. Campus Pond ranged from 6.7 to 8.4 with
a mean of 7.5. Cypress ponds were acid with a range of 6.4 to
6.8 and a mean of 6.6. Temperature was not considered to be
significant.

The data in Table 1 indicate a variability in habitat preference
for members of the order. The genus Trachelomonas was repre-
sented in cypress ponds by 25 species. This is nearly twice the
number of trachelomonads present in any other habitat. When
considering that 12 of these species were recorded only from
cypress ponds, a decided preference for such a habitat seems in-
dicated. The acid condition prevalent in cypress ponds may be
significant in this regard. According to Hall (1965), pH may affect
the utilization of nitrogen sources by flagellates.

Single-habitat preference appears to be lacking for most of the
other genera. The genus Euglena was represented in Campus Pond
by 17 species, and in cypress ponds, sinks, Hogtown Creek, and the
Santa Fe River by 12, 7, 7, and 7 species respectively. The mini-
mum number of species of Euglena recorded from any one habitat
was five, as contrasted with one for Trachelomonas, and relatively
few species of this genus were recorded from only a single habitat.
Distribution of the six species of Lepocinclis was comparable to
that for Euglena.

The genus Phacus was found to be well represented in and
nearly restricted to three habitats, i.e., Campus Pond, cypress ponds,
and sinks, with approximately the same number of species found in
each. All of the 34 recorded species were found in at least one of
these habitats, and 17 were found in more than one. Cryptofilena
was restricted to these same three habitats.

The remaining genera, Anisonema, Entosiphon, Menodium,
Distigma, Peranema, and Euglenopsis, are colorless forms, and are
generally similar in their patterns of distribution. Eight of the ten
recorded species were found in sinks. Species were also recorded
from cypress ponds and Campus Pond.

198 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SUMMARY

1. Eleven genera representing 121 species of the Order Euglenida
were recorded from various aquatic habitats in Florida.

bo

Cypress ponds, sinks, and a pond receiving final efhuent from
a sewage disposal plant were found to contain the largest
numbers of species.

3. Single habitat preference, i.e., cypress ponds, was indicated
for Trachelomonas, but most of the other euglenoids were more
widely distributed.

ACKNOWLEDGEMENTS

The author is greatly indebted to Dr. James B. Lackey of the
University of Florida for his encouragement and help during the
course of this research. I am indebted to Dr. Richard M. Johnson
of Asheville-Biltmore College for aid in collecting and his many

helpful criticisms.
LITERATURE CITED

Hari, R. P. 1965. Protozoan nutrition. Blaisdell Publishing Co., New York.
90 pp.

Department of Biology, Asheville-Biltmore College, Asheville,
INEZG.

Quart. Jour. Florida Acad. Sci. 29(3). 1966 (1967)

Two New Subfamilies of Monogenenetic Trematodes
C. E. PRicr

An evolutionary sequence of genera within the monogenetic
trematode family Dactylogyridae will go far toward improving the
current state of classification of this family. As some point of
reference is a basic necessity, the most variable and most readily
observable structures are best utilized as the main bases of such a
proposal. Among the monopisthocotylean monogenetic trematodes
two highly variable entities exist, the copulatory complex (sclero-
tized male genital organs) and the attaching armament of the
haptor (posterior attaching disc). Of the two, the sclerotized por-
tions of the haptor seem to furnish more significant information.

During the course of this study, it has become apparent that a
definite need exists for establishment of two additional subfamilies
within the Dactylogyridae. Recognition of these subfamilies should
aid in better understanding the evolution within this family.

CuRRENT TAXONOMIC STATE OF THE FAMILY

In his monograph concerning the Monogenea, Yamaguti (1963)
lists four subfamilies as comprising the Dactylogyridae: Dactylogy-
rinae Bychowsky, 1933; Ancyrocephalinae Bychowsky, 1937; Lingu-
adactylinae Bychowsky, 1957; and Geneticoenterinae Yamaguti,
1963.

One of the major characters differentiating the various sub-
families is the number of anchors (large haptoral hooks) present
in the haptor. As pointed out by Yamaguti, the number of anchors
is a sound trait on which to base subfamilial classification.

Members of the Dactylogyrinae possess a single pair of anchors.
The species belonging to this taxon are morphologically homo-
geneous and seem to warrant subfamily status without question.

More study is required before acceptance or refutation of the
proposed subfamilies Geneticoenterinae and Linguadactylinae. Both
are apparently closely related to the subfamily Ancyrocephalinae.

It is the subfamily Ancyrocephalinae which represents what is
probably the most confused area among the many confused areas
of classification within the suborder Monopisthocotylea. Presently
containing more than 50 genera, the Ancyrocephalinae is over-
burdened with taxonomic errors, inconsistencies, and synonymy. It

200 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

is hoped that the present work will make a worthwhile contribution
to the taxonomy of this group.

PRopPOsED NEw SUBFAMILIES

All but three of the 50 or so genera currently included in the
Ancyrocephalinae possess four anchors in the haptor. Heteronchoc-
leidus Bychowsky (1957) and Trianchoratus C. Price and Berry
(1966) are monotypic genera whose members possess three anchors.
The genus Anacanthorus Mizelle and C. Price (1965) contains three
species, all of which lack anchors entirely.

The atypical architecture of the haptors of these three genera
is here interpreted by the present author as being of a highly
divergent nature, and these genera should not be grouped with
parasites possessing four anchors. These divergent genera quite
definitely belong in the Dactylogyridae, but in subfamilies other
than the Ancyrocephalinae.

As the three involved genera were originally placed in the sub-
family Ancyrocephalinae on several bases, thus establishing them
as showing affinities with that group, the subfamily diagnoses can
be given quite simply.

HETERONCHOCLEIDINAE n. subfam. Dactylogyridea: Dactylogy-
ridae. With characters of the superfamily and family, some char-
acters of the Ancyrocephalinae. Three anchors, one median and a
bilaterally symmetrical pair posterolateral. Hooks 14 or 16. In-
cluded genera: Heteronchocleidus and Trianchoratus.

ANACANTHORINAE n. subfam. Dactylogyridea: Dactylogridae.
With characters of the superfamily and family, and some char-
acters of the Ancyrocephalinae. Anchors lacking. Hooks 18. Only
included genus: Anacanthorus.

Key TO SUBFAMILIES OF DACTYLOGYRIDAE

As more study is required concerning the validity or proper
status of the subfamilies Geneticoenterinae Yamaguti and Lingua-
dactylinae Bychowsky, they are omitted from the following key.

1 Eaton without amc tons see se eee eee Anacanthorinae
Taz. Haptor..with anchors) 22 ea oe ee ee 2
DE rlaptor withe2ranc Ors: Glycine Dactylogyrinae
War Eeyore \iaid oie 8) Cun oe ee Heteronchocleidinae

Die Haptonawath:4ancnorsn (Ce pains) esa eaten ean enn ee Ancyrocephalinae

Price: New Trematode Subfamilies 201

SUMMARY

Three divergent genera of monogenean trematodes have been
removed from the subfamily Ancyrocephalinae and placed in newly
established subfamilies of the family Dactylogyridae. This taxono-
mic adjustment is made primarily on the number of anchors present
in the haptor. The genera Heteronchocleidus Bychowsky and
Trianchoratus C. Price and Berry are placed in the new subfamily
Heteronchocleidinae; members of these genera possess three hap-
toral anchors. The genus Anacanthorus Mizelle and C. Price, the
members of which lack anchors entirely, is placed in the new sub-
family Anacanthorinae.

LITERATURE CITED

BycHowsky, B. E. 1957. Monogenetic trematodes-their systematics and
phylogeny. (Russian text, translated into English). American editor
W. J. Hargis, Jr. Sponsored by AIBS. Graphic Arts Press, Inc., Wash-
ington, D. C., 627 pp.

Mizexe, J. D., AND C. E. Price. 1965. Studies on monogenetic trematodes.
XXVIII. Gill parasites of the piranha, with proposal of Anacanthorus
gen. n. Jour. Parasitol. vol. 51, no. 1, pp. 30-36.

Prick, C. E. anp W. S. Berry. 1966. Trianchoratus, a new genus of Mono-
genea. Proc. Helminthol. Soc. Washington, vol. 33, no. 2, pp. 201-203.

YaMacutTl, S. 1963. Systema helminthum. Vol. IV: Monogenea and Aspi-
docotylea. Interscience Publishers, New York, 699 pp.

Department of Biology, North Texas State University, Denton,
Texas 76203. Present address: Augusta College, Augusta, Georgia
30904.

Quart. Jour. Florida Acad. Sci. 29(3) 1966 (1967)

Mayfly Nymphs from Northwestern Florida
ROBERT F. SCHNEIDER

SINCE publication of The Mayflies of Florida (Berner, 1950),
13 species of Ephemeroptera have been added to the fauna of the
northwestern part of the state. This paper gives the occurrence
of species now known from each of the nine major rivers, from the
Perdido on the western border of the state, eastward to the Es-
cambia, Blackwater, Yellow, Shoal, Choctawhatchee, Holmes, Chi-
pola, and Apalachicola, respectively. The records of Berner
(1950, 1955, 1958) are included, as well as new distributional data
obtained by the writer during investigation of water pollution for
the Florida State Board of Health from 1960-1965, and differenti-
ated in the list by asterisks. The habitat in which the nymphs were
collected is also given.

Family EPHEMERIDAE

Hexagenia bilineata (Say). Apalachicola River. Leaf detritus
and silt bottom.

Hexagenia munda elegans Traver. Apalachicola River. Leaf
detritus and silt bottom.

Hexagenia munda marilandica Traver. All nine rivers. Leaf
detritus and silt bottom.

Pentagenia vittigera (Walsh). Apalachicola River. Silt bottom.

Family NEOEPHEMERIDAE

Neoephemera compressa Berner. Apalachicola. Vegetation.
Neoephemera youngi Berner. *Perdido, *Yellow, *Chipola, Apa-
lachicola. Vegetation and silt bottom.

Family CAENIDAE

Brachycercus sp. A. Shoal, Choctawhatchee, Apalachicola. Leaf
detritus, silt bottom, sand bottom. Recorded as Brachycercus sp. A
by Berner (1950).

Brachycercus sp. *Escambia, *Yellow. Leaf detritus, silt, sand.

Caenis diminuta Walker. *Escambia, *Yellow, *Shoal, *Choc-
tawhatchee, Holmes, Chipola, Apalachicola. Leaf detritus, silt
bottom.

SCHNEIDER: Mayfly nymphs 203

Caenis hilaris (Say). *Perdido, *Escambia, *Yellow, *Shoal,
*Choctawhatchee, *Holmes, Chipola, Apalachicola. Leaf detritus,
clay bottom, vegetation, log or board, sand bottom.

Family BAETISCIDAE

Baetisca escambiensis Berner. Escambia. Silt bottom, sand
bottom.

Baetisca becki Schneider and Berner. *Perdido, *Blackwater,
*Shoal. Gravel or rock, sand bottom.

Baetisca gibbera Berner. Escambia, *Shoal, *Choctawhatchee,
Apalachicola. Log or board, sand bottom.

Baetisca obesa (Say). *Blackwater, *Yellow, *Choctawhatchee,
*Chipola, Apalachicola. Leaf detritus, vegetation, log or board,
silt bottom.

Baetisca rogersi Berner. Perdido, Escambia, Blackwater, Yellow,
Shoal, Choctawhatchee, Apalachicola. Gravel or rock, log or board,
silt bottom, sand bottom.

Family LEPTOPHLEBIIDAE

Choroterpes hubbelli Berner. Choctawhatchee. Silt bottom,
sand bottom.

Habrophlebiodes brunneipennis Berner. *Perdido, *Escambia,
Apalachicola. Leaf detritus, silt bottom, sand bottom.

Habrophlebia vibrans Needham. *Perdido, *Escambia, * Black-
water, Apalachicola. Leaf detritus.

Leptophlebia intermedia (Traver). Perdido, Escambia, Shoal,
Choctawhatchee, Chipola, Apalachicola. Leaf detritus.

Paraleptophlebia bradleyi (Needham). Perdido, *Escambia,
“Yellow, *Shoal. Leaf detritus, vegetation, silt bottom.

Paraleptophlebia volitans McDunnough. Perdido, Escambia,
Blackwater, Shoal, Choctawhatchee, Holmes, Apalachicola. Leaf
detritus, gravel or rock, vegetation, log or board.

Family EPHEMERELLIDAE

Ephemerella choctawhatchee Berner. *Perdido, *Escambia,
*Blackwater, Choctawhatchee, Chipola. Vegetation, sand bottom.
Ephemerella hirsuta Berner. Perdido, *Escambia, Apalachicola.

204 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Leaf detritus, vegetation.
Ephemerella rotunda Morgan. Apalachicola. Vegetation.
Ephemerella deficiens Berner. Escambia, Blackwater, Yellow,
Shoal, Chipola, Apalachicola. Leaf detritus, vegetation.
Ephemerella trilineata Berner. All nine rivers. Leaf detritus,
vegetation, log or board.
Ephemerella dorothea Needham. *Choctawhatchee, *Holmes.
Leaf detritus, vegetation.

Family TricoRYTHIDAE

Tricorythodes albilineatus Berner. Perdido, Escambia, Black-
water, Yellow, Choctawhatchee, Chipola, Apalachicola. Clay bot-
tom, vegetation, log or board, silt bottom, sand bottom.

Family SIPHLONURIDAE

Isonychia fattigi Traver. Apalachicola. Leaf detritus.

Isonychia pictipes Traver. *Perdido, Holmes, Chipola, Apalachi-
cola. Gravel or rock, log or board, sand bottom.

Isonychia sp. A. Perdido, Escambia, Blackwater, Shoal. Leaf
detritus, log or board, sand bottom.

Isonychia sp. B. *Shoal, Holmes. Vegetation, log or board.

Isonychia sp. G. *Chipola, Apalachicola. Leaf detritus, log or
board, sand bottom.

Family HEPTAGENIIDAE

Heptagenia flavescens (Walsh). *Perdido, *Shoal, Apalachi-
cola. Leaf detritus, gravel or rock, vegetation.

Stenonema exiguum Traver. All nine rivers. Gravel or rock,
vegetation, log or board.

Stenonema interpunctatum (Say). Apalachicola. Vegetation.

Stenonema proximum Traver. All nine rivers. Gravel or rock,
vegetation, log or board.

Stenonema smithae Traver. All nine rivers. Leaf detritus,
gravel or rock, vegetation, log or board.

Family AMETROPODIDAE

Siphloplecton speciosum Traver. Perdido, Blackwater, Shoal,
Choctawhatchee, Chipola, Apalachicola. Vegetation.

SCHNEIDER: Mayfly nymphs 205

Family BAETIDAE

Baetis australis Traver. Perdido, Chipola, Apalachicola. Leaf
detritus, vegetation, log or board, silt bottom, sand bottom.

Baetis ephippiatus (Traver). Perdido, Escambia, Choctawhat-
chee, Holmes, Apalachicola. Vegetation.

Baetis intercalaris McDunnough. Holmes, Chipola, Apalachi-
cola. Vegetation, sand bottom.

Baetis propinquus (Walsh). Apalachicola. Gravel or rock,
vegetation.

Baetis spiethi Berner. Perdido, Choctawhatchee, Holmes, Chi-
pola, Apalachicola. Vegetation, log or board.

Baetis spinosus McDunnough. All nine rivers. Leaf detritus,
vegetation, log or board, silt bottom, sand bottom.

Callibaetis floridanus Banks. *Perdido, “Escambia, *Choctaw-
hatchee, *Holmes, Chipola, Apalachicola. Leaf detritus, gravel or
rock, vegetation.

Centroptilum hobbsi Berner. *Perdido, *Escambia, *Black-
water, *Yellow, *Shoal, *Choctawhatchee, Chipola, Apalachicola.
Vegetation, log or board.

Centroptilum viridocularis Berner. “Escambia, *Blackwater,
*Yellow, Choctawhatchee, Holmes, Chipola, Apalachicola. Vegeta-
tion, sand bottom.

Cloeon rubropictum McDunnough. Apalachicola. Vegetation,
log or board.

Cloeon sp. A. Holmes, Apalachicola. Vegetation, log or board.

Pseudocloeon alachua Berner. *Escambia, *Holmes, *Chipola.
Vegetation.

Pseudocloeon bimaculatum Berner. Perdido, Escambia, Black-
water, Yellow, Shoal, Choctawhatchee, Chipola. Leaf detritus,
gravel or rock, vegetation, sand bottom.

Pseudocloeon parvulum McDunnough. *Holmes, Chipola, Apa-
lachicola. Gravel or rock, vegetation.

Pseudocloeon punctiventris McDunnough. *Blackwater, *Choc-
tawhatchee, Apalachicola. Gravel or rock, vegetation, sand botton.

Family BEHNINGIDAE.

Dolania americana Edmunds and Traver. *Sweetwater Creek
(a tributary of Blackwater River). Sand bottom. The only pre-
vious record of this family in the New World was the original

206 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

description of D. americana, based on five nymphs from the Sa-
vannah River system in South Carolina (Edmunds and Traver,
1959).

ACKNOWLEDGEMENT

Sincere appreciation is expressed to Dr. Lewis Berner for his
suggestions and criticisms in the preparation of this manuscript.

LITERATURE CITED

BERNER, Lewis. 1950. The mayflies of Florida. Univ. Florida Press, Biol.
Sci. ser., vol. 4, no. 4, pp. 1-267, 88 figs., 24 plates, 19 maps.

—. 1955. The southeastern species of Baetisca (Ephemeroptera: Baeti-
scidae). Quart. Jour. Florida Acad. Sci., vol. 18, no. 1, pp. 1-19, 23 figs.

. 1958. Mayflies from the lower Apalachicola river drainage. Quart.
Jour. Florida Acad. Sci., vol. 21, no. 1, pp. 25-3], 1 fis.

EpMuNDbs, GEORGE F., JR., AND JAY R. TRAvER. 1959. The classification of
the Ephemeroptera, I. Ephemeroidea: Behningiidae. Ann. Ent. Soc.
America, vol. 52, no. 1, pp. 43-51, 32 figs.

Bureau of Sanitary Engineering, Florida State Board of Health,
Pensacola, Florida. Present address: U. S. Department of the In-
terior, Federal Water Pollution Control Administration, Southeast
Water Laboratory, Athens, Georgia.

Quart. Jour. Florida Acad. Sci. 29(3) 1966( 1967).

Caribbean Recruitment of Florida’s Spiny Lobster Population
Haroip W. Sims, JR., AND RoBert M. INGLE

Tue Florida spiny lobster, Panulirus argus (Latreille) is an animal
of considerable importance to Florida fisheries. The principal
fishery is centered in the Florida Keys and southeastern Florida,
but it ranges throughout the Gulf of Mexico and along the east
coast of the United States to Beaufort, North Carolina (Moore,
1962).

Previous reports have been published on various phases of the
biology of this animal. Crawford (1922) wrote on spawning
habits and artificial hatching. Crawford and De Smidt (1922)
covered life history, utilization, and the findings of a spiny lobster
hatchery formerly operated by the U. S. Fish and Wildlife Service
at Key West. Smith (1948a, 1958) dealt with the fisheries generally
and made notes on life history, methods of capture, markets, and
legal regulations. Dawson and Idyll (1951) reported on conserva-
tion methods. Lewis (1951) described the phyllosoma larvae and
was first to mention that foreign recruitment may take place. Later
Lewis, Moore, and Babis (1952) reported on the post-larval stages.
Robinson and Dimitriou (1963) published on the present and past
status of the fishery. Ingle et al. (1963) proposed a possible Carib-
bean origin of Florida’s spiny lobster populations and were the first
to emphasize the importance of the Yucatan Straits in such recruit-
ment. Several mimeographed reports have been prepared by the
University of Miami on the production of marketable lobsters. The
present summary provides an extension of the work reported by
Ingle and co-workers.

MeETHODS AND MATERIALS

Between August 1962 and August 1963 eight trips were made
through Florida Straits to the Yucatan Straits for the purpose of
collecting plankton. Samples were taken during the day and night
when work was underway. Some cruises and sampling were termi-
nated before reaching the Yucatan Straits, as weather and rental
boat fees governed the number of days work on each cruise. During
the eight trips 413 samples were taken and a total of 6931 phyllo-
soma larvae belonging to the genus Panulirus were collected.

Standard tows were made at random with a conical California-

208 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 1

Vertical distribution and diurnal migration of phyllosoma
Cruise number 7, June 1963. Number collected by each net by day or night

Station Day Night
Number *300/s Surf. *300/s Surf.

I 5 0

2 14 0

3 liat 0

4 0 25
5 ili 1

6 iS 0

7 5 1

8 0 0

9 1 0

10 1 62
It 5 80
112 24 245
13 4 21
14 Trawl Sample

15 35 4

16 14 0

1G 14 0

18 4] 0

19 36 5
20 24 153
21 28 162
2D, It 4
DS} 0 0
24 Ih 0

25 0 0

26 5 ol
DY i! 272,
28 Try Net

29 Try Net

30 1 0

*Oblique tow 300 foot to surface.

Vertical distribution and diurnal migration of phyllosoma

SiMs AND INGLE: Spiny Lobster Recruitment

TABLE 1 (Continued )

209

Cruise number 7, June 1963. Number collected by each net by day or night

Station
Number

31
32
33
34
35
36
37
38
39
40
41
42
43
44
A5
46
47
48

*300/s

22

D®|W WW

Day

Surf.

oS = Ot

*Oblique tow 300 foot to surface.

Night
*300/s

Surf.

14

107

1a

210 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

type plankton net with a diameter of one meter at the mouth and
about 15 meters in length. The main body of the net was con-
structed or number 30 grit nylon gauze with a finer section of
number 56 grit nylon gauze at the cod end. It was lowered into
the water to a depth of 150 or 300 feet and was then brought to
the surface over an oblique tow which brought it from scheduled
depth to surface in 30 minutes. At the same time another net of the
same type, or of 2 meter in diameter in some cases, was towed just
below the surface. Phyllosoma larvae were collected in both nets.
Samples were preserved immediately in 5 per cent formaldehyde.
Later they were washed, then stored in 3-5 per cent formaldehyde
buffered with borax and marble chips.

In the laboratory all phyllosoma larvae were removed from
plankton collected in each sample. Various known and unknown
species were separated and the number of larvae for each species
recorded. Total length, measured from the apex of the angle
formed by the eye stalks to the tip of the telson, and the possible
stage of development were recorded. Tables 2-6 deal with the
phyllosoma larvae of only Panulirus.

THE PHYLLOSOMA LARVAE

The phyllosoma larvae of the Palinuridae (typical spiny
lobsters) and the Scyllarides (sand or shovel-nosed lobsters) are
leaf-like and transparent plankton.

The colloquial name stems from Leach (1816), who gave such
larvae the generic name Phyllosoma. Other workers notably Guerin
(1830), Richters (1873), and Ortmann (1893), carried on the name
in the belief that these forms were adult. White (1847) gave the
adult form of a spiny lobster the generic name Panulirus. It was
not until the early 1900’s that many zoologists simultaneously dis-
covered that the phyllosoma were the larval stages of spiny lobsters.
Although Phyllosoma has priority, it has been used exclusively for
larvae, and the International Commission on Zoological Nomen-
clature recently suppressed Phyllosoma in favor of Panulirus (Von
Bonde, 1932).

The long larval life of phyllosoma is well documented by Smith
(1948b), Sheard (1949), Lewis (1951), Johnson (1960a), and
many others. The wide geographic dispersal, via ocean currents,
is shown by Gurney (1936), Smith (1948a), and Thorson (1961).

Sims AND INGLE: Spiny Lobster Recruitment 211

Because P. argus is the commonest spiny lobster in the area of
this study as well as in the Caribbean, the area of possible recruit-
ment, it is assumed that the majority of phyllosoma reported upon
here belong to that species. Probably some are of the species
P. guttatus (Latreille) and P. laevicauda (Latreille ), both of which
are reported in scattered populations throughout the areas of study
by Holthuis and Zaneveld (1958), Holthuis (1946), and Smith
(1948b). There is no published literature describing laboratory-
reared phyllosoma of P. argus, P. guttatus, or P. laevicauda, and no
one has ventured to describe the larvae of P. guttatus or P. laevi-
cauda from planktonic collections. Lewis (op. cit.) described from
plankton 11 stages in the larval development of P. argus. Although
our data are not in complete agreement with those of Lewis, we
provisionally accept his designations. It appears there could be
intermediates between many of the stages Lewis describes, but
little more than conjecture could be added without laboratory
raised material.

For 300 Panulirus phyllosomas selected at random, an attempt
was made to separate the three species meristically, by comparing
such body parts as fore-body width to total length and fore-body
width to hind-body width. These ratios failed to show any specific
differences. To further check this method the same Panulirus
phyllosomas were plotted against an equal number of phyllosomas
of Parribacus, of the family Scyllaridae. Phyllosoma of these two
genera are similar in body shape, but the two are easily separable
on morphological characteristics. The graphs failed to show a
difference between the two genera until the very late stages, and
then only because Parribacus grows much larger as a phyllosoma.

From work of Saisho (1964), who compared the first stages of
several species of Panulirus, and from our own observations on
Parribacus (Sims, 1965), it does not appear to be possible at present
to distinguish between the phyllosomas of closely related species.

SURFACE CURRENTS AND LARVAL MIGRATION

Ocean currents affect the distribution of food for fish, they warm
or cool areas of the sea so that certain species of marine life might
dwell there, and they aid in the distribution and transmigration of
the eggs and larvae of these species. Consequently, a knowledge
of currents is essential for the understanding of the variations in
abundance of marine animals that depend on them.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

212

Wea
‘O'1
‘WN
Wa
WH
Ou
‘Ou
‘Ou

aseud
uoo|

G8E

ies

‘BAY

066

8G

0'SG

0'SG

9°GG

9°8¢

0'6¢

L6G

‘duo
‘BAY

09

OOT

IVAICT

YIM

}U99 Og

eV

OL

sopdures
Joaquin

ST 9 SLI
LT I st'0
= L 9 ET
ol = ae
O'S S LL
IZIS 98R1$ UOleIAIg
[PPOW =- [Ppoyw urd

G

9341S
uvdI\

N 066-0G 29pnqe] ‘J vory 10; vzep ATYQUOT,,

G6 WIAV.L

Ctes I

So ee Il

CG S8I-GG

Gauls

Oc Ss I

osury
ANS

66L

paroy[op
JoquinN

t96l
‘Sny
C961
ounf{
ef61
‘aidy
C961
Ue [|
cS61
‘09q
cS6L
0
6961
‘ydas
C961
‘SnY

97eq

SIMs AND INGLE: Spiny Lobster Recruitment 213

Published studies on the surface currents of the Caribbean Sea
and Gulf of Mexico (Sverdrup et al., 1949; Leipper, 1954; Stommel,
1958; Drummond and Austin, 1958; Stewart, 1962; Ichiye, 1962;
Day, 1961; Salsman and Tolbert, 1963; Tapenes, 1963; Wust, 1964 )
indicate that the currents of the Caribbean, via the Yucatan Straits,
may be of importance in seeding Florida and the Gulf of Mexico
with tropical marine life. These wind driven currents of the
Caribbean, after going through numerous local eddies and gyres
along shorelines, flow into the narrow straits between Cuba and
Yucatan. After leaving the Yucatan Straits the water fans out into
various parts of the Gulf of Mexico. Water on the eastern edge of
the straits turns to the right to enter the Florida Straits and Gulf
Stream, water mid-way in the straits flows northward into the Gulf
of Mexico where it (1) enters the so called “loop current” (Salsman
and Tolbert, 1963; Univ. of Miami, 1957), which flows first north-
ward toward the United States coastline, then turns south and
enters a clockwise eddy flowing along the continental shelf of
Florida’s west coast; or (2) after a northward flow turns to the left
and enters a counterclockwise eddy that flows along the coasts of
Mississippi, Louisiana, and Texas. Water on the western side of
the straits flows into the southwestern Gulf and Campeche area.
There is an indication that some of this water leaves the Gulf by
subsurface and counter currents (Leipper, 1954; Salsman and
Tolbert, 1963). Leipper (1954) gives a generalized current pat-
tern of the surface currents of the Gulf of Mexico and Yucatan
Straits.

There is biological evidence that tropical Caribbean fish and
invertebrates are carried to Florida by these currents (Caldwell
and Briggs, 1957; Briggs, 1958; Caldwell, 1959; and Dawson and
Idyll, 1951). Cuppy (1917) and Arendt (1963) show that ocean
currents transport large numbers of “sea beans” from the Caribbean
to the coasts of the Gulf of Mexico and east Florida.

With this oceanographic and biological evidence in mind we
initiated a drift-bottle study which would help establish the rate of
flow of the surface currents in which the phyllosoma probably
travel. The data discussed herein cover work during the spring
and summer of 1963, with notes on a small drop made in the fall
of 1962. |

The release of drift-bottles was made in conjunction with our
plankton sampling cruises. Bottles of various types were dropped

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

214

e961

Wa = 066 66 vi GG L Lv L SoG L ect ‘sny
e961
‘O'T O'8E T'86 OL nS LT 9 74 9 EO GSS CPrsl oun{
e961
‘WN O'8E O'LG SV 9S Lt I SVG i OIG FT 8ST ‘aidy
e961
‘WH VSE 96 19 61 GL 9 9GI 9 CSCO I61 ‘uel
C961
WH L8e VGC OF GE G8 L 9 OL L TS¢9 I v9T RKC
c961
Ou QLe £66 68 LG OV V LOT V GEG T LGV ‘PO
G96L
(Oul '8e 9°66 v9 O€ GS L U'81 L 096C I COV ‘ydog
G961
Ord 9 LE T0€ 98 vl get V el V 0'ST-9'T 9ET ‘ony
asveud [eS ‘dulay, ovaiey sojduirg 9ZIS a8e}S UOlVIADGd 98v}S asury pe,a][0D aed
uooj ‘SAY ‘SAY yaa roquinN [epoyy [epoy uvoy| Uva aZIS roquin Ny
U9 Jog

N oFG-BS OPMINL] “TT Vor 10F vyep ApyIUOYW
€ AIaVL

Sims AND INGLE: Spiny Lobster Recruitment 215

during cruises in the months of September and October, 1962, and
April, June, and August of 1963. Fig. 1 shows the approximate
location of each release. Table 7 gives a complete list of the re-
turns to date.

Drirt BotrLE RELEASE AND RETURN DATA

During the fall of 1962, 20 bottles were released in the Yucatan
Straits. One was recovered 18 days later at Snake Creek bridge in
the Florida Keys, a distance of more than 360 miles, with a mini-
mum rate of 20 miles a day. Another was found at Key Largo 119
days after release; as this bottle was found high on the beach it may
have been there for some time. A third bottle released toward the
western side of the Yucatan Straits, was recovered after 210 days at
Matagorda, Texas.

In April, 1963, 212 drift-bottles were released in groups of 6
along a course from Dry Tortugas to Alacran Reef and then south-

GULF OF MEXICO

KEY

YUCATAN ey sor APRIL @
oy -) JUNE O

4aw_ |AUG.
S OTHERS &

-O-

Fig. 1 Drift-bottle release locations (from U. S. Coast and Geodetic
Survey Chart No. 1007).

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

216

Wa
‘Onl
‘WN
Wu
We
‘Ou
Onl
Ou

soseyd
uooyj

got

eS
‘SAY

066 L8 Sill

6 86 LG IG
0:96 OS SI
TVG GV b
LG CV OG
L8G GL Lb

L 66 IV GI
GLE OOT V
‘dutoyt, ovaiwy = sopduirg
"SAY Uy Joquiny

JU99 19g

OV

91

Ll

IZIS

[EReN

e961

26 8°96 G C9IGLTI GLI “SnVy

C96I

I Die il Glee PSI oun{

C961

G GGL ¢ 0'0G-9'T C8 Idy

e961

L LST ib GOGGE JE ‘uv

cS6l

9 Gs Ail 9 6LI-S TI OF “09d

G96I

V S él V C6I-GT 6S PO

C961

I GGG I 0'61-S'T VS ‘ydos

C961

I ©0 T g¢o-C'T 661 ‘ony

98V1G UOlIAVqQ asvIS osury pepe][oD eae @|
[PPOW uvIIN uvoyN 9ZIS JoquinN

N «9G-¥G ‘III PO1V OF ep ATYJUOW

V ATAVL

Sims AND INGLE: Spiny Lobster Recruitment 21:7

west to the Yucatan Straits. At some stations soft drink bottles
(161) were released for the U. S. Fish and Wildlife Service Bio-
logical Laboratory, Galveston, Texas. It is interesting to note that
most of the bottles released between Dry Tortugas and 88°35’ west
longitude were recovered in the Florida Keys and on the Florida
east coast. Those released near the Tortugas showed more returns
in the Florida Keys than did bottles released west of there. The
one exception was station 20, whence one recovery was made at
Grand Isle, Louisiana, and another at Veracruz, Mexico. All the
bottles released west of 88°35’ W, were recovered from the coasts
of Texas and Louisiana. Bottles released along the north coast
of Yucatan and in the western Yucatan Straits were found in
Florida, the Bahama Islands, Texas, and Louisiana. According to
current charts all these bottles should have stranded in the north-
western Gulf. Observed returns can only be explained by the
existence of uncharted eddies or by the effect of local wind and
tidal conditions after the initial dispersal. The effect of wind on
drift-bottles is shown by Chew et al. (1962), who reported on the
abnormalities in returns of drift-bottles released in the wake of
hurricane Carla. The general effect of wind on surface and sub-
surface currents is discussed by Sverdrup et al. and by Leipper.
The fact that no recovery was made from station 32 in less than
92 days suggests that the bottles may have spent some time travel-
ing around before being caught in a current that carried them
ashore. To date 24 per cent of the bottles released in April, 1963,
have been recovered.

In June 1963, 434 of our own drift-bottles and 135 bottles pro-
vided by the Fish and Wildlife Service were released, mostly in or
slightly south of the Yucatan Straits. The quickest return from
this area was 65 days, and the mean number of days for all re-
coveries was 120. This would suggest that the bottles may have
been retained for it was shown by the fall release that such a trip
could be made in as few as 18 days. Again, bottles released on the
western side of the Yucatan Straits were recovered from Jackson-
ville, Florida, to southwestern Texas. To date returns have reached
23 per cent of those released.

Only 192 bottles were dropped in August, 1963. They were
released at four sites in and just north of the Yucatan Straits. Only
two stations have yielded returns. From Station A2 recoveries were
made on the Florida east coast and at Port O'Connor, Texas. From

AUG. 1962 SEPT. 1962
N= 799 N= 0

OCT....1962 DEC. -<I9'62
N =9 N = Il

JAN. 1963 APRIL 1963
N =0 N = 1001

JUNE 1963
N = 716

|
STAGE

Fig. 2. Monthly occurrence of larval stages in Area I.

Sims AND INGLE: Spiny Lobster Recruitment 219

Station A4 returns were made from the east coast of Florida only.
The transport appeared to be much faster at the time of this re-
lease. One bottle reached Delray Beach, Florida, in 20 days, a rate
of not less than 30 miles a day. The mean rate for all returns was
50 days or about 12 miles a day. We have recovered only 7 per cent
of those released.

PosstIBLE INTERPRETATIONS OF DRIFT-BOTTLE DATA

Data thus far developed from this study indicate that the rate
of flow of the surface current from the Yucatan Straits to Florida
is sometimes, at least, as fast as 30 miles a day. Such rapid trans-
port could have important bearing on the recruitment of phyllosoma
larvae. Lewis (1951) feels that the length of the planktonic life
of P. argus is more than 6 months. Other workers (Gurney, 1936;
Johnson, 1960a; Prasad and Tampi, 1959) predict similar lengths
of time for other species. All estimate that 9-13 stages develop dur-
ing this interval. Saisho (1962) was able to raise the larvae of
P. japonicus to the eleventh phyllosoma stage in 90 days. How-
ever, his stage XI is equal in size and development to stage V as
described by Lewis. Saisho interprets this to mean that there must
be many more stages in the development of the larvae than past
studies have indicated. The opinion shared by Saisho (1962),
George and Cawthorn (1963), and Inoue and Nonaka (1963) is
that the length of time between ecdyses varies with age and be-
comes increasingly longer as the phyllosoma mature. Saisho’s work
on P. japonicus showed that the time between stage I and II aver-
aged 7.4 days, while the time between stage X and XI averaged
13 days. From these data we may assume that the length of time
from stage I to the last stage in P. japonicus must be more than 6
months and that the total number of stages must be more than 11.
The same is likely true of P. argus.

When applied to our drift-bottle data, these findings have in-
teresting cannotations. For example, the data shown by drift-
bottle No. 77A, which traveled from the Yucatan Straits to Snake
Creek bridge in the Florida Keys in 18 days, suggest that even at
this rate of transmigration any first stage phyllosoma hatched in the
Yucatan Straits undergoes two ecdyses before reaching Florida.
Therefore, any first stage phyllosoma larvae found in Florida waters
must be the spawn of local populations. Deductions of this type

AUG.

OCT.

JAN. 1963
N = 191

JUNE 1963
N = 1843

SEPT. 21962
403

DEC.

APRIL 1963
N = 158

AUG.

Fig. 3. Monthly occurrence of larval stages in Area II.

Sims AND INGLE: Spiny Lobster Recruitment 221

should be of value in locating offshore populations. It is possible
that older stages could pass from the Caribbean to Florida without
ecdyses; however, it may be deduced from the drift-bottle study
that some of the objects carried by the surface currents of this area
may spend some time in the eddies and slow moving currents that
retard their northward movement.

It is possible that larval development is accelerated in the warm
tropical waters of the Caribbean and lower Gulf of Mexico. If so
the distance from the source of recruited larvae to Florida would
be shorter than the distance in higher latitudes.

VERTICAL DISTRIBUTION AND DIURNAL MIGRATIONS

A phyllosoma larva is not a static object like a drift-bottle. AI-
though planktonic in nature and at the mercy of prevailing cur-
rents, the larvae may inadvertently retard their northward move-
ment by undergoing changes in vertical distribution. Harada
(1957), George and Cawthorn (1963) and Johnson (1960a) sug-
gested that the phyllosoma larvae follow the typical daily vertical
migration of zooplankton (Moore, 1958). Our observations both
in the laboratory and at sea agree with these data. In tanks at the
Pigeon Key Field Station, early stage phyllosoma swam the length
of a 12 foot tank to congregate in the dim light surrounding a flash-
light beam. If the beam was moved to the other end of the tank
all the larvae followed within 5-10 minutes.

Table 1 lists the data collected during cruise number 7, made
in June 1963. During this cruise two one-meter nets were fished
simultaneously at each station in an attempt to show a variation
in the catch of each net during night and day. One net was fished
just below the surface and well behind the boat, to reduce the
chance of catching sargassum weed, which was abundant in the
area. Every attempt was made to keep the top of the net not
deeper than one foot. The second net was fished in the oblique
manner, 300 feet to surface in 30 minutes. Both nets were fished
for the same length of time. The deep net was brought out of the
water as soon as it reached the surface to prevent it from taking
surface plankton, as closing devices were not available. The data
clearly show that the largest number of phyllosoma were collected
by the deep net during day sampling, 7 Am to 7 pm, and by the
surface net during the night. Almost no larvae were taken on the

AUG. 1962
l29

OCT. 1962

VO JUNE 1963
I54

l23456789 10123456789 10II
STAGE

Fig. 4. Monthly occurrence of larval stages in Area III.

Sms AND INGLE: Spiny Lobster Recruitment 223

surface during day samples, and the few larvae that were collected
by the deep net at night may have been inadvertently captured in
surface water as the net was lowered and raised.

Differences in direction between surface and subsurface waters
are not uncommon and are frequently described in oceanographic
reports. These differences in current direction coupled with the
vertical distribution and migration of phyllosoma larvae could aid
in the retention of phyllosoma in a given area and would augment
chances for some local recruitment in Florida. This must remain
a speculation, however, until supporting evidence is developed.

SEASON OF HATCHING AND OCEANOGRAPHIC DISTRIBUTION

The first phyllosoma stage was collected every cruise except
the one made in January when the youngest stage was stage II.
The last stage (XI) was taken every trip except April. First stage
post-larvae were collected only in April, June, and August. In no
cruise did the range fail to extend from Stages II to X.

Such data suggest that hatching takes place throughout the year
at various points in and below the area of study. Supporting evi-
dence is given by Smith (1948b) and Holthuis and Zaneveld
(1958), who found gravid female spiny lobsters in the Caribbean
from October to June, and by Lewis (1951), who reported them in
Florida from March to November.

In an attempt to show a progression in the modal sizes of
phyllosoma transmigrating from the Caribbean to Florida and from
Florida north, four sampling areas were set up; Area I, from lati-
tude 20-22° N, including the Yucatan Straits and the waters north
of the Yucatan Peninsula; Area II, between latitude 22-24° N, in-
cluding the waters west of Cayo Arenas and east to Alligator Reef
in the Florida Keys; Area II], the waters between latitude 24-26° N;
and Area IV, a straight line due east of St. Augustine, Florida ex-
tending to the middle of the Gulf Stream. Area IV was sampled
only during the months of July and August, 1962. Tables 2-4 give
the data collected in each of the areas. Table 5 combines Areas
I-I1]. Table 6 provides information obtained in the Gulf Stream
east of St. Augustine. Monthly size frequency graphs are provided
for each area (Figs. 2-5).

These tables and figures show no modal progression in size or
stage between areas or from month to month. Geographically the

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

224

Wa a

‘Om GLe
‘WN e Le
Wa st
Wa gst
‘Ou L’Le
Ol 6 LE
‘Ou L'LE
poetic IGS
uoo| ‘RAV

066

6 8G

SENG

6 VG

OG

8°86

€'66

T0€

‘duo J,

‘SAV

06

OL

|S,

LS

OV

6L

LG

€6

OVAIV'T
YIM
yu99 10g

66 O'S iS
HTL Lit 9
LL LT I
9G GL 9
9S G8 L
6 0'V V
GV GG L
86 O'S Si
sopduins IZIS 93R4S
ETON MEX AT LE ROA

V6I S| 6 9G T
est 9 Cc aGsc: [
S09 Il O1ISVT
LGl 9 CGGEG
6GI L ISSiGecal
GI V GEGoG I
L0G L 0'9¢6-o'1
/é; \U It c OSGeaiT
UOHRIADG, 98v)5 OSU
uevsyN ueeWN 9Z1§

CoS

CTL]

VVGI

861

SOG

GGG

LGV

V9OOL

pepos][on
roquin Ny

C961
‘ony
C961
oun[
C961
‘ady
C961

‘ur {
C961
‘0d
C961
0
CS6l
"ydos
CS6I

‘ony

ay @|

poyesrodi0out III-[ S¥vety 10F vyep A[YWUOYY

G WIAaVL

Sims AND INGLE: Spiny Lobster Recruitment 229

larvae are spread almost uniformly throughout the areas sampled
except for the area in the Yucatan Straits, where the narrow channel
and less surface area probably account for a concentration of the
larvae and more are taken per sample.

A small number of early stage phyllosoma larvae belonging to
the genus Panulirus have been collected in the Gulf of Mexico west
of St. Petersburg, Florida.

of AREA IV

JULY (8, 1962 JULY 23, 1962
32 Zoi:

JULY 27, 1962 JULY 3l, 1962
28

r2345678 90 12 34567 8B9ION
STAGE

Fig. 5. Monthly occurrence of larval stages in Area IV.

HyDROGRAPHY AND DISTRIBUTION OF LARVAE

Studies made at the Pigeon Key Field Station indicate that
hatching takes place on offshore reefs on the ocean side of highway
U.S. 1. Samples of the plankton taken over these reefs during the
spawning period yielded numerous first stage phyllosoma larvae.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

226

i 8E m5 G VST G 8 cls TI
Il 6 CG G 0 G CG

Sl 8 aa = 0 oe CCST
oT 9 O€ € Sie € 0'9-0'G
OS GI aaa 9°¢ G9 9°¢ SETOE
ce at eG G TV G 03:9 1

AVAIL] YIM U01R1S IZIS 98S UOLIAVG 98S 9ZIS

}U90 Jog = Joquinyy [PPoW [RPOW uvoW uvdW asuryy

VpLO[y ‘ouNsnsny 3S JO ysvo onp “AT vory OF VIP ATYWUOT

9 HWIAVL

por][0D
roquinN|

ZOG6L ‘A[nf
GIGI “CS “SsNV
Z9GI ‘TE A4[n[
ZOGBI “LS Ap
ZOGL “CS ATOL
ZOGI ‘ST Aqnf

07°

Simms AND INGLE: Spiny Lobster Recruitment 2277.

In samples taken farther offshore the percentage of stage I lessened
and a wide size range was collected. From the bridges along U. S.
1 in the Florida Keys, 1237 samples produced only 7 phyllosoma
larvae, viz.: three stage I collected in May 1963, and single speci-
mens of stages I, III, VI, and VIII collected in August 1963. Prasad
and Tampi (1957) and Johnson (1960b) reported similar findings
and feel that there is a rapid offshore movement of the first stage.
If this is the case in the Florida Keys, then this stage must enter
the Gulf Stream to be carried toward northern waters. Some of
these larvae may be able to re-enter coastal populations by being
caught in one of the numerous eddies that lie along the eastern
coast of the United States and in the Bahamas. Great numbers
must drift northward with the Gulf Stream to oblivion in the North
Atlantic.

Twenty-six per cent of the phyllosomas collected during the 8
cruises were in the first stage. The largest single catch of this stage
was made in April, 1963, when 712 were collected in a 30 minute
sample taken just northeast of Cabo Catoche, Yucatan. All the
other samples containing large numbers of stage I were collected
within 50 miles of land, but it was not uncommon to collect this
stage more than 100 miles offshore.

The middle stages, IV through VII, accounted for the greatest
number of phyllosomas collected. These were collected every
month and in every area sampled indicating a good mixing of larval
populations and a constant recruitment.

The last stage accounted for only one-half of one per cent of
the total number collected. Small numbers were collected both in-
shore and offshore and on the surface as well as in the deep water.

Post-larvae were collected in the offshore waters of the Florida
and Yucatan Straits during the months of April, June, and August
and were collected in the inshore waters of the Florida Keys from
April to August. Large numbers of post-larvae were reported in
Miami from January to March (Lewis et al., 1952), and they occur
throughout the year in the Indian River at Stuart, Florida (Witham
et al., 1965). The post-larvae apparently may enter the inshore
waters to undergo juvenile development, but their occurrence both
inshore and offshore suggests metamorphosis takes place regardless
of the suitability of location for survival.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

228

‘Vp ‘Tuelyy “ND spurs fo *g ‘Aoy JON[A Ipls “yf 60T €9/T1/8 rE'FS ZO'FS 69/%6/P II LY

Lplo,y ‘yoveg rummy 2.9 ¢9/o¢/9 9 #8 60'FZ 69/¥0/¥ Or GP

VplOL | ‘Osiv'] Aoy “pus ‘N jo “gy ‘Aoy ysyposuy = s_LTE r9/9/¢ 8c'Eg SLPS 69/¢3/F 6 9¢

epHopy ‘os1e'y] Aoy ‘puo “N jo “y ‘AoY ysyposuy TTT ¢9/c/6 goes CL FS 69/¥3/P 6 cE

eplopy ‘Tuer ‘(pue "N) oy sopoyy PIO FS ¢9/8t/¢ 8o'eg SLPS 69/73/P 6 re

VPHOP A ‘SeGVD [RIOD —s- g@G c9/oe/L 8c'Eg ETFS 69/ea/¥ 6 ras

VPHOLy ‘Seqrg [RIoy —s- Fg 69/13/9 SEES ISPs 69/P6/F¥ 8 0g

Vplop | ‘Aoy UoNrzur[_ opis wey ZB 69/ct/2L SEER ISPS 6o/¢o/¥ 8 6Z

VplOLy “eprourr[s] “ospoy nqyrw QT 69/¢/¢ SEES ISPS 69/P3/F 8 GS

VpHOpy ‘shoy epHopy ‘Aoy AFTPuIAA ~—> QE 79/61/¢ ZTE 6S FZ 69/¢o/F i 6

VpHopy ‘oyepropney] 34 ‘yovrog epef gz c9/Pr1/¢ ET'ES 62 FS 69/P6/F ih IZ

VpHOpy ‘sAoy VpHopy ‘Aoy voryd voog 6P 69/61/9 ETE 6ST 69/¥6/F¥ L 03

VpHOpy, ‘sesnqyoy AI QT 69/6/01 00°8 SEES 69/¢3/¥ 9 LI

VPHOPA YU] OIG We 0% c9/et/¢ 00°E8 SES 69/¢3/F¥ 9 QT

VpHo,y ‘yoreg AvypoqT gy ¢9/11/¢ 00°8 SEES 69/¢3/¥ 9 al

epuopy ‘vuryuey — pT 69/¢3/¢ 00°E8 SEPT 69/¢3/F 9 eT

shoy eplo[y ‘Aoy uonrzwr[q 6% ¢9/0z/¢ OF'S8 LEV 69/es/F G Tl

epuo,y “uoyeY voog gy] ¢9/6/¢ OF'S8 LEFT 69/¢3/F G 8
suxo], “eplosryw JO wey OIG c9/¥/¢ 90°98 GOSS 29/6/01 6h OV@

VpHOpy ‘OBiey] Avy ‘Joo urovQ ~~ TT 69/¥/1 00°98 00'°Sz 69/9/6 SF V6L

SUE OAL CI/EGGT SEM) EMIALS TO UEMONC Fa) zo/ez/6 = 0098 — ss o's_—ié 90/6 8h VLL
[PASLQey Jo vory posdeyq [PASLQOY ‘M ‘SUOT ‘N77 asvaloy uoeyg sI9quinNy
SAC fo OVEC SUOT}LOO'T ISvI]OY fo 9}EC LOG UUTL NY 9[HOg

vyep UINjJoL 9]}0q-JUIG

L ATEaV.L

229

SrMs AND INGLE: Spiny Lobster Recruitment

sexo], “epnsulueg IBAlog

‘eT ‘UOTOWUeD

SVX], “UO}JSOATLS)

vplo,y “yorog vuoyAreq

BpHo[y “Yoveg surmmoqyeayy ‘sesioutey iepey
eplopy Tuepy ‘Avg oudvostg

BpHO[Y “seA AVY “QnlD 10y4FO IO[AR], “77
eplo,y ‘yovog ounsnsny ‘3S

VPLIO[ Y “IOLUIOARC I],

eploly ‘eletg 3 “Yd Joddag ‘KN soprut ¢
‘XOW “ZNID BIOA JO Avg ‘eSal[ey vjop ofloo1y
‘BT “‘o[sy puri

Bpholy “Yorog tue ‘yrwg ounojnepy
Bplloyy “Yoveg O10A

vpHo[y qURID Iesu “eplo[y “epuy ueyseqos
epllopy “‘yorog ourduog

eBpllo[y “oyepiopney] 147

epllo,y “yYorog r1uieryy

VpHO[y “YWOAA oye] ‘Yreg sddiyg

eplo,y “Yovog uyeg

‘eplo[ yy ‘episuvs0Q ‘Os1e7T Aoy YON

6E1
Sel
cel
Tél
Lol
961
OGI
6IT
VIT

eS eR eI Lie ee wg eS ee = tee at a LSM NES meee BME a eae ee
Toquun Ny Jaquin Ny

[PASI Y fo volV

aa ¢9/6/9 6168 ese ¢9/9¢/F
IS e9/91/9 00°68 63'S 69/96/F¥
Ch ¢9/o01/9 00°68 6S'SS 69/96/F
OST ¢9/¢3/6 CF'88 LESS 69/¢a/F
61Z c9/6I/IL SF'88 1€'BS 69/cz/¥F
III 69/4/6 9T'98 GPSS 69/¢3/F
Z6 69/L3/L 91°88 IFS 69/9¢/F
9C1 ¢9/6¢/6 91°88 IF 3S 69/96/¥
FOI ¢9/38/8 C388 FESS ¢9/90/¥
LOT 69/6/01 CTL8 6S'SS 69/¢6/F
60 79/836/% cG'98 OGG 69/¢3/F
LOT ¢9/o1/8 GC'98 LOSS 69/¢3/¥
LG 69/%3/¢ 9€°98 ST'eZ 69/¢a/¥
ce c9/oe/s 9E°98 ST's 69/¢o/F
Le ¢9/1/9 9¢°98 ST'Es ¢9/¢¢/F
SF ¢9/s1/9 L198 IGG 69/ca/¥
08 69/FI/L LT'98 TAGS 69/c0/F
Z9 ¢9/9¢/9 L198 ISS 69/ca/¥
Ig c9/et/L GE's CEES 69/c¢/F
€8 c9/91/L cE'cg Geez 69/ca/¥F
cg ¢9/66/9 8Z'S8 NES 69/¢c/¥
poesdeyy [PAPI TOY AYN ‘SsuOT “N ‘yey 9SeI[OY
sAeq fo OE C] SUOT}890'T oSeI[OY fo 91°C

uoHe}S

ap0g

SS ]SSSSS—SSS———————LL_
eyep UIMNJor 2[H0g-YNG

(709) 2 ATAV.L

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

230

vpnsulueg IwATog ‘sexo yp, “yovog [vysAip
eplo,y ‘yovog [e1oAvury odey

sexo, ‘TYstyD sndioy ‘purysy oipeg
eplopy ‘Tey Ywqg wopuriy

vplop.y “yovog O10,

Oolxeyy ‘sedipneure y,

vpllop yy “UozyeYy voog

Sexo], “YsiyD sndio+y Jyo “purysy orpeg
vplHOLY “yovog rueiyy

eplopy ‘oulsnsny "yg ‘Yovog ouvyiA
VplOp J ‘Osaie'T Aoy JO apis urv0Q

‘VT “ssvq AYSIU AA

Sexo], “UO}JSOATLS) ‘e[NsUIUOg IwATOg
sveulvyeg “Avd Ie A\-O-uvyyy

vUvISINO'T “puvy]sy NoT[IVD

sexo], “epnsuruog eproseyeyy ‘nodvg s uoatg
sexo], ‘yorog JuosINg

Sexo], “Vpslosrjzryy

sexo], ‘odooiy “yovog optsying

Sexo], “JSLIYOTID

[PASLIOY «JO vory

GL
€6
LIT
8V
901
686
88
CGI
OS
16
6rI
SOT
ial
CVG
OTT
€L
v9
89
IV
6P

posdeyly
sAvq

69/3/6
69/6/6
69/ct/or
69/1/8
¢9/0z/9
r9/e/¥
¢9/91/6
¢9/¢3/01
69/8/8
¢9/61/6
¢9/¢c/6
¢9/1/6
¢9/06/6
69/16/61
£9/¢3/8
69/01/L
ec/1/L
69/P/L
e9/9/9
69/F1/9

[PAOLQOY
JOSE

00°L8
66 G8
6G G8
66 G8
6G G8
6G G8
GC G8
6G G8
66 G8
6G G8
80°98
80°98
80°98
80°98
80°98
To L8
1G L8
IG L8
10°88
61°68

SUOT}LVOO'T OSBOTOY

vyep UIMJoI 9[}I0G-71Iq

(*yU09) 2 ATAVL

"M ‘SUOT ‘N ‘1e7T

CE°0G ¢9/61/9
Ch'0Z ¢9/0z/9
CFO ¢9/02/9
CF'0G ¢9/06/9
CF'0G ¢9/0z/9
CF'0S ¢9/06/9
CFO 69/06/9
CF'0G ¢9/06/9
Ch'0G ¢9/06/9
CF'0G ¢9/06/9
Geil 69/66/F¥
Geng 69/66/F
GEIS ¢9/66/F¥
CEI 69/6¢/F
GETS 69/6¢/F
0'SS ¢9/83¢/F
0'SS 9¢/86/¥
70'S 69/83/F
BC'S ¢9/96/F¥
Ware 69/96/F

ISBIOY,

jo OCC]

8b
VV
VV
VV
VV
VV
VV
VV
vV
VV
CE
CE
GE
Ge
CE
[€
I€
ie
66
86

008
L6G
966
P66
68
98
V8G
08S
SLZ
LLG
60
S0G
GOS
IST
rol
S91
19T
6ST
6h
OFT

Joq Un NY TOQUIN NY

uolryg

aTnog

SIMs AND INGLE: Spiny Lobster Recruitment 231

SUMMARY

A large number of plankton samples were taken with | meter
and % meter plankton nets in the waters of the Yucatan and
Florida Straits. The absence of a progression of model size or stage
for phyllosoma larvae of Panulirus argus from southern to northern
stations or from month to month indicates that year around spawn-
ing takes place in the Caribbean. This assumption is supported by
the continuous presence of gravid females. The wide size range
throughout the period sampled is also consistent with widespread
recruitment from the Caribbean.

Drift-bottle data indicate that the rate of flow of the surface
water from the Yucatan Straits to Florida may be as fast as 30 miles
a day, but many floating objects take longer because of eddies and
slower currents. Moreover, in a laminated current system in which
contiguous layers move in opposing directions, horizontal move-
ments of vertically migrating plankton may be retarded. Phyllosoma
larvae undergo such vertical migration.

Successful migration of planktonic animals over long distances
is largely dependent upon length of larval life and speed of cur-
rents. Prevailing hydrographic conditions in the Yucatan and
Florida Straits give strong evidence of the seeding of Florida
waters with Caribbean hatched phyllosoma larvae. Information is
now needed to show how these larvae enter coastal populations
and what methods may be employed to retain greater numbers of
them to insure their survival and growth. The international scope
of current and future investigations emphasizes the need for co-
operative efforts throughout the Caribbean area. Even though
spawning in Florida is probably greatest in spring and summer,
the year around delivery of larvae from the Caribbean and con-
tinuous recruitment of newly formed post-larvae makes year classes
difficult to distinguish.

LITERATURE CITED
ARENDT, H. E., 1963. A wandering beachcomber returns. Florida Naturalist,
volmao; no, ll, pp. 5-7.

Briccs, J. C. 1958. A list of Florida fishes and their distribution. Bull.
Florida State Mus., vol. 2, no. 8, pp. 223-318.

CALDWELL, D. K., AND J. C. Briccs. 1957. Range extensions of western
North Atlantic fishes with notes on some soles of the genus Cymnachirus.
Bull. Florida State Mus., vol 2, no. 1, pp. 1-11.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

232

eplopy “yovog O10,

eployy ‘Turerpy FO YMoS

Vploly ‘yoveg vuojAeq

epHOL,y ‘ouljsnsny 4g Tou pURIsy visejseuy
eplOoLy ‘oulsnsny "3S “YoRog }Usdse17)
eplioyy “‘Yovog ounjdoyy ‘yovog epioA 9}U0d
SBVxOT, “UOJSOATLL) “PURIST URT]IG

sexo], ‘soyovog Jjng) 10uu07) OG WOg

VPHOL[Y ‘SAVY VPLOp[ “IOTUIOAR IT,

vuvIsIno'y ‘puvjs] diay, puri

Oxo SZNID) BIOA

epllo, yy “yorog Aviad

epHO[y “ootolg “Ww

epllo[y ‘ouAvosig Aoy

sexo], ‘siyD sndioy ‘purysy s1peg

vplopy ‘yovog a[fAsnqty, “odey os[ey

Sexo], “IOUUO0D OQ MOg ‘purys] eVpsosejieyyy
eplo,y “purysy Joydn{

SVx9T, “o][[IAsSUMOIg

Vpllo[y “Yovog evurAulg MON

[PASLQeY Jo vory

89

LIG
16

LOT
L8

IOL
GLG
eSI
IéI
8cI
GGG
<9

901
901
661
OGI
Sil
Sil
ILE
6G1

pesdely
sAvq

¢9/1z/8
79/e1/%
¢9/61/6
¢9/¢/01
c9/ct/6
¢9/83/6
r9/L1/¢
¢9/61/TI
69/81/01
69/43/01
79/P1/%
¢9/3/6
¢9/F/0r
¢9/¢/01
69/12/01
¢9/st/ol
69/Z1/T1
¢9/91/01
79/c6/¥
¢9/cz/0r

[PASLQeY
jo o1eq

ce'gs
cess
cess
cecs
cess
99°98
gc'98
99°98
99°98
99°98
c¢c'98
gc'98
€1S8
cl S8
€TSs8
€1 Ss
el Ss
€LSs
€TSs8
€1S8

OV 0G
OV 0G
OV 0G
OV 0G
OV 0G
0£ 0G
GE 0G
O€' 0G
Of 0G
O€'0G
GE 0G
GE 0G
LV 06
LV'0G
LV'0G
LV'06
LV 06
LV 06
LV'06
LV 0G

vyep UINjol 9[H0g-YuNCG

(709) 2 ATAVL

"M “SU0T “N 2eT
SUO}vIO'T osKa[oyY

¢9/06/9
¢9/02/9
¢9/06/9
¢9/02/9
69/02/9
¢9/61/9
¢9/61/9
¢9/61/9
¢9/61/9
¢9/61/9
¢9/61/9
¢9/61/9
¢9/06/9
¢9/06/9
¢9/0z/9
¢9/0¢/9
¢9/0¢/9
¢9/06/9
¢9/0¢/9
¢9/¢c/9

asvaloy
jo aeq

OLV
LOV
907
1lOV
66E
LEE
C6
16€
98
O8E
LLE
GLE
VLE
eLe
99
69E
O09
9S
6St
vot

IOQ UN NY IQ un NT

wONeIS

a]y0g

233

Sims AND INGLE: Spiny Lobster Recruitment

epllopy ‘sea, Aoy Jfo ‘sAoy Young s[ppes
VpHO[y “Osie'T Aoy JO apis uvs00 ‘purjsy elnf
eplo,y ‘yYovog poomAT[op{

eVplHOpy ‘sesnyioy, “Aoy uopresy

eployy “yoveg eurAuig MaN YyNOS

eplo[y ‘sesnyoy, “Aoy uopies

epho,y “yovog vuoyded

eVplHOp[Y ‘sesnjyloy, “Aoy Uwapres

BPO, “TUR

sexo], “‘puvysy oipeg

eplHO[y “so1oyg uoyyeIeyy ‘Yovog unt wooo
BpHopy “seM Aoy

eplHO[y “JseA, Avy “Yoveg sioyzeUIg

eplo[y “yovog oe[IAsnzy,

eplo,y “ounsnsny “3s

vplojy “qovog eucjyded

eplo,y ‘ounsnsny 34S

eplo[y “yowog Avijoqd

eplo,y ‘1oydn{

eplo,y “yovog uosuof

[RPAsINeY JO vory

IéI
col

66
801

pesdely
sAeq

69/8¢/TI
69/06/TI
¢9/6/6

69/1 /9
e9/91/01
e9/9t/9
69/8/01
e9/91/9
¢9/1/6

r9/61/F
c9/Pt/L
¢9/8/6

69/T1/L
¢9/96/6
69/06/11
¢9/11/01
¢9/13/6
¢9/96/6
¢9/8¢/6
¢9/2/01

[PAoInEY
jo a3eq

Loss
VI'98
VI'98
008
VI'98
00°€8
VI'98
00'€8
VI'98
OG ES
OG ES
OG ES
OG ES
90°S8
90°S8
90°S8
le Sie}
€T'S8
€TS8
GlLSs

eyep UIMJoI 9[}0q-YUG

(709) 2 ATAVL

GV 0G
CI'Gsé
CI'GG
SE VG
CI'GG
SE VG
CI'G%
8E° VG
STG
O€'9G
O€'9G
O€'9G
O€'9G
9V IG
9V 1G
9V' IG
IO'1G
IO'1G
I0'1G
1016

“AA ‘Bu0'T "N WeT
UOT}BOO'T VSvofOY

¢9/0/9
¢9/11/9
e9/11/9
¢9/#1/9
¢9/11/9
¢9/#1/9
e9/11/9
69/1 /9
69/11/9
69/¢ez/9
¢9/¢3/9
¢9/¢s/9
69/¢3/9
¢9/1¢/9
¢9/1¢/9
¢9/1¢/9
¢9/1¢/9
¢9/1¢/9
¢9/12/9
¢9/13/9

asealoy
jo a7eq

G6V
O67
887
L8V
987
C8V
O87
OLY
VLYV
697
89F
LOV
6S7
CVV
OV
9ET
CV
SGV
LGV
IGP

joquinyy JoquinNy

uoTeIS

amiog

VpLoLy, “yoreg puowsiQ TTT 69/8/01 GG'98 PE 0G ¢9/61/9 ce 9L¢
Vplopy “Yorog vuoyeq ZI ce9/LT/0L Sg'98 PE0G ¢9/61/9 ce CLS
VPHO[] “IOWIOART, gg ¢9/¢3/6 Gc'98 FE0G ¢9/61/9 ce TLS

VpHopy ‘Tuer ‘Aoy spurs gg 69/21/01 9Sci98 PE0Z ¢9/61/9 cE OLS
sexo], “IOUU0).Q WOd FZE 79/8/F¥ CC'98 PE'0S ¢9/61/9 cE 69¢
VplLop,] ‘Seloyg woe, [QT 69/6z/6 CZ'98 IF'0G ¢9/0¢/9 OF GZG
BPO, “Yowog puouug ETT c9/L1/0I S398 IF'0G ¢9/06/9 OF gs

epHop | ‘Aoy oqumooyeyy IOMOT 6g 69/13/6 GS'98 IF'06 ¢9/06/9 OF 6I¢
VpLOp | ‘SeBn}O], ‘Aoy uepiey | e9/st/9 00°E8 6E'FS 69/#1/9 9 Ig
epuopy ‘soy eplopy ‘Aoy NC —-goT 69/¢/01 LG°S8 SFOS ¢9/0¢/9 GP rats
epHOo[y “yowog uesuef gy c9/Tte/8 LG'S SFO ¢9/06/9 SP Ig

BPHO[y “Yovog BuUIAUIg MON GG ¢9/13/6 yigies SF'0G ¢9/06/9 rai org

sexo], “AsiyO sndioD ‘purjs] e1ped = gc eg/es/Il Lg'¢8 SF'0G ¢9/0¢/9 CP 60S
BqnD ‘euvaryy “Yorog vse1Q rood = gg €9/Ga/cl Lees SF'0G ¢9/06/9 GP LOS
BpHOoLy “yovog [eioavurD odeQ = egy €9/ Iie/ Ol ee Less CFO ¢9/0z/9 OF 90¢
VpHO[y “seroyg UoTIeIeIN gay eo/ez/1l _—sLg'e8 SP'0G ¢9/06/9 CP ery

VpHo[y “Yoreg Puousg JON 96 ¢9/9¢/6 GE"Sg SP'0G ¢9/06/9 OP GOS

epHoLy ‘sAoy epHopy “purysy eynf oz 79/8/¢ LG"S8 SPO ¢9/0¢/9 SP S6F
BpHopy ‘Shoy poly “Aoy uoneqwed goy 69/9/01 LG'S SF'0G ¢9/06/9 SF 96F
EPMO[Hy YOeod PuoUN® — OgT €9/S1/01 JG a8 SF'0G ¢9/06/9 OP G6F

[PASLIOY JO vory posdeq [PADIIOY INN SHOOT) ING Hf asvapoy = Joquinyy Joquinyy

sAvq jo a3eq SUOI]BOO'T 9Sva[oY jo o7eq uonrg os9no0g

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

—_

¢

234

Rep UMIoI 9[}}0q-lUG

(WU09) 2 ATAVL

SIMs AND INGLE: Spiny Lobster Recruitment. 235

CALDWELL, D. K. 1959. Observations on tropical marine fishes from the
northeastern Gulf of Mexico. Quart. Jour. Florida Acad. Sci., vol. 22,
no. l, pp. 69-74.

CuHew, F., K. L. DRENNAN, AND W. J. DEMoRAN. 1962. Drift-bottle return
in the wake of hurricane Carla, 1961. Jour. Geophys. Res., vol 67,
HOM MDD et1o-271 10,

Crawrorp, D. R. 1922. Spawning habits of the spiny lobster Panulirus
argus with notes on artificial hatching. Trans. Amer. Fish. Soc., vol.
50, no. 2, pp. 312-319.

CrawForp, D. R. ann W. J. J. DE Smipr. 1922. The spiny lobster
Panulirus argus of southern Florida: Its natural history and utilization.
Bull. U.S. Bur. Fish., vol. 38, pp. 281-310, fig. 260-273.

Dawson, C. E. anp C. P. Ipyty. 1951. Investigations of the Florida spiny
lobster Panulirus argus (Latreille). Florida Bd. Conserv., tech. ser., no.
2, pp. 1-39.

Day, C. G. 1961. A literature survey of hydrography, bathymetry and
fisheries of the Atlantic Ocean under the Atlantic Missile Range with
an appendix on the Mona Island region. Woods Hole Ocean. Inst.,
no. 61-36, pp. 1-114.

DrumMmMonp, K. H., anp G. B. AusTIN, Jr. 1958. Some aspects of the
physical oceanography of the Gulf of Mexico, physical, chemical data
from Alaska cruises. U. S. Fish and Wildl. Ser. Spec. Sci. Rept. Fish.,
no. 249, 417 pp.

GeorcE, R. W., AND P. CAwTrHOoRN. 1953. Investigations on the phyllosoma
larvae of the Western Australian crayfish. Report for 1962, pp. 1-9
(mimeo. ).

Guerin, F. C. 1830. Crustaces et arachnides. Voyage de la Coquille, 1822-
1825, vol 2, no. 2, pp. 9-319.

Guppy, H. B. 1917. Plants, seeds, and currents in the West Indies and
Azores. Williams and Norgate, London, 531 pp.

Gurney, R. 1936. Larvae of Decapoda Crustacea. Part III. Phyllosoma.
Discovery Rept., vol. 12, pp. 400-440.

Harapa, E. 1957. Ecological observations on the Japanese spiny lobster,
Panulirus japonicus (von Siebold), in its larval and adult life. Pub. Seto
Mar. Biol. Lab., vol. 6, no. 1, pp. 99-119.

Hoxttuuis, L. B. 1946. The decapod Macrura of the Snellius Expedition. I.
The Stenopodidae, Nephropsidae, Scyllaridae and Palinuridae. Biol.
Res. Snellius Exped. XIV, Temminckia, vol. 7, pp. 87-178.

Ho.ttruuts, L. B., AND J. S. ZANEVELD. 1958. De _ Kreeften van de Neder-
landse Antillen. Zoologische Bijragen, no. 3, 26 pp.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

236

eployy “yovog vuojAed

eplIo,y “ouljsnsny "4S

eplopy ‘yovog evurdulg MON

eployy ‘Asyory Og MeN

eployy ‘Aoyory Og MaN ‘Yoveg Aej-V-1O[
eplopy ‘Aoyory WOg Mon “Yovog Iey-V-IO[ 4
vplop yf ‘Aoyory WOg MON ‘yovog 1ourpy
vplo,y ‘Aoyory Wog Many ‘Yovoeg Iey-VY-1O[
eplto,y “Aeyory WOg MON ‘Ady UaeI5)
eplOopy “Weng ‘yoyuy stony “4s

BPO] “UOYeIK

epHO,y “soa Ady “Aoy UBUIO AA

VPlO[Y ‘919g “yf

seuleyeg “Ae ysturds

eplo,y “Aoy youq

sexo, “YsiyD sndioy ‘purjs— sipeg

VplOo[y “Yovog euIAUIG MON

sexo, “‘YsuyoD sndioy “purysy, o1peg

epHOoly ‘Yovog BVuIAUIg MON

eplopy “Yovog rureryy

Bpllo,y “aenys

[PAZIIJOY «JO voIy

OTT 69/LI/0T 90°S8
POT ¢9/¢/0T 90°S8
86 €9/13/6 90°S8
CT 69/8/L LG'S
CT 69/8/L LG'S
GT 69/8/L LG'S
19 €9/G6/8 LOSS
LG 69/06/L LG'SS8
GT €9/8/L LG'S
Negi 69/86/01 CPEs
IP 69/¢/8 Ol'es
PG 69/LI/L O1'E8
06 €9/61/6 OL'Es
CLG £9/06/€ OL'E8
6IT 69/06/01 OL'Es
6ST 69/¥#6/TT F198
OTT €9/G1/01 IF'98
ZOl 69/836/6 €S 98
60T ¢9/01/01 Gc'98
9II 69/61/01 Gc'98
16 69/¥6/6 GC'98
pesdely [BAIT TOY “M ‘SuO'T
SARC] jo 93eq

IV 1G
9V IG
9V IG
00° LG
00° LG
00° LG
00° LG
00° LG
00° LG
CV 9G
€V 9G
CV 9G
tV 9G
CV 9G
CV 9G
99°06
99°06
IV 06
VE OG
VE OG
VEOG

‘NO We]

SUOT}VOO'T ISvIOY

69/€636/9
€9/16/9
69/13/9
69/63/9
69/€63/9
69/€66/9
69/¢6/9
69/€63/9
69/66/9
€9/€6/9
69/€6/9
€9/€3/9
€9/16/9
€9/€6/9
€9/€6/9
69/81/9
69/81/9
€9/81/9
€9/61/9
€9/61/9
¢9/61/9

asvajoy
jo o1eq

>
wt

NNNANNANAA Pte eA eT e
Sag

ct
ct
Gt

Toq Un NT
UOT}

899
G99
€99
699
099
6S9
899
LG9
gc9
OS9
679
vVV9
cVv9
GVO
69
VEO
669
G19
68S
08S
8LS

joaquin
a[0g

vyep UINjol 9[}OG-UG

(7u09) 2 ATAV.L

~
iN

SiMs AND {NGLE: Spiny Lobster Recruitment

eplo,y ‘ounsnsny 3s

vVplHOLY ‘Yoveg Jo[sv[y pue puowidg UusesMjog
vplopy ‘yorog wyed SoA
epHopy “yovog 93z]Je}es
eplHo,y “qovog 9a][Auosyor[
VpHO,y “Yowog PvuIAUIG MON
Sexo], “purysy sipeg yWwos
eplo,y “yovog Jepsrpy

eplio, yy “yovag vooo0n

eplopy ‘yorog Aeijoqd

sexo], “Iouu0D OC WO “puvyjsy epsiosezeyy
eVplo[y “sAyoel[ soperps19Ayq W0g
epho[y “‘Yorog vurlAwig MoN
eplo[y ‘Yovog sumoqyeyfy
eplopy “yovog evo007

vpHo[y ‘[etoavuey odeg
sexo], ‘pueysy oped

sexo], ‘purls] oipeg

epHo,y “Yovog vuoyseqd
eplo[y ‘Yovog PooMAT[OPy
eploy,y ‘euljsnsny 3S

[BAIIJOY «JO voly

96

roma
CEG
611
col
901

pesdely
sAeq

€9/16/6
¢9/8/0I
¢9/1T/6
69/92/6
69/1/01
69/¢G3/6
$9/¥/¥
¢9/6/0I
€9/06/TT
69/08/8
£9/S/F¥
69/L1/6
69/¥6/6
69/¥6/6
69/¢6/6
€9/G6/6
69/LT/0L
£9/6/%
€9/81/01
¢9/Z/0I
¢9/¢/0I

[BAI oY
jo ayeq

OT L8
O18
OT'L8
OL L8
OT L8
86 &8
O'L8
OT L8
OT L8
OI L8
OV'S8
OV'S8
OV S8
90°S8
90°S8
90°S8
90°S8
90°S8
90°S8
90°S8
90°S8

"“M ‘sUuO'T
SUOI}VOO'T osvaleYy

vyep UINjJol 9]}90q-31IG

(WU09) 2 ATAV.L

Ieee
T€'eG
TES
I€€G
Ieee
VE EC
L&€¢
Teo
[eC
Ieee
GG CG
GG CG
G6 GG
9V IG
9V'IG
9V'1G
9V 1G
OV IG
9V' 1G
9V IG
OV IG

‘NWT

¢9/01/8
¢9/01/8
€9/01/8
¢9/01/8
€9/0I/8
¢9/01/8
¢9/01/8
69/01/8
69/01/8
€9/01/8
£9/6/8

¢9/6/8

¢9/6/8

69/16/9
69/13/9
¢9/61/9
€9/1G/9
69/13/9
69/12/9
€9/16/9
€9/16/9

asvoloy
jo 93eq

VV
VV
VV
VV
VV
VV
VV
VV
VV
VV
GV
GV
GV
LV
LV
LV
LV
LV
LV
LV
LV

868
988
LL8
GL8
OLS
698
898
L98
098
898
VSL
VLL
692
S89
V89
€89
089
8L9
TL9
OLO
699

joqunyN sJequinyy

U01}L3S

anog

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

238

plop, “OSsurUte,y

svxay, “yYorog vpsosejieyy

eplo[y ‘sSAoy vpriopy ‘Aoy punoyAoi5y
eplo[y ‘[etoavuey Wog

eplop,y “yorog eurdluig MoN

BpHo[y “PoomAyjoy

eplopy “SAoy eprliop.y “Aoy UorjeyuRyg
eplo,y “yovog Avijoqd

eplo,y “Yovog euIAUg MON

eplo,y “olueyperpuy

eplo,y “Yovog eurdulg MON

BPLIO[ YY “AO1NIG “Wy

VplIO[y ‘SoA, Aoy IVvou ‘Aoy epnorvsreg
eplIo,y “oy[IASnLL,

epllo, yy “yovog poomAyTjop{

eplopy “Yovog wyeg

epHo,y “yovog puouligd

eplopy Tey “orog ueploy
eplo, yy “vuevjuey

eploy,y ‘Ayunop speq ‘yovog ueploy

[PASI Y jo voly

PPI ¢9/01/II
CSI 69/06/01
06 69/91/6
66 69/13/6
Tau 69/01L/0L
OZT €9/L1/01
OFT 69/66/6
ZOE £9/S1/G
6FI €9/G3/6
GST €9/86/6
ZT €9/83/6
L&T 69/¢/01
60% 69/¥6/TT
PLI €9/06/01
OLT ¢9/9T/OI
CL 69/01/L
Gle €9/LZ/TT
68 69/06/L
PPS 69/86/61
931 69/6/6
posde[y [PASLQOY
sAeq jo 93eq

gc'98
6G G8
IV'98
GG G8
IvV'98
IV 98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98
80°98

"MM ‘sUuO'T

VE0G
SV'0G
99°06
CcV'0G
99°06
99°06
Ces
Gels
ces
ces
Gel%
Cels
cels
cei?
Cel’
ces
ces
cols
cele
CE1G

‘NIT

SUOT}COO'T OSvofOY

¢9/61/9
69/61/9
€9/81/9
69/06/9
¢9/81/9
69/81/9
69/66/F
69/66/F
69/66/F
69/66/F¥
69/66/F
69/66/¥F
69/66/F
69/66/F¥
69/66/F¥
69/66/F¥
69/66/F
69/66/F
69/66/F
69/66/F

asvaloy
jo 91eq

ce
VV
cE
VV
ee
cE
GE
GE
Ce
GE
GE
GE
GE
GE
CE
(eS
(oS
GE
Ce
GE

GGV8
108
8608
0608
9192
TvvL
9608
SS6L
9E6L
8682
C68L
9881
9L8L
OI8L
8082
S991
TS9L
IV9L
Sc9L
VCOL

Toq UN AY Ioquun NY

UO1}L4S

vyVp WML 9[}0qG-yUd

(709) 2 ATAVL

ano

‘Sims AND INGLE: Spiny Lobster Recruitment 239

IcurvE, T. 1962. Circulation and water mass distribution in the Gulf of
Mexico. Geofisica Internacional, vol. 2, no. 3. pp. 47-76.

IncLE, R. M., B. EvprREp, H. W. Sims, Jr., AND E. ELprRED. 1963. On the
possible Caribbean origin of Florida’s spiny lobster populations. Florida
Bd. Conserv., tech. ser., no. 40, 12 pp.

InovE, M., anp Nonaka, M. 1963. Notes on the cultured larvae of the
Japanese spiny lobster Panulirus japonicus (vy. Siebold). Bull. Jap.
soem Scieetish:, vol. 29, no. 3, pp. 211-218.

Jounson, M. W. 1960a. Production and distribution of larvae of the spiny
lobster Panulirus interruptus (Randall) with records of P. gracilis
Streets. Bull. Scripps Inst. Oceanogr. Univ. Calif., vol. 7, no. 6, pp
413-462.

1960b. The offshore drift of larvae of the California spiny lobster
Panulirus interruptus. Symposium on the changing Pacific Ocean in
1957 and 1958. Calif. Coop. Oceanic Fish. Invest. Rept., vol. 7, pp.
147-161.

Leacu, W. E. 1816. Tuckey’s narrative of an expedition to explore the
River Zaire. London. Appendix 4, pp. 413-418.

Lerpper, D. F. 1954. Physical oceanography of the Gulf of Mexico. In
Galtsoff, P. S., Gulf of Mexico. Its origin, waters and marine life.
U. S. Fish and Wildl. Ser., Fish Bull., no. 89, pp. 119-137.

Lewis, J. B. 1951. The phyllosoma larvae of the spiny lobster, Panulirus
argus. Bull. Mar. Gulf and Carib. Fish. Inst., vol. 1, no. 2, pp. 89-103.

Lewis, J. B., H. B. Moore, AND W. Basis. 1952. The post larval stages of
the spiny lobster, Panulirus argus. Bull. Mar. Sci. Gulf and Carib.
Huish. Inst), vol. 2, no. 1, pp. 324-337.

Moors, D. R. 1962. Notes on the distribution of the spiny lobster Panulirus
in Florida and the Gulf of Mexico. Cruataceana, vol. 3, no. 4, pp.
318-319.

Moore, H. B. 1958. Marine ecology. John Wiley & Sons, New York, 493
pp.

ORTMANN, A. E. 1893. Ergebnisse der Plankton. Expedition der Humboldt.
Decapoden und Schizopoden, pp. 89-90, pl. 7, fig. 3.

PrasapD, R. R., AND P. R. S. Tampr. 1957. On the phyllosoma of Mandapam.
Proc. Nat. Inst. Sci. India, vol. 23, pp. 48-67.

. 1959. On a collection of Palinurid phyllosomas from the Laccadive
Seas. Jour. Mar. Biol. Assn. India, vol. 1, no. 2, pp. 143-164.

RicuHters, F. 1873. Die Phyllosomen, ein Beitrag zur Entwicklungs-Gesch-
ichte der Loricaten. Zeitschr. f. Wiss. Zool., band 23, no. 4, pp.
623-647.

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

240

BpHOpy “Tuer, Ivou ‘opis “Yq “Ao HON L0G 79/61/T 00'°L8 C60 €9/61/9 Ses GSFS
eplo[y ‘yoreg PUOULIQ (6 €9/L1/6 9¢°98 0 0G ¢9/61/9 9¢ PSPS
Vplop,y ‘yorog ourdwog “yovog O10qsT['H 60T 69/1/01 CT'S8 L¥'0@ €9/06/9 CY O8F8
Vplo,y “Yovog vurAUg MON [OT 69/86/6 GG’98 PC 0S €9/61/9 GE Gcrs
sexo], ‘purjs] eIped El 69/¥/TT 80°L8 9¢°0% €9/61/9 US LYS

[BAoLIOY Jo voly posdey[y yRAsmyeYy AA BUTT "N 387 asvopoy = JoquinNy JoquinNy

sAeq fo 93°C] SUOT}VOOT OSvITOY jo 93eq UO01}EYS gyno,

vyep UIMjJoL 9[}I0q-1ICG

(*yu09) 2 ATAVL

SIMs AND INGLE: Spiny Lobster Recruitment 24]
Rosinson, R. K., AND D. Dimitriov. 1963. The status of the Florida spiny
lobster fishery 1962-63. Florida Bd. Conserv. tech ser. no. 42, pp. 1-30.

SatsHO, T. 1962. Notes on the early development of phyllosoma of Panulirus
japonicus. Mem. Fac. Fish. Kagoshima Univ., vol. 11, no. 1, pp. 18-23.

1964. The first phyllosoma stage of the spiny lobster, genus Panuli-
rus. Mem. Fac. Fish. Kagoshima Univ., vol. 12, no. 2, pp. 127-134.

SALSMAN, G. G., AND W. H. Totsert. 1963. Surface currents in the north-
western Gulf of Mexico. U.S. Navy Mine Defense Lab., Panama City,
Florida, Research and Development Report 209.

SHEARD, K. 1949. The marine crayfishes (spiny lobsters), family Palinuridae,
of Western Australia with particular reference to the fishery on the
Western Australian crayfish (Panulirus longipes). Austral. Council Sci.
Indust. Res. Bull. no. 247, 45 pp.

Sims, H. W., JR. 1965. The phyllosoma larvae of Parribacus. Quart. Jour.
Florida Acad. Sci., vol. 28, no. 2, pp. 142-172.

SmirH, F. G. W. 1948a. The spiny lobster industry of the Caribbean and
Florida. Carib. Res. Coun., Carib. Comm. Port-of-Spain, Trinidad, Fish.
Ser., no. 3, 49 pp.

1948b. The spiny lobster and scale-fish industry of British Hon-
durus, with recommendations for its control and development. Rept.
to the Government of British Honduras, Coral Gables, Florida, 29 pp.
(mimeo. ).

——. 1958. The spiny lobster industry in Florida. Florida Bd. Conserv.,
ed. ser., no. 11, 36 pp.

STEWART, H. B., Jr. 1962. Oceanographic cruise report of USC and GS
ship Explorer-1960. U.S. Dept. of Commerce and Coast and Geodetic
Survey.

STOMMEL, H. 1958. The Gulf Stream; a physical and dynamical description.
Univ. California Press, Berkeley, 202 pp.

SVERDRUP, H. V., M. JoHNSON, aND R. H. FLEMMING. 1949. The oceans,
their physics, chemistry, and general biology. Prentice-Hall, Inc., New
York, 1087 pp.

Tapanes, J. J. 1963. Afloramiento y corrientes cercanas a Cuba. Centro de
Invest. Pesq., Cont. no. 17, 30 pp.

TuHorson, G. 1961. Length of pelagic larval life in the marine bottom
invertebrates as related to larval transports by ocean currents. In
Oceanography, Amer. Assoc. Adv. Sci., Washington, D. C., pp. 455-474.

UNIveRSITY OF Miami. 1957. University of Miami Marine Laboratory. Final
Report. Red tide studies., no. 57-18, 13 pp., 8 figs.

242 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Von BonpE, W. 1932. Post-brephalus development of some South African
Macrura. Fish. and Mar. Biol. Surv., Union of South Africa, Report no.
8, pp. 2-42.

Wuire, A. 1847. List of the specimens of Crustacea in the collection of the
British Museum, pp. i-viii, 1-143.

WirHaM, R., R. M. INGLE, AND H. W. Sms, Jr. 1964. Notes on post-larvae
of Panulirus argus. Quart. Jour. Florida Acad. Sci., vol. 27, no. 4,
pp. 289-297.

Wust, G. 1964. Stratification and Circulation in the Antillean-Caribbean
Basins. Part I. Columbia Univ. Press, New York, 201 pp.

Florida Board of Conservation Marine Laboratory, St. Peters-
burg, Florida. Contribution No. 95.

Quart. Jour. Florida Acad. Sci. 29(3) 1966 (1967)

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL Members For 1966

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Presbyterian College
Florida Southern College
Florida State University
Jacksonville University
Marymount College
Miami-Dade Junior College
Mound Park Hospital Foundation
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida
University of Tampa

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1966

President: Manrcarer GILBERT
Department of Biology, Florida Southern College
Lakeland, Florida

President Elect: Jackson P. SICKELS
Department of Chemistry, University of Miami
Coral Gables, Florida

Secretary: Joun D. Kirtsy
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: James B. FLEEK |
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: Pierce BRopKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges
should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

C7
BA
eres
Sv Quarterly Journal

of the

Florida Academy of Sciences

Vol. 29 December, 1966 No. 4

CONTENTS

Organotin esters and their reaction with Grignard reagents
McDonald Moore and F. C. Lanning

Wavellite-cemented sandstones from northern Florida
Frank N. Blanchard and Stephen A. Denahan

Notes on spiny lobster larvae in the North Atlantic
Harold W. Sims, Jr.

Account of an octopus bite Arthur C. Wittich
A small Miocene herpetofauna from Texas J. Alan Holman
Diet of the bowfin in central Florida Michael C. Diana

First Gulf of Mexico record for Lutjanus cyanopterus
Martin A. Moe, Jr.

The molid fish Ranzania laevis in the western Atlantic

C. Richard Robins

Iodine toxicity in relation to hormones L. R. Arrington,
R. N. Taylor, C. B. Ammerman, and A. C. Warnick

Variation in some snakes from the Florida Keys Dennis R. Paulson

Officers and members of the Academy

TH MA;
4 om HS SOWNayS .
=

243

248

257
265
267
276

285

287

290
295
309

Mailed February 12, 1968

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Mareins should be 144 inches all around.

Tirtes should not exceed 42 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and
Appkess in paragraph form following Literature Cited.

LITERATURE CrTeED follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLEs are charged to authors at $16.70 per page or fraction. ‘Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

LEcENDs for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($15.80 per page, $14.00 per half
page for zinc etchings; $14.50 per page, $13.25 per half page for halftones).
Drawincs should be in India ink, on good board or drafting paper, and let-
tered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. PHoToGcRAPHS should be of good contrast, on glossy paper. Do not
write heavily on the backs of photographs.

Proor must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

Vol. 29 December, 1966 No. 4

Organotin Esters and their Reaction with Grignard Reagents
McDonatp Moore Ann F. C. LANNING

IN a previous paper, Moore and Lanning (1966) reported the
preparation of two organotin esters and that tin tetrabenzoate re-
acted with phenylmagnesium bromide to produce tetraphenyltin,
triphenylearbinol, and a ketone. The ketone produced, from the
reaction of two times the stoichiometric amount of phenylmagne-
sium bromide with tin tetrabenzoate (Moore and Lanning, 1966),
was identified as benzophenone by conversion of the ketone to a
semicarbazone derivative.

In this work a further study has been made of the reaction of
organotin esters with Grignard reagents.

An ether solution of dibutyltin diacetate reacted with twice the
stoichiometric amount of butylmagnesium bromide to yield, after
hydrolysis, tetrabutyltin, 5-methyl-5-nonanol, and 2 hexanone. The
results may be accomplished by the proposal shown in Fig. 1. No
2-hexanone was formed when four times the stoichiometric amount
of the Grignard reagent was used.

Owing to the high eletropositivity of the tin atom, the tin is at-
tacked by the butyl group of the Grignard reagent. The ketone is
produced when insufficient amounts of Grignard reagent are used.

The reaction of tin tetrabenzoate with twice the stoichiometric
amount of phenylmagnesium bromide was carried out, and benzo-
phenone was isolated. These results, when considered in connec-
tion with the report of Moore and Lanning (1966) on reaction of
tin tetrabenzoate with phenylmagnesium bromide, suggests that the
reaction occurs according to Fig. 2.

In the present report, unsuccessful attempts were made to pre-
pare the organotin esters of oxalic, malonic, and succinic acids from
their sodium salts with stannic chloride in anhydrous benzene. Lan-

244 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ning (1953) could not prepare any of the silicon esters of the lower
members of the dicarboxylic acids from their sodium salts with sili-
con tetrachloride in anhydrous diethyl ether.

Bu Mg'Br_
z ee
Buz Sn*(O-C-CH3)2+2 Bu Mgt Br) ——————> [Bu,Sn*(0-C-CH3), |
O°
Bug Sn+ 2CH3-C-OMgBr
2Bu'Mg‘Br
OH O HO OMgBr Bu
Bu,Sn+CH,-C-Bu,+Bu-C-CH, <——— Bu,Sn+CH,-C-Bu,+CH.,-C- (OMgBr),
-MgBrOH
Fig. 1. Proposed mechanism for the reaction of dibutyltin diacetate with
butylmagnesium bromide.
Br-Mgt-Ph”
Oo” Onsen
(Ph-C-0), Sn +4 Ph Mgt Br>--——> [(Ph-C-O), Snt]

oO
PH, Sn+Ph-C-OMgBr
4 PhMgBr
Q H,0 De
Pha Sn+Ph3COH+Ph,C <——— Pha Sn+Ph3-C-OMg Br+Ph,-C(OMgBr)o

-MgBrOH

Fig. 2. Proposed mechanism for the reaction of tin tetrabenzoate with
phenylmagnesium bromide.

EXPERIMENTAL

Reaction of Organotin Esters with Grignard Reagent. The re-
actions of dibutyltin diacetate with Grignard reagent were carried
out with two and four times the stoichiometric amount of ethyl-
magnesium bromide.

In each case the Grignard reagent was prepared from bromo-
butane in the usual manner. It was then added through an addi-
tional funnel into an ether solution containing 0.03 mole of dibutyl-
tin diacetate. A stoichiometric amount would be 2 molecules of the

Moore AND LANNING: Organotin Esters 245

Grignard reagent to one of the organotin ester. The mixture was
stirred mechanically and maintained at the boiling point for one
hour after the addition was completed. The Grignard complex was
hydrolyzed in an ammonium chloride solution containing ice. Some
hydrochloric acid was added afterward to react with the magnesium
precipitate. The organic layer was separated and dried with Drier-
ite. The ether was removed under reduced pressure. This left a
viscous light yellow oil. This oil was distilled and the results are
given in Table 1.

TABLE 1

Product obtained from reaction of dibutyltin diacetate with
2.0 times the stoichiometric amount of ethylmagnesium bromide

Product (C,H,),-Sn (C,H,),COHCH, C,H,COCH,
% Yield 88.4 49.0* a3"
Boiling Point
Lit. 14510 84-519 PATE
Found 145-610 86-710 128-9
Index of Refraction
Lit. Np2°=1.4736 Nyp?29°=1.4341
Found Np?2=1.4730 Np?2—1.4360
2,4 Dinitrophenylhydrazone
Lit. 106
Found 105-6

* The per cent yield was based upon all the acetate-formed alcohol.
» The per cent yield was based upon all the acetate-formed ketone.

The above reaction was repeated with four times the stoichio-
metric amount of Grignard reagent and only tetrabutyltin and 5-
methyl-5-nonanol were isolated. The analyses and yields are given
im) Walle: 1.

The infrared spectrum of tetrabutyltin was identical to an au-
thentic sample. The spectrum of 5-methyl-5-nonanol showed sig-
nificant absorption bands at 1150 and 1310 cm‘ that are due to
tertiary alcohols. The 2,4 dinitrophenylhydrazone was prepared in
the usual manner from 2 hexanone and purified from ethanol.

Reaction of tin tetrabenzoate with phenylmagnesium bromide
was carried out in a .05 molar solution using twice the stoichio-
metric amount of Grignard reagent according to Moore and Lan-

246 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ning (1966). A stoichiometric amount would be four molecules of
the Grignard reagent to one of the organotin esters. The Grignard
complex was hydrolyzed in an ammonium chloride solution con-
taining ice. Some hydrochloric acid was added afterward to react
with the magnesium halide precipitate. The organic layer was sep-
arated and dried with Drierite. The benzene-ether solution was re-
moved under pressure leaving triphenylcarbinol and tetraphenyltin
along with an oil. The oil was dissolved in ethanol and a semi-
carbazone was made in the usual manner. The semicarbazone was
purified from ethanol and had a melting point corresponding to
benzophenonesemicarbazone 162-3 (Lit. 164).

TABLE 2

Product obtained by reaction of dibutyltin diacetate with

4.0 times the stoichiometric amount of ethylmagnesium bromide

(Cnesn (C,H, ),COHCH,

% Yield 92.6 73.4
Boiling Point

(ort 14510 84-510

Found 145-4610 86-710
Index of Refraction

at Np2°=1.4736 Np2°=1.4341

Found Np22=1.4730 Np22=1.4370

Experimental Attempted Preparation of Organotin Esters of the
Dicarboxylic Acids. The method was a modification of that used
by Moore and Lanning (1966) to prepare the organotin esters of
two monocarboxylic acids. Stannic chloride (5.8 ml .05 m), dis-
solved in 50 ml of benzene, was added dropwise into slurry of 1%
times the calculated amount of anhydrous salts dispersed in 300 ml.
The stannic chloride was added slowly over a one hour period. The
mixture was stirred during the addition of stannic chloride and for
2% hours thereafter. The benzene solution was allowed to stand
for 12 hours and gave a positive chlorine test. The mixture was
stirred and kept at the boiling point for two hours. The reactions
did not take place as a positive chlorine test was obtained again
after the solution had stood for another 12 hours.

Moore AND LANNING: Organotin Esters 247

ACKNOWLEDGMENTS

The authors are indebted to the National Science Foundation for
financial support of this investigation (RPCT-GY-2422) and the
Peninsular Chemresearch, Inc., for an authentic sample of tetrabutyl-
tin. They also wish to express their appreciation to Dr. Mary Guy
of the University of Florida for the infrared spectra and to Robert
Swaine, student at Central Florida Junior College, for his coopera-
tion.

LITERATURE CITED

LANNING, F. C. 1953. Preparation and properties of silicon tetrapropionate.
Jour. Amer. Chem. Soc., vol. 75, p. 1596.

Moors, M., anp F.. C. LANNiING. 1966. Organotin esters. Quart. Jour. Florida
Acad. Sci., vol. 29, pp. 73-76.

Department of Chemistry, Central Florida Junior College, Ocala,
Florida (present address: Erling Riis Research Laboratory, Inter-
national Paper Co., Mobile, Alabama 36601); Kansas State Univer-
sity, Manhattan, Kansas.

Quart. Jour. Florida Acad. Sci. 29(4) 1966( 1968 )

Wavellite-Cemented Sandstones from Northern Florida
FRANK N. BLANCHARD AND STEPHEN A. DENAHAN

MICROCRYSTALLINE wavellite is known to be a constituent of the
aluminum phosphate zone of the Bone Valley Formation of central
Florida, and brief reference has been made to microcrystalline
wavellite in the Hawthorn Formation (Altschuler et al., 1956 and
Espenshade and Spencer, 1963). Megascopic spherulitic wavellite
aggregates and spherulitic nodules of quartz grains cemented by
microcrystalline wavellite, from the Bone Valley Formation, have
been described by Bergendahl (1955). The purpose of this article
is to report a number of occurrences of wavellite, similar to that
described by Bergendahl, in the Hawthorn Formation and in other
formations of Miocene age in northern Florida, and to present evi-
dence for the formation of wavellite by pseudomorphic replacement
of quartz and feldspar (not previously recorded as far as we know),
phosphate pellets, and clay minerals, and by open filling of pores
and fractures.

LOCATIONS AND OCCURRENCE

Most of the wavellite-bearing rocks described in this article were
found in weathered phosphatic argillaceous sandstones of the Haw-
thorn Formation in Alachua and Marion Counties, Florida at the
following locations:

1. Fisher sink, sec. 36, T. 8 S., R. 18 E., Alachua County
. Spider sink, sec. 2, 1.9 S., R. 18 E., Alachua’ County,
Old Thomas sink, sec. 28, T. 9 S., R. 18 E., Alachua County
Road cut (I-75 at north Gainesville interchange), sec. 19, T. 9 S., R.
19 E., Alachua County
Cadillac hill (road cut), sec. 3, T. 9 S., R. 18 E., Alachua County
Road cut (I-75 at State Road 236), sec. 18, T. 7 S., R. 18 E., Alachua
County
. Road cut (new U. S. 441, 8.1 miles north of intersection, in Ocala, of

U. S. 441 and State Road 40), Marion County

Wavellite-bearing specimens from locations 5, 6, and 7 were
found in place in poorly indurated sandstone, whereas specimens
from the other locations were found mainly or entirely as nodules
and concretionary masses in unconsolidated sediment and soil.
Other occurrences of wavellite include:

8. Road cut (State Road 146), sec. 1, T. 2 N., R. 6 E., near Ashville,

Jefferson County

rm CO LO

BS UW

i

BLANCHARD AND DENAHAN: Watvellite-Cemented Sandstones 249

Fig. 1. Photomicrographs of wavellite (w), replacing feldspar (f), micro-
cline (m), and quartz (q), respectively. Figures at left are with crossed pol-
ars; those at right are the corresponding views in plain light. Scales in milli-
meters. Specimens are from location 1, Fisher sink, Alachua County, Florida.

250 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Among the nodules and concretions, the material found at Fisher
sink is typical; there, irregular concretionary masses of sandstone
are composed of quartz grains tightly cemented with wavellite and
contain only traces of feldspar, phosphate pellets, limonite, and clay.
Some of the material looks like an ordinary well-indurated argil-
laceous sandstone, while other specimens are composed of an inter-
lacing network of irregular and closely spaced veinlets of wavellite-
cemented quartz grains, that separate poorly indurated clay-filled
cavities. The white resistant wavellite veinlets stand out in relief
as a result of differential weathering. Material from the Jefferson
County location is similar, except that some specimens consist of
white masses of nearly pure wavellite, and much of the material is
in macroscopic spherulitic aggregates. In hand specimens the cryp-
tocrystalline and microcrystalline wavellite resembles a white argil-
laceous cement and cannot be identified positively without chemi-
cal or instrumental techniques; the macrocrystalline wavellite is
white, occurs in radiating acicular crystals with well-developed
prismatic cleavage, and is easily identified in hand specimen.

IDENTIFICATION

In most of the sandstones studied, the wavellite crystals are
large enough and abundant enough so that they can be identified
readily in crushed fragments or thin sections by optical properties.
In other specimens cryptocrystalline wavellite was suspected from
observation of thin sections and confirmed by x-ray and differential
thermal analysis. Biaxial positive interference figures, with moder-
ately large 2V, are easily obtained from the more coarsely crystal-
line wavellite; crystals are elongate parallel with c, are length slow,
and have extinction parallel with the best cleavage (spherulitic ag-
gregates show “pseudointerference figures”, as in Fig. 8); measured
refractive indices for material from Jefferson County are X=1.527,
Y—1.535, and Z—1.554. These properties and values correspond
closely with data recorded for wavellite by Palache et al. (1951)
and for Florida wavellite by Bergendahl (1955).

X-ray diffractometer patterns, run on powdered whole-rock
samples and on the minus-200 mesh fraction, were compared with
the pattern yielded by wavellite from Montgomery County, Arkan-
sas, and with data from A.S.T.M. card +2-0075; these comparisons
supported the identification.

Fig. 2. Photomicrographs of wavellite (w), replacing quartz (q) and car-
bonate-fluorapatite (a), respectively. Figures at left are with crossed polars;
those at right are the corresponding views in plain light. Scales in millimeters.
Specimens are from location 1, Fisher sink, Alachua County, Florida.

22, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Differential thermal analysis (run to 800° C.) of each of the
wavellite-bearing rocks (both whole rock and minus-200 mesh frac-
tions) gave curves with strong endothermic peaks at 250° C.
Wavellite from Montgomery County, Arkansas showed the same
peak as well as an additional weak endothermic peak at 275° C. Di-
agnostic DTA curves can be obtained with either coarsely crystal-
line or cryptocrystalline northern Florida wavellite, even with small
percentages of the mineral. A partial review of the literature on
differential thermal analysis in general (Kauffman and Dilling, 1950,
and others) and specifically on phosphate minerals (Manly, 1950,
and others) suggests that this technique provides an excellent me-
thod for identification of wavellite, especially where the mineral is
cryptocrystalline and not positively identifiable in thin section.

MINERALOGY, PETROGRAPHY, AND ORIGIN

Examination of thin sections shows that wavellite has formed
by pseudomorphic replacement or original grains of the sandstone
(including quartz, feldspar, and phosphate pellets ), replacement of
matrix clay minerals and possibly secondary phosphatic cement, and
by open filling in pores and fractures. Replacement involves micro-
scopic or submicroscopic solution of one mineral species followed
by almost immediate deposition of another species in the space pro-
vided. Evidence of replacement consists of recognition of relict tex-
ture, or residuals of the original mineral, or both. If the form of the
original mineral is preserved, the replacement is described as pseu-
domorphic.

Fig. 1 (upper) shows the outline of a former feldspar crystal
which shows pseudomorphic replacement by wavellite. Replace-
ment is clearly controlled by the feldspar cleavage, as the coarse
wavellite crystals have grown with traces of their prism faces paral-
lel with traces of the cleavage of the feldspar. The wavellite

Fig. 3. Upper: Microcrystalline wavellite cement (w) and quartz (q).
Wavellite probably formed by replacement of an original clay matrix. Crossed
polars. Scale in millimeters. Specimens from location 1, Fisher sink, Alachua
County, Florida.

Lower left: Macrocrystalline wavellite cement (w) and quartz (q). Wavel-
lite probably formed mainly by cavity fillings. Crossed polars. Scale in milli-
meters. Specimen from location 8, Jefferson County, Florida.

Lower right: Photomicrograph showing spherulitic aggregates of wavellite
(note pseudointerference figure) cement (w) and quartz (q). Crossed polars.
Scale in millimeters. Specimen from location 8, Jefferson County, Florida.

BLANCHARD AND DENAHAN: Watvellite-Cemented Sandstones 253

254 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

which has replaced the feldspar is distinctly more coarsely crystal-
line than the microcrystalline wavellite of the matrix. Fig. 1 (lower)
shows a microcline crystal (characteristic twinning shows faintly )
also pseudomorphically replaced by wavellite. Within the large
mass of coarsely crystalline wavellite there are several residuals of
microcline with identical optical orientations, indicating that the
microcline was originally a single crystal grain. Quartz grains have
been replaced by coarsely crystalline radiating wavellite; “floating”
residuals of quartz (lower right corner of Fig. 2, upper) within the
large mass of wavellite, have identical optical orientations indicat-
ing that they were originally part of a single-crystal grain of quartz.
The quartz grain in the upper left corner of Fig. 2 (upper) appears
to be partly replaced by wavellite. Fig. 2 (lower) shows a large
phosphate pellet almost completely replaced by wayvellite.

Fig. 3 (upper) shows quartz grains embedded in a cement ma-
trix of microcrystalline wavellite; infrequent spherulites of more
coarsely crystalline wavellite (Fig. 3, upper left) are also present.
From Figs. 1-3 (upper) it seems probable that at least some of
these spherulites represent former grains which have been replaced
completely by wavellite. The microcrystalline nature of much of
the matrix wavellite is thought to be inherited from an original clay
matrix, now replaced by wavellite. This contention is strengthened
by observations of rocks from locations 5 and 6 where all degrees
of transition can be seen from clay to slightly and completely wavel-
litized clay.

The wavellite replacements are described as pseudomorphic
(in part) because the texture of the original rock is preserved and
reflected by differences in crystal size of the wavellite; more coarse-
ly crystalline wavellite formed by replacement of grains and from
precipitation in open space while cryptocrystalline wavellite formed
by replacement of clay.

A combination of DTA, x-ray, and optical studies of the wavel-
lite-bearing sandstones show small amounts of variscite (Blanchard
and Denahan, 1967), kaolinite, feldspars, carbonate-fluorapatite
(as grains), and infrequent “heavy” minerals associated with the
wavellite and quartz. Crandallite and millisite are reported to be
abundant in the aluminum phosphate zone of parts of the Bone
Valley Formation in central Florida, (Altschuler, et al., 1956 and
Owens, et al., 1960), but we have not yet been able to detect these

BLANCHARD AND DENAHAN: Wavellite-Cemented Sandstones 255

minerals associated with wavellite in the Hawthorn Formation in
northern Florida.

From field observations and from mineralogy and texture ob-
served in thin section, we recognize three fairly distinctive types of
wavellite-bearing rocks:

(1) Bedded sandstones were found in place at locations 5, 6, 7,
and 8. These rocks occur near the present ground surface and bear
obvious indications of intense weathering. Feldspar is almost com-
pletely lacking, kaolinite is present in at least small amounts and is
the dominant clay mineral, and phosphate pellets and grains have
been nearly or completely destroyed. Wavellite in these rocks is al-
most entirely cryptocrystalline to microcrystalline, and is commonly
mixed intimately with kaolinite and traces of relict phosphate. The
wavellite seems to have formed from pre-existing minerals, especi-
ally clay minerals and carbonate-fluorapatite grains, by reactions
with downward moving acidic solutions (as described by Altschuler
ebal., 1956).

(2) Wavellite was found as concretions and nodules in uncon-
solidated soil and sediment at locations 1, 2, 3, 4, and 8. In most of
these rocks microcrystalline to macrocrystalline wavellite cements
sand grains in interlacing networks of irregular and closely spaced
veinlets, which separate poorly indurated clayfilled cavities. The
resistant wavellite veinlets stand out strongly as a result of different-
ial weathering, and this boxwork texture resembles that seen in clay
in which desiccation cracks have been filled with sand that has
washed in. These concretions and nodules appear to have formed
largely by precipitation of wavellite in open spaces and to a lesser
extent by replacement of original minerals. The aluminum phos-
phate has been carried downward through permeable zones until
an environment favorable for precipitation was encountered.

(3) The third type of occurrence seems to be a more extreme
case of the second type. In these rocks macrocrystalline wavellite
cements quartz grains (Fig. 3, lower left) and also occurs as rather
large pure masses, some of which contain macroscopic spherulitic
aggregates of wavellite (Fig. 3, lower right).

ACKNOWLEDGMENTS

This work was supported in part by NSF Grant No. GY-330
(undergraduate research participation ) and in part by the Graduate

256 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

School of the University of Florida, through a summer research ap-
pointment (1966) granted to the senior author. Dr. J. R. Under-
wood helped in editing the manuscript.

LITERATURE CITED

ALTSCHULER, Z. S., E. B. JAFFEE, AND FRANK CutTtTiTTA. 1956. The aluminum
phosphate zone of the Bone Valley Formation, Florida and its uranium
deposits. U. S. Geol. Survey Prof. Paper 300, pp. 495-504.

BERGENDAHL, M. H. 1955. Wavellite spherulites in the Bone Valley Forma-
tion of Central Florida. Am. Mineral., vol. 40, pp. 497-504.

BLANCHARD, FRANK N., AND STEPHEN A. DENAHAN. 1967. Variscite from
the Hawthorn Formation. Quart. Jour. Florida Acad. Sci., vol. 29, no.
3, pp. 163-170.

ESPENSHADE, G. H., AND C. W. SPENCER. 1963. Geology of phosphate de-
posits of northern peninsular Florida. U. S. Geol. Survey Bull. 1118,
Di paAthgs! 12 pls:

KAUFFMAN, A. J., JR.. AND E. Don Dmtiinc. 1950. Differential thermal curves
of certain hydrous and anhydrous minerals, with a description of the
apparatus used. Econ. Geol., vol. 45, pp. 222-224.

MaANty, Ropert L., Jr. 1950. The differential thermal analysis of certain
phosphates. Am. Mineral., vol. 35, pp. 108-115.

OwENs, J. P., Z. S. ALTSCHULER, AND R. BERMAN. 1960. Millisite in phos-
phorite from Florida. Am. Mineral., vol. 45, pp. 547-561.

PaLACHE, C., H. BERMAN, AND C. FRONDEL. 1951. Dana’s System of Mineral-
ology. John Wiley and Sons, New York, vol. 2, 1124 pp.

Department of Geology, University of Florida, Gainesville,
Florida.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

Notes on Spiny Lobster Larvae in the North Atlantic
Haroip W. SIMs, Jr.

Various species of Palinuridae, spiny lobsters, and Scyllaridae,
sand lobsters, are found throughout tropical and subtropical areas
of the world as well as in certain areas warmed by tropical currents.
Although adult migrations may be extensive, they tend to follow the
coast and deeper water seems to form a barrier to adult distribution.
Widespread distribution of the species is largely accounted for by
the dispersal of pelagic larvae by ocean currents.

The phyllosoma, leaf-like and transparent, is well adapted to a
planktonic existence. The long larval life is well established by
Smith (1948); Lewis (1951); Johnson (1960) and many others.
Their wide geographic distribution is shown by Gurney (1936);
Smith (1948b) and Thorson (1961, p. 468).

In the eastern Atlantic, adult spiny lobsters and sand lobsters
range from as far south as southern Brazil, throughout the Carib-
bean, and as far north as Beaufort, North Carolina (Chace and Du-
mont, 1949; Moore, 1962). Larvae have been reported as far north
as Bermuda by Lebour (1949). To my knowledge this paper con-
stitutes the first report of phyllosoma larvae captured in the North
Atlantic Current.

My report deals with a collection of phyllosoma larvae made by
Dr. Rudolf S. Scheltema, of Woods Hole Oceanographic Institution.
Collections were taken over a two year period from October 1962 to
October 1964, during various cruises made between Woods Hole,
Massachusetts and the Azores. Samples were taken using a 3/4
meter plankton net fished in an oblique or vertical manner from 200,
100, or 30 meters to surface. The time of each tow varied from
10 to 30 minutes. The samples were preserved in formaldehyde.
The phyllosomes were later separated from other organisms and
shipped to the author. Thirty-three phyllosomes of four genera
were captured, the majority of them between 30-45° N. lat. and 72-
64° W. long. However, live specimens were caught as far northeast
as 30° 45’ N., 32° 05° W. A list of stations yielding phyllosoma
larvae is shown on Table 1. Taxonomic difficulties prevented iden-
tification beyond genus.

Of the four genera represented, Panulirus accounted for the
largest number, 27. They ranged in size from 3.4-23.0 mm (Stages

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

258

rFerorrroeoorroorr ©

poseuedg

68
€8
68
G9
6°66
001
OCI
0'€S
89
GL
Col
8
9°8
89
68
0OL
06
e8
VL
Cor
681
GS
VE
GGL
uy

“IIe J

‘ued

eel {UI QOT-0

‘ued

“IIe g aovyING

‘ued

‘ued

Aenea + 00G-0

‘ued

‘ued

‘ued

‘ued {Ul YOZ-O

‘ued

Ue! + 00Z-0

ed {Ut 00-0

eis +4 00Z-0

‘ued fUI Q0Z-0

‘ue {UL YOYSZ-O

eer {UL QOSZ-O
OS {UI 00-0

“AOS {Ut OOT-O

‘ued «Ul QOT-0

pu xt QOT-O

‘ued «Ul QOT-O
yuay MAO J,

6'8I

SC
9°06
GEG
T'€%
Ges
6:06
GIG
9'8T
6 GG
CCG
TGS

‘duro J,

OVCO

VOOG

Gc69

SO6I

OILTT
CGIG
CECG
8666
VOLT
GIGI
L000
CVLG
O€60
0080
SVv60

OUIL ],

C9XI9 MAVCOL NAVC6E
C9OXIL M0089 N/STo9t
VPOIIIG NM JSGolTL N/GGott
VITITALS MEGoLI N/GGo9E
VOXG M/GhoVS NVo6E
POX M,O000G6S NJLEobE
POXIOE M/GSoLV N6To6E
VOXI6G MSPoth NOE BE
VIOXI6G NM JAVoGr NWVEo6E
VPOXIIG M/S00GE N/SVo6E
VOXIV M1009 NOEolV
POXIG M6EoVI NCOP
COLAG M/SGoLIO NJLIoLt
COLAP MLGoOL NSToLt
COAVG MATo9S N00GE
9}EC] UOT}VOO'T

OL-V8cVv

AG 786V

6-L-IIV

6 GLILV

MESS STDIN
OS-e€T TV
8V-ET IV
Or eT Tv
SV el Iv
OF- el Tv
SS LSIUINY
Gao telihy)
SGivaclmle@
Vrcr eo [Pd
JESSE) XG

oULTY YON ey} UL pommjzdeo ovarry vUosoT[AYyd Io0F vyep uONRIS

Lt etal

uOHeIS

eit

VI

ol

NA
La

on
ret ro

oN OHNO hr OOD

ON

259

Sms: Spiny Lobster Larvae

rereooe eo ©

9681S

69 Weg 1UL_ OF=0
6 GL ued

OL Wed.

GL sO

Ls Medi

L’8 ‘urd {UL QE-(—

soynUIU QT

ies OS ee OED
cs urd

SGI ued ft OOT-O
Wy j-uoy MO J,

o1UR]yV YON

sapiuppyhiag “hog ‘snuppphag “og “snongiudog “iIeg “snaynupg “ueg :e1auay}t

9} UL poinjdvo ovAILT VULOSO A d 107 veep UO}
I IP I esl F eIep UNIS

(-WU09) T ATAVL

enbyqot

[PONT A »
== = | A800 COUN M,FToS9 NS8P06G 9-S-€S6I-V 61
== aU Oa M,90089 N/ITo9E MLVSCV ST.
Soe pe eee COIS M9€069 N,LTo8€ DAO Sey, 3 1
== Cee eODaih M,90089 NI109€ M9-F8S3V OT
‘duro J, OUurL J, o1eq UOI}VOO'T UOTzLIS ‘ON

260 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

3-10 according to Lewis, 1951). Sims and Ingle (1966) extended
the size range for Lewis’s stages of Panulirus larvae. Even these
maxima were extended in each stage by larvae caught in the North
Atlantic.

BERMUDA

Fig. 1. Location of North Atlantic stations which produced phyllosoma
arvae.

The water temperature, at time of sampling, averaged 10 de-
grees centigrade lower than the average temperature in which
southern larvae were collected. The lower temperature may in
some manner bring about larger sizes for the various stages. There
is a possibility that some of these larvae belong to the species Pan-
ulirus guttatus which is common in Bermuda and may have larger
phyllosoma stages. There are no published data on morphological
differences between the larvae of P. argus and P. guttatus and at-
tempts to separate Caribbean collections meristically have not been
successful.

One stage 6, 8.3 mm larva of Panulirus was collected at Station
AII-13-45, which is 2100 nautical miles from the known northern
range of the genus. This apparent dispersal distance is consistent
with previous estimates made by Ingle et al. (1963).

One phyllosome, genus Scyllarus, was captured at Station AII-
13-5, about 600 miles due east of Cape Cod. The larva appeared

Sims: Spiny Lobster Larvae 261

to be in the last or close to the last stage. There are no published
data on the complete phyllosoma stages of the scyllarids and to
place this larva into a species is impossible at this time. Adult Scyl-
larus americanus, S. chacei, and S. nearctus are reported to occur
as far north as North Carolina ( Holthuis, 1960).

The genus Scyllarides was represented by two specimens. One
from Station AII-13-3, 400 miles east of Cape Cod and another
from Station A284-10KG, about 300 miles east of Cape May Light.
Holthuis (1946) reports the adult of Scyllarides americanus (nodi-
fer) and S. guineensis var. bermudensis from Bermuda. The larvae
of these species have not been described. The phyllosome from
Station AII is larger and unlike other Scyllarides larva I have ex-
amined. The fore-body is as wide as long and twice as wide as the
mid or hind-body which are formed as one. The antennae are flat
and paddle-like composed of 2 segments. Antennule segments 4,
with an endopod. The second maxilla is widely expanded in the
shape of an anvil, but bears no setae. Maxilliped 1 is finger-like,
extending behind the second maxilla. Maxillipeds 2 and 3 are typi-
cal, both lack exopods. Long coxal spines are found on the third
maxilliped and on all pereiopods. Pereiopods 1-4 appear to be well
developed, all bearing setose exopods. Most of pereiopods 3 and 4
are missing on this specimen. Pereiopod 5 is formed of 3 segments
and is shorter than the tail. The uropods are formed as two pairs
budding just forward of the telson which is formed of two long
spines radiating at an acute angle. Four pairs of pleopods are
formed on short stalks. The larva is without gills. A long spine
projects upward on the dorsal carapace at the intersection of pereio-
pods 1-4. This larva is placed in the genus Scyllarides following the
key suggested by Gurney (1936).

The second Scyllarides larva, collected at Station A284-10KG is
similar to the larva in Gurney’s Figure 26, p. 431.

Three specimens of the genus Parribacus were collected: one
(7.4 mm, stage 6, at Station AII-13-10) just west of Flores in the
Azores; one (22.9 mm, stage 9, at Station A284-5K ) about 400 miles
northeast of Cape Hatteras; and one (8.9 mm, stage 6, at Station
A284-1G) about 1100 miles east of Atlantic City, New Jersey. The
larvae were similar to ones I collected in the Yucatan and Florida
Straits and fit well within the size ranges as shown in my descrip-
tions of phyllosomes of Parribacus (Sims, 1965). Adults of Parri-

262 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

_bacus antarcticus (Lund) are not reported north of Florida and
there is no record of the species occurring in the Azores.

DISCUSSION

In his interesting paper on species dispersion by ocean currents,
Gunnar Thorson reiterates the fact that phyllosomes are truly long-
distance larvae, drifting 90-120 days before settling. Consequently
these larvae should be found in plankton far from the range of the
adults. Studies in the Florida Straits showed that large numbers of
phyllosomes are carried through the Yucatan Straits where they
enter the Florida Current. As samples were taken northward in the
current, fewer larvae were taken per tow. It is probable that some
of them settle in Florida and Bahamian waters, and others are re-
tained in the cyclic eddies that gyrate shoreward or into the Sar-
gasso Sea. Some apparently are carried with the axis of the Gulf
Stream into the North Atlantic where larval development is prob-
ably slowed as the water cools and great mortality must occur. It
is evident, however, that some larvae can survive temperatures
down to 18.6°C as was noted at Station AII-13-3.

Thorson (1961) states, “A surface current of average velocity
will take some 22 to 23 weeks to pass from Cape Hatteras to the
Azores. Guppy (1917) reports drift-bottles drifting 273 days to
cover the same route. Although these figures may not be typical of
normal current speeds the fact remains that a drift of this rate was
recorded and it could be expected that a few isolated phyllosomes
could be transported in the same manner. It is probable that the
modal time is longer than that mentioned above. If so, all the
young larvae collected east of Cape Hatteras probably originated
from there or from Bermuda. Phyllosomes recruited from the
Caribbean or Florida would probably be well into the last stages
before reaching high latitudes; unless, of course, the lower tempera-
tures slow larval development. If we use Thorson’s figure of about
18 nautical miles a day, then a phyllosome hatched at Bermuda
should be near the last stage by the time it reaches the Azores. The
fact that early stages of the genus Panulirus were captured within
1000 miles of the Azores suggests that the larval life may be longer
in these latitudes. There is always the possibility that deep water
individuals produced the larvae we obtained, but there is no evi-
dence to support this theory. It is not possible to evaluate an

Sims: Spiny Lobster Larvae 263

Azores source since only a few adult samplings have been attempted
there and these have been negative. The occurrence of a stage 6
Parribacus phyllosome, just west of Flores suggests that this genus
occurs further north than Florida. A vigorous search for adult
spiny lobsters in this area would be helpful.

LITERATURE CITED

Cuace, F. A., Jk. AND W. H. Dumont. 1949. Spiny lobster—identification,
world distribution, and U. S. trade. Commercial Fish. Rev. U. S. Fish
and Wildl. Ser., vol. 11, no. 5, pp. 1-12.

Guppy, H. B. 1917. Plants, seeds and, currents in the West Indies and Azores.
William Norgate, London.

GurRneEy, R. 1936. Larvae of decapod crustacea. Discovery Reports, vol. 12,
pp. 377-440.

Ho.ituuis, L. B. 1946. The decapod macrura of the Snellius Expedition. I.
The Stenopodidae, Nephropsidae, Scyllaridae and Palinuridae. Biol.
Res. Snellius Expdn. XIV, Temminckia 7, pp. 1-178.

Ho.ttuutis, L. B. 1960. Preliminary description of one new genus, twelve new
species, and three new subspecies of Scyllarid lobsters (Crustacea De-
capoda Macrura). Proc. Biol. Soc. Washington., vol. 73, pp. 147-154.

INGLE, R. M., B. ELprep, H. W. Sims, JR. AND E. ELDRED. 1963. On the pos-
sible Caribbean origin of Florida’s spiny lobster populations. Florida
Bd. Cons. Tech. Ser., no. 40.

Jounson, M. V. 1960. Production and distribution of larvae of the spiny
lobster, Panulirus interruptus (Randall) with records on P. gracilis
Streets. Bull. Scripps Inst. Oceanogr. Univ. California, vol. 7, no. 6,
pp. 369-379.

Lresour, M. V. 1950. Notes on some larval decapods (Crustacea) from Ber-
muda. Proc. Zool. Soc. London, vol. 1, no. 2, pp. 369-379.

Lewis, J. B. 1951. The phyllosoma larvae of the spiny lobster Panulirus argus.
Bull. Mar. Sci. Gulf and Carib., vol. 1, no. 2, pp. 89-103.

Moore, D. R. 1962. Notes on the distribution of the spiny lobster Panulirus
in Florida and the Gulf of Mexico. Crustaceana, vol. 3, no. 4, pp. 318-
319.

264 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SHEARD, K. 1949. The marine crayfishes (spiny lobster), family Palinuridae,
of Western Australia with particular reference to the fishery of the
Western Australian crayfish (Panulirus longipes). Austral. Council Sci.
Indust. Res., Bull. no. 247, pp. 1-45.

Sms, H. W., Jr. 1965. The phyllosoma larvae of Parribacus. Quart. Jour.
Florida Acad. Sci., vol. 28, no. 2, pp. 142-172.

Sms, H. W., JR. AND R. M. INGLE. 1967. Caribbean recruitment of Florida’s

spiny lobster population. Quart. Jour. Florida Acad. Sci., vol. 29, pp.
207-242.

SmitH, F. G. W. 1948a. The spiny lobster and scale-fish industry of British
Honduras, with recommendations for its control and development. Rept.

to the Government of British Honduras. Coral Gables, Florida, Mimeo.,
pp. 1-29.

—-— 1948b. The spiny lobster industry of the Caribbean and Florida. Carib.
Res. Coun. Carib. Comm. Port-of-Spain, Trinidad, Fish. Ser., no. 3,
pp. 1-49.

TuHorson, G. 1961. Length of pelagic larval life in marine bottom inverte-
brates as related to larval transport by ocean currents. In Oceanography,
Amer. Assoc. Adv. Sci., Washington, D. C., pp. 455-474.

Marine Laboratory, Florida Board of Conservation, St. Peters-
burg, Florida. NFS Grants GB 2207 and BG 861; Florida Board of

Conservation Contribution No. 110.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

Account of an Octopus Bite

ARTHUR C. WITTICH

It has been long known that bites of certain species of octopuses
are poisonous (Halstead, 1959), but actual descriptions of the bites
and the sensations experienced by the persons bitten are rare in the
literature. Consequently, the following account appears worth
reporting.

While sorting biological specimens from a trynet sample, during
a regular sampling run of the Florida Board of Conservation Re-
search Vessel R/V Hernan Cortez, I received a bite from a small
female octopus (Octopus joubini Robson), with a body length of
50 mm. The sample was taken at 1855 hours on 3 March 1966,
about 18 miles west of Egmont Key, Florida (27° 37’ N, 83° 07’ W),
at a depth of 10 fathoms.

The animal adhered to the back of my left hand, rapidly
stretched all eight tentacles, and then contracted tightly. This
produced an astringent effect on my skin. Suddenly a sharp, pierc-
ing sensation occurred in my hand directly below the animal's
bulbous body. The combination of the animal's tight grip, its slip-
pery body, and the fact that my other hand was wet caused difh-
culty in removing the octopus quickly. During this slight time
interval of approximately four seconds a severe pain proceeded
from the bite region up my arm to the deltoid area. Here the pain
terminated and diffused into a vague and generalized sensation
which remained in the shoulder region.

- The wound consisted of two small circular holes which emitted
a slight stream of blood. Almost instantly a pure white welt of
about 25 mm diameter formed around the bite. The region began
to swell, and the pain in my arm subsided slightly. A commercial
antiseptic was applied to the wound.

During the following hour I experienced constant pain localized
in the bite region, accompanied by nausea, headache, and fever.
This condition remained for the first eight hours, progressively
lessening during the next 12 hours. Neither food nor liquids were
desired for about 24 hours, during which time I remained in bed.

On the morning of March 5, 36 hours after the bite, I was able
to get up and take nourishment. The next day I felt well with no

266 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

apparent adverse effects except a sensitive and swollen wrist. Nor-
mal appetite and body functions resumed.

The bite region remained swollen and had a noticeable itchiness
for the following three weeks. During this period there was a ser-
ous discharge from the unhealed wound. At the end of the third
week these conditions subsided, leaving a 3 mm wound surrounded
by a 10 mm red periphery. One month after the bite was received,
these measurements were reduced to 2 mm and 6 mm, respectively,
and healing proceeded rapidly.

The specimen had the following measurements: top of head to
tip of longest arm, 72; longest arm, 51; maximum radius, 99; mantle
length, 50 mm.

A report of another bite by the same species, O. joubini, ap-
peared in a recent number of Sea Secrets (Anonymous, 1965). The
sensations recorded by the victim of this bite, Edward A. Schumann,
M.D., were very similar to the symptoms I experienced. Both bites
occurred on the hand.

ACKNOWLEDGMENTS

Thanks are due to W. G. Lyons, of the Florida Board of Con-
servation Marine Laboratory, who identified the octopus, and to
Martin A. Moe, Jr., and Edwin A. Joyce, Jr., also of the Marine
Laboratory, who reviewed the manuscript.

LITERATURE CITED

ANonyMous. 1965. Sea Secrets, vol. 9, no. 7, p. 5.

HausteaD, B. W. 1959. Dangerous marine animals. Cornell Maritime Press,
Cambridge, Maryland, pp. 34-37.

Florida Board of Conservation Marine Laboratory, St. Peters-
burg, Florida (present address: 2116 Nekle, Tampa, Florida).
Marine Laboratory Contribution No. 104.

Quart. Jour. Florida Acad. Sci. 29(4) 1966( 1968 )

A Small Miocene Herpetofauna from Texas
J. ALAN HoLMAN

Recorps of North American Miocene amphibians and reptiles
are so few that the discovery of a small herpetofaunal assemblage
from the Miocene of the Texas Gulf Coastal Plain is of much inter-
est. The fossils were found by Bob H. Slaughter, Curator of the
Shuler Museum of Paleontology of Southern Methodist University,
who collected, washed, and sorted the material from the site, and
kindly sent the herpetological remains to me for study. These fos-
sils represent at least two salamanders, two frogs, a crocodilian, a
lizard, and two snakes. All of the six genera identified have living
representatives, but the two forms discussed at the specific level are
extinct.

A recent flood has buried the site under a considerable amount
of silt, but Mr. Slaughter will continue to study the locality, and
plans a future publication that will detail the site and its mam-
malian remains. The following information has been supplied by
Mr. Slaughter. The site lies within the Flemming formation and is
stratigraphically slightly below the Coldspring local fauna which
has been provisionally referred to the late middle Miocene by Quinn
(1955). The deposit is on the property of Mr. F. S. McGee and is
near the base of a feature called Pine Isle which was formed by the
truncation of a regional high by the Trinity River. The site lies at
longitude 95° 03’, latitude 30° 41’, and is seven miles northeast of
the town of Coldspring on the western bank of the Trinity River in
San Jacinto County, Texas.

I wish to take this opportunity to thank Mr. Slaughter for allow-
ing me to study these fossils that he collected while working under
the support of National Park Service (Southwestern Region) Con-
tract Number 14-10-0333-1712. My work was supported in part by
National Science Foundation Grant GB-5988. Donna Rae Holman
made the drawings.

Class AMPHIBIA
Order URoDELA
Family Sirenidae
Siren sp. indet.

Material. Thoracic vertebra, SMP-SMU 61869, Fig. la.

Fig. la. SMP-SMU 6189, vertebra of Siren sp. indet., ventral view; b and c,
SMP-SMU 61870, holotype vertebra of Notophthalmus slaughteri n. sp. (b,
posterior view, c, dorsal view); d, SMP-SMU 61871, holotype left ilium of
Hyla miocenica n. sp., lateral view; e, SMP-SMU 61872, right ilium of Rana
cf. R. pipiens, lateral view; f, SMP-SMU 61874, left dentary of Eumeces sp.
indet., lingual view.

Remarks. This fragmentary vertebra represents the family Siren-
idae and is assigned to the genus Siren rather than to the genus
Pseudobranchus on the basis that the fossil has the lower margin of
its centrum very slightly curved rather than concave (Goin and

Hotman: A Miocene Herpetofauna 269

Auffenberg, 1955). The Texas fossil is similar in size to Miocene
and Pliocene species of Siren and is larger than fossil and recent
species of Pseudobranchus.

Unfortunately, certain important parts of the fossil are missing.
Thus, I feel that it is unwise to attempt a specific designation. The
possibility exists that the Texas vertebra represents a new form, as
the only other pre-Pleistocene Siren fossils known are Siren hesterna
Goin and Auftenberg from the lower Miocene of northcentral Flor-
ida, and Siren simpsoni Goin and Auffenberg from the middle Plio-
cene of northcentral Florida. Both the Florida and the Texas sirens
are small forms. The centrum length of the Texas Siren is 2.50 mm.
This length is 2.67 mm in the lower Miocene form, S. hesterna, and
it is about 3.0 mm in the middle Pliocene S. simpsoni (estimated
from Fig. 1 in Goin and Auffenberg). The Texas fossil probably
represents an adult as it has an interrupted notochordal canal.

Following is a description of the fossil. The neural arch, the ali-
form processes, and the neural spine are missing. The left prezyga-
pophysis is missing. The right prezygapophysis is ovaloid in shape
and its prezygapophyseal face is flat and has its margin slightly
elevated. The centrum is amphicoelous, constricted at its middle,
and has it notochordal canal interrupted by a bony partition. The
left transverse processes are broken. The right transverse processes
are posteriorly directed. The lower margin of the centrum is only
slightly curved when viewed from the side, and the bottom of the
centrum has a well-developed keel. There is a large fossa on each
side of the keel at about its middle. The right fossa is smaller and
slightly posterior to the left one. Marked depressions occur on each
side of the keel.

Family Salmandridae
Notophthalmus slaughteri n. sp.

Holotype. Dorsal vertebra, SMP-SMU 61870, Fig. Ib and Ic.

Diagnosis. A Notophthalmus that resembles Notophthalmus ro-
bustus Estes of the lower Miocene of northcentral Florida in its low
neural spine, but differs from N. robustus and is similar to modern
species of Notophthalmus in having its rib bearers longer and less
stubby.

Etymology. The species is named in honor of its collector, Bob

270 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

H. Slaughter, Curator of the Shuler Museum of Paleontology of
Southern Methodist University.

Description. The vertebra is relatively complete, but it has part
of its right postzygapophysis and the anterior part of its neural
spine missing. The nural spine is relatively low and its top portion
is a thickened cap of pitted dermal bone. The prezygapophyses
have round faces. The neural arch is relatively low and the neural
canal is slightly depressed. In anterior view, the prezygapophyses
have their edges turned upward slightly and the rib-bearers are
relatively long and narrow. The cotyle is round and the noto-
chordal canal is interrupted. In posterior view, the dorsal part of
the neural arch is quite robust, but the ventral portion is thinner.
The bottom of the centrum is only slightly constricted at its middle
and is quite distinct from the transverse processes which have their
bases highly sculptured.

Remarks. Estes (1963) has pointed out that of the living genera
of the Salamandridae, only Notophthalmus and Cynops have the
same peculiar pattern of the dermal cap of the neural spine, and
furthermore that Cynops lacks the extensive pitting on the dermal
cap that is found in Notophthalmus. The fossil vertebra agrees
with the latter genus in this character. It seems quite possible that
N. slaughteri is the temporal equivalent of N. robustus, as the major
difference between the two is the more robust rib-bearers of the
latter species. Notophthalmus slaughteri resembles N. robustus and
differs from the recent species in having a lower neural spine, al-
though some vertebrae of a recent N. viridescens (Rafinesque )
from Brown County, Indiana have neural spines that are almost as
low as in N. slaughteri.

In summary, it should be emphasized that the vertebrae of all
known Notophthalmus from Early Miocene to the present are very
similar in detailed structure and differ mainly in relative propor-
tions.

Order ANURA
Family Hylidae
Hyla miocenica n. sp.
Holotype. Left ilium, SMP-SMU 61871, Fig. 1d.

Diagnosis. A moderately large Hyla that shows similarities to
Hyla arenicolor Cope, Hyla squirella Sonnini and Latreille, and

HoLtMAN: A Miocene Herpetofauna 271

Hyla versicolor Le Conte, but differs in that its dorsal protuberance
is more oval in outline than in the former two species, and is less
flattened and more produced from the shaft than in the latter
species.

Description. The terminology of this section follows Chantell
(1964). The ilium is complete except that part of the dorsal sur-
face of the dorsal protuberance is broken, and the tips of the dorsal
and ventral acetabular expansions are partially broken. The dorsal
prominence is only moderately developed. The dorsal protuber-
ance is oval in outline, is well produced laterally, and is longer than
it is high. The surface of the dorsal protuberance is rounded, and
the protuberance is moderately distant from the acetabular border.
The acetabulum is well excavated and has a distinct border. The
ventral acetabular expansion is extensive and its anterior border
makes a wide angle with the shaft. The shaft is slightly curved and
lacks either a crest or a ridge.

Discussion. The Texas fossil Hyla resembles some of the recent
species of Hyla more than it does hylids that have been described
from the lower Miocene of Florida and from the “Mio-Pliocene”
of Nebraska. Hyla miocenica differs from Hyla goini Auftenberg
from the lower Miocene of northcentral Florida in that its dorsal
protuberance is longer and is farther from the acetabular border.
Hyla miocenica differs from Pseudacris nordensis Chantell in having
a much more extensive ventral acetabular expansion and in having
its dorsal protuberance closer to the acetabular border. Upon com-
parison with species of North America Hylidae the fossil appears
most similar to Hyla arenicolor, H. squirella, and H. versicolor all
of which have individuals that are rather close in structure to H.
miocenica. Nevertheless, H. miocenica may be separated from in-
dividuals of these species available to me on characters given in the
diagnosis section. Unfortunately, it is impossible to say to which
of the recent species the fossil species is most closely allied.

Family Ranidae
Rana cf. Rana pipiens Schreber

Material. One left and one right ilium belonging to separate in-
dividuals, one sacral vertebra, SMP-SMU 61872, Fig. le.

Remarks. These ilia are not distinguishable from those of recent

272 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Rana pipiens (Holman, 1965) and are tentatively assigned to this
species. The ilia represent two small individuals.

Based on the report of Tihen (1954) the Texas fossil sacrum is
also similar to that of recent Rana pipiens. The length of the cent-
rum of the fossil is 2.0 mm, the width is 2.0 mm, thus the ratio of
the length of the centrum divided by the width of the centrum is
1.0. This indicates the fossil falls into Tihen’s (op cit. p. 219, Fig.
1) Rana pipiens group. The intercondylar space of the fossil meas-
ures 0.6 mm. The ratio of the width of the centrum divided by the
intercondylar space is 3.33. This ratio also indicates the fossil is
similar to Tihen’s (op. cit. p. 220, Fig. 2) R. pipiens group. I can
find no subjective characters to distinguish the fossil sacrum from
those of recent Rana pipiens at hand.

This is only the second Miocene locality in the New World that
has yielded the genus Rana. Two extinct species of Rana and Rana
cf. R. pipiens have been identified from the early Miocene beds of
northcentral Florida (Holman op. cit.). One of the extinct forms,
Rana bucella, shows no close affinities to recent or fossil Rana spe-
cies, but the other form, R. miocenica, is similar to recent R. pipiens.

Class REPTILIA
Order CrocopiLiA
Family Alligatoridae
cf. Alligator sp. indet.

Material. Two small teeth, SMP-SMU 61873.

Remarks. These teeth have a thick enamel layer with a finely
striated surface and have a sharp anterior and posterior keel as seen
in recent Alligator mississipiensis (Daudin ) teeth.

Order SQUAMATA
Family Scincidae
Eumeces sp. indet.

Material. Left dentary, SMP-SMU 61874, Fig. 1.

Remarks. This partial dentary is referable to the genus Eume-
ces, but it is not assigned to species because of its fragmentary
nature, and because several species of recent Eumeces are not avail-
able as skeletons. The bone represents about the anterior one-half
of the lower jaw of a skink a little larger than a recent Ewmeces
obsoletus (Baird and Girard) with a skull length of 19.7 mm as

Houtman: A Miocene Herpetofauna 273

measured from the tip of the rostrum through the occipital condyle.

A description of the fossil dentary follows. The Meckelian
groove is open and the ramus above this groove is robust. Nine
teeth and remnants of teeth are present. The teeth are columnar
and regularly spaced. The crowns are swollen and quite distinct
from their bases. The tooth crowns are striated and show patterns
of wear only on the lingual side. These patterns of wear on the
crowns take the form of grooves. None of the recent skeletons I
have examined show as much wear on the tooth crowns. Each of
the two posterior complete teeth is about one-fourth produced
above the dorsal rim of the dentary, whereas each of the two an-
terior complete teeth is about one-third produced above the dorsal
rim of the dentary. In labial view, the bases of the teeth are quite
constricted and have their surfaces very smooth. There are four
foramina on the labial side of the dentary. The most anterior fora-
men is at the tip of the bone and is ventral, the most posterior one
is dorsal, and the middle two are intermediate in position. Eumeces
sp. from the early Miocene of northcentral Florida (Estes, 1963) is
the earliest record of the genus. There are no other Miocene rec-
ords.

Family Columbridae
Colubridae sp. indet.

Material. Fragmentary vertebra, SMP-SMU 61875.

Remarks. This vertebra is too fragmentary for sufamilial identi-
fication, but on the basis of its being longer than wide, its moder-
ately high neural spine, and the thin base of its broken hypapo-
physis it is referred to the family Colubridae.

Family Viperidae
Viperidae sp. indet.

Material. Vertebra, SMP-SMU 61876.

Remarks. This vertebra represents the first Miocene record of
a viperid snake from North America. Previously, the earliest rec-
ords are those of Brattstrom (1954) who reported Agkistrodon
contortrix (Linnaeus) and Crotalus viridis (Rafinesque) from the
early Pliocene of Nebraska.

A description of the vertebra is as follows. In dorsal view, the
vertebra is wider than long. The neural spine is broken off. The
right prezygapophysis is broken and the left prezygapophyseal

274 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

shape. The accessory process of the left prezygapophysis is broken.
In anterior view, the neural arch is robust, the cotyle is round and
is larger than the neural canal, and the depressions on either side of
the cotyle are moderately deep. A rather large fossa is present in
the left of these depressions. The transverse processes are almost
entirely broken. In ventral view, the centrum is triangular in shape
and is wider anteriorly than it is posteriorly. The hypapophysis is
broken at its base, but it is thick as in modern viperids. Moderately
excavated depressions occur on either side of the bases of the hy-
papophyses. In posterior view, the neural arch is robust and the
condyle is round and about the same size as the neural canal.

DISCUSSION

Perhaps the most striking aspect of the fossil herpetofauna is
that its genera are modern. In fact, all six of them can be found in
or near the area today. It is also interesting to note (although
snake material is scarce) that the Colubridae and the Viperidae are
the only snake families represented, for earlier North American
Cenozoic deposits have their ophidian faunas almost completely
dominated by boid snakes (Holman, 1964).

Based on the habitat preferences of the majority of living spe-
cies of most of the genera identified in the present study (Siren,
Notophthalmus, Hyla, Rana, and Alligator), a quiet water situation
such as a marsh or a swamp was probably an important feature of
the area during the time of the accumulation of the fossils, The
type of terrestrial situation represented is unknown since the specific
affinities of Ewmeces and the snakes are obscure.

LITERATURE CITED

BRATTSTROM, B. H. 1954. The fossil pit-vipers (Reptilia: Crotalidae) of North
America. Trans. San Diego Soc. Nat. Hist., vol. 12, pp. 31-46.

CHANTELL, C. J. 1964. Some Mio-Pliocene hylids from the Valentine forma-
tion of Nebraska. American Midl. Nat., vol. 72, pp. 211-225, 4 figs.

Estes, R. 1963. Early Miocene salamanders and lizards from Florida. Quart.
Jour. Florida Acad. Sci., vol. 26, pp. 234-256, 4 figs.

Gorn, C. J., AND W. A. AUFFENBERG. 1955. Fossil salamanders of the family
Sirenidae. Bull. Mus. Comp. Zool., vol. 113, pp. 497-514, 3 figs.

HotMaAn: A Miocene Herpetofauna 275

Houtman, J. A. 1964. Fossil snakes from the Valentine formation of Nebraska.
Copeia, 1964, pp. 631-637, 3 figs.

1965. Early Miocene anurans from Florida. Quart. Jour. Florida Acad.
Sci., vol. 28, pp. 68-82, 2 figs.

Quinn, J. H. 1955. Miocene Equidae of the Texas Gulf Coastal Plain. Bureau
Econ. Geol. Univ. Texas Publ. 5516, pp. 1-102, 5 figs., 14 pls.

THEN, J. A. 1954. A Kansas Pleistocene herpetofauna. Copeia, 1954, pp. 217-
2 Ne 2eties:

Department of Biological Sciences, Illinois State University,
Normal, Illinois 61761 (present address: The Musuem, Michigan
State University, East Lansing, Michigan 48823).

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

Diet of the Bowfin in Central Florida
MIcHAEL C, DIANA

Stupy of the diet of bowfin (Amia calva) in central Florida was
undertaken in hopes that it would add to the scarce knowledge of
the local food habits of the species, and give an indication of its
role in managing fish populations. The study was conducted while
I was employed by the Florida Game and Fresh Water Fish Com-
mission as part of the federal aid project F-17-R according to the
Dingell-Johnson Act. I am indebted to Roger A. Martz and Ru-
dolph H. Howell for assistance in capturing the fish, and to Dr.
John C. Briggs for his help in preparing the manuscript.

METHODS

Specimens were procured by electric shocking, using a 220-volt
A.C. generator, during the first three months of 1965 between the
hours of 8:00 A.M. and 5:00 P.M. Only the fish along the littoral
zone of the lake to a depth of about eight feet were captured. No
attempt was made to shock fish in deeper water. A shocked fish
could be scooped out of the water using a dip net. Individual fish
were given a number, and records were kept of length, weight, and
sex in addition to the exact locality of the capture, the date, and the
time of day. The whole digestive tract was then cut out and im-
mersed in 10 per cent formalin solution.

Very near the conclusion of the study it was found that merely
placing the eviscerated digestive tract in 10 per cent formalin was
inadequate. It was found that tape worms in the stomach were
able to live up to five hours after the digestive tract was immersed.
There may possibly be some food deterioration after the stomach
is immersed in formalin. In future studies it would be advisable to
inject the stomachs with formalin before immersion.

In the laboratory, the stomach was separated from the intestine
at the pyloric constriction. The volume of the stomach contents
was measured by water displacement. Individual food items were
separated, identified, and counted. The volume of each food type
was determined, as well as its frequency of occurrence.

To examine the diet in relation to management, fish found in
the stomach contents from Lake Griffin and Lake Harris and Hel-
ena Run were categorized as rough fish or game fish.

D1ANA: Diet of the Bowfin 277

Stupy AREAS

In this investigation, no attempt was made to determine the
availability of food types from each location. However, observa-
tions while collecting yielded the following information which
might help to explain diet dissimilarities in the different locations.

Number

Volume

Dorosoma petenense 72%

Dorosoma petenense 64%

Unidentified
10%

Dorosoma cepedianum
24%

entrarchidae 0. 6%

Ameiuridae 0.7% sats Shee 4%
Cyprinidae 0. 7% a eae

Volume Number

Ameiuridae 40% Centrarchidae 50%

Unidentified 32%

Cyprinidae 1%

Centrarchidae
21%

Ameiuridae 10% Cyprinidae 23%

Unidentified 17%

Fig. 1. Number and volume of fish from stomachs of bowfin (Amia calva).
Upper figures are based on data from Lake Griffin, lower figures from Lake
Harris and Helena Run.

Helena Run, partly a spring run fed by Bugg Spring, is located
in Lake County and drains Lake Denim into Lake Harris. A swamp

278 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

borders the collecting area on both sides of the run. There are
many dead-end canals extending perpendicular to the run. Many
bowfin were collected in these canals, but the majority were col-
lected in the main part of the run.

Lake Griffin is a 8,800 acre lake in Lake County drained by the
Oklawaha River at the northern end. The northern third of the
lake is well covered with Nuphar. The main collecting area was
at the southwest shore. Most of the bowfin were collected in canals
and along the shore in the lily pads and maiden cane. Lake Grif-
fin is the most fertile of the collecting areas, its water very dense
with phytoplankton. This fertility is partly caused by drainage
from the Leesburg Sewage Treatment Plant and the Minute Maid
Company Citrus Processing Plant. Twin Palms is a fish camp mid-
way up the western shore of Lake Griffin.

Little Lake Harris is actually the southern end of Lake Harris
situated between Howey-In-The-Hills and Astatula. The collecting
areas were in the narrow coves and canals near Astatula. Many
snakes were observed sunning on the Nuphar and Eichornia grow-
ing along the water's edge.

Lake Harris is a 17,000 acre lake south of Leesburg. It is less
fertile than Lake Griffin but still has an abundant shad population.
The bowfin from this location were collected along the shore to the
north and south end of the mouth of Helena Run. This is pri-
marily an area of emergent grass and cypress trees.

Venetian Gardens is a pleasure area located on Lake Harris.
The bowfin from this area were collected in the deep clear canals
lacing the area.

The area entitled “boat docking area” was also located on Lake
Harris very near Venetian Gardens. It is a small deep cove well
grown with subsurface algae appearing to be Spirogyra. A good
population of adult bass was evident.

The St. Johns River location was not in the Leesburg area. The
collecting area was out from Crow’s Bluff along the shoreline of
Norris Dead River among numerous Nuphar. This water had a
higher salinity than any other area examined. In some cases the
shocker could not be used because of the high salinity.

Lake Dora is a very fertile lake located in Lake County south
of Tavares. The fish from this area were collected along the south-
west shore in the maiden cane. > ee

Diana: Diet of the Bowfin 279

Lake Eustis is a 7,400 acre lake in Lake County located at Eus-
tis. The bowfin collected were shocked from the shore area among
the maiden cane.

RESULTS

During the course of this research, 131 bowfin were collected,
18 of which had empty stomachs. Upon examination, the full
stomachs demonstrated a wide variety of food items. Some of
these will be described to help clarify the data.

A red rubber fishing worm was found in a bowfin from Helena
Run. Since it was not nutritious, it was not entered in the data.

Two instances of bird remains were discovered in two different
stomachs, one from Little Lake Harris and one from Lake Griffin.
The first was a black bird found coiled up in the spiral valve of the
intestine, with only 12 feathers remaining in the stomach. The
other was a coot’s leg (Fulica americana). It is possible that the
black bird was eaten alive, and that the leg was torn from a live
coot, but this is doubtful. The coot’s leg was in the stomach with
distal end extending out of the lower esophageal region. If the
coot had been alive the leg’s position in the stomach should have
been reversed. Of course, the bowfin might have dropped the leg
and then reversed its position, but in my opinion both should be
considered carrion.

Another item which could also be considered carrion was found
in two bowfin from the Venetian Gardens location. This was some
type of previously cooked meat, tentatively identified as a pork
chop.

It should be noted that there is a high volume of detritus from
Helena Run. This is because one of the bowfin contained 10 ml
of mud and detritus. The same stomach also contained the re-
mains of what seemed to be either a mud siren or eel. Perhaps
the mudfish devoured the mud and detritus while in pursuit of the
food item.

In several instances, vertebrae were found that could not be
identified. They may have been small snakes, salamanders, or
lizards. In the data they are designated as unidentified vertebrates.

It should also be noted that amphipods, and snails are included.
Those found were very small and, like detritus, are probably acci-
dental food items.

280 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The detailed data giving the numbers, volumes, and frequen-
cies of food items from each location are given in the following
paragraphs.

The stomach contents of 26 bowfin from Helena Run contained:
25 fish with a volume of 123.8 ml and a frequency of occurrence of
14, 30 crayfish (82.3 ml, 15), 13 prawns (1.2 ml, 5), 30 insects
(3.76 ml, 13), 10.77 ml of detritus with a frequency of 5, 2 amphi-
pods found in one stomach, 1 snail (0.05 ml), 2 unidentified verte-
brates (0.4 ml, 2), 4 unidentified items (4.3 ml, 2), 1 snake (175
ml), and | mud siren or eel with a volume of 50 ml.

Twenty-three bowfin from Lake Griffin contained: 51 fish (288
ml, 19), 22 crayfish (32.14 ml, 12), 286 prawns (16.02 ml, 19), 10
insects (1.42 ml, 7), 0.81 ml of detritus with a frequency of 10, 10
amphipods (0.04 ml, 6), 3 snails (0.05 ml, 4), 1 salamander (1.8
ml, 1), 1 frog (1.9 ml), and the leg of a coot found in one stomach
for a volume of 18 ml.

The 18 bowfin collected from Little Lake Harris were found to
contain: 10 fish (48.98 ml, 8), 24 crayfish (36.15 ml, 15), 12 prawns
(0.5 ml, 5), 7 insects (1.1 ml, 5), 0.04 ml of detritus from 3 stom-
achs, 1 amphipod, 2 snails (0.01 ml, 1), 4 unidentified items (1 ml,
3), 2 snakes (123 ml, 2), 1 salamander (0.4 ml), 1 frog (1.9 ml),
and the remains of a black bird.

Twelve stomachs from Lake Harris contained: 11 fish (206.4 ml,
8), 11 crayfish (16.73 ml, 9), 9 prawns (1.28 ml, 4), 9 insects (1.38
ml, 4), 4 unidentified vertebrates (4.57 ml, 4), 1 snake (54 ml),
and 1 salamander with a volume of 0.6 ml.

The ten stomachs from Lake Dora contained: 9 fish (132.5 ml,
9), 1 prawn, and 1 mud siren or eel (2.5 ml).

Nine stomachs from the St. Johns“River contained: 2 fish (230
ml, 2), 3 crayfish (9 ml, 1), 7 prawns (0.42 ml, 2), 2.6 ml of detri-
tus from 2 stomachs, 1 amphipod (0.02 ml), 1 snake (50 ml), and
2 fiddler crabs (0.9 ml, 2).

From Venetian Gardens, the five bowfin collected were found
to contain: 2 fish (27 ml, 3), 2 crayfish (10.6 ml, 2), 0.8 ml of detri-
tus from one stomach, and cooked meat remains found in two
stomachs for a volume of 47.2 ml.

Five stomachs from Lake Eustis contained: 5 fish (2.95 ml, 3),
5 crayfish (5.4 ml, 3), 82 prawns (12.4 ml, 5), 13 insects.(1.6 ml,
2), 5 amphipods found in two stomachs, and 2 snails (0.02: ml,-I).

Diana: Diet of the Bowfin 281

Four stomachs from Twin Palms contained: 5 fish (2.43 ml, 4),
3 crayfish (12.2 ml, 2), 273 prawns (22 ml, 4), 1.5 ml of detritus
from two stomachs, 10 amphipods (0.03 ml, 2), 1 snail (0.02 ml),
and 1| unidentified item with a volume of 0.8 ml.

Two stomachs from the “boat docking area” contained: 1 fish
(mle crayfish (1:7 ml, 2), and 0.2 ml of detritus from both
stomachs.

The stomach from Lake Yale contained: 1 prawn, and 1 un-
identified vertebrate with a volume of 1.8 ml.

The total of 113 full stomachs collected during the study con-
tained: 121 fish (1064 ml, 72), 102 crayfish (206.2 ml, 61), 684
prawns (53.82 ml, 46), 69 insects (9.26 ml, 31), 16.52 ml of detritus
from 23 stomachs, 28 amphipods (0.09 ml, 13), 8 snails (0.2 ml, 8),
7 unidentified vertebrates (6.77 ml, 7), 9 unidentified items (6.1 ml,
f)) oesmakes| (402,ml, 5), 3 salamanders (2.8 ml, 3), 2 frogs
(25.9 ml, 2), 2 fiddler crabs (0.9 ml, 2), 2 instances of bird remains
(18 ml, 2), 2 mud sirens or eels (52.5 ml, 2) and cooked meat for
a volume of 47.2 ml found in two stomachs.

Further identification of the fish taken from the stomach con-
tents of the bowfin from Helena Run and Lake Harris showed the
following groups of fish present: 15 centrarchids ranging 1-6 inches
in length, 3 catfish from 2-8.5 inches long, and 7 cyprinodontids
from 1-1.5 inches long.

In comparison, the stomach contents of bowfin from Lake Grif-
fin contained 30 threadfin shad, 2-5 inches long, an 8% inch gizzard
shad, 3 cyprinodontids, each 1 inch long, one 3 inch catfish, and
one 1% inch centrarchid.

DISCUSSION

The bowfin has been generally reported to be a piscivorous fish,
and the present study showed the diet to be 56 per cent fish (Table
1). However, Berry (1955) noted that out of 77 stomachs from
Newnan's Lake, Florida, only two were found that did not contain
fish. A similar situation was found only in Lake Dora (Table 1).

The occurrences of crayfish, frogs, insects, carrion, and snails
are similar to the variety of food items found by Lagler and Hubbs
(1940) and by Lagler and Applegate (1942) from southern Michi-
gan waters. The rather high percentage of prawns and snakes may
be peculiar to the diet of bowfin in central Florida (Table 1).

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

282,

(G6) ST
(60) LI
(Z) 8I
(GT) 9T
(EI) FAL
(9) 8
(FI) 9

SNOQUPT[IOSTIN,

(G0) 8

(1) 0@
(70) maa

(8'0) 8Z

BJOOSU]

) g9
) IF

(G0) SI
(€'0) 6I
(L) OL
(ZO) €T

SOJVDUOULILTE

*(eumnyoa Aq sasoyjuered ul pue) Joquinu Aq UdaATS ore SoseyUI0IIg ;

(OL) OL
(KS) Il
(9) GZ
(IL) JUS
(L) g
(61) 8Z

oeplorysy

soyuedi1ag

1(Da]D9 Duuy ) UYMOg JO syU9}UOD YORUIO}S

T WIav.L

(9G) ZI suouttoods |TV
(6L) 8I TOATY suyol 3S
(86) &8 BIO Oe]
(Y6/l,)) V6 SILIv ET oye'T
(13) OT StueyE oyxe'T 9yIT
(08) IT Uy) 9e"]
(83) FZ uny Pus[oH
SOOSIg

Diana: Diet of the Bowfin 283

Neither of these items was reported by Scott (1938), Lagler and
Hubbs (1940), Lagler and Applegate (1942), or Berry (1955).

In 1955, Berry found that the bowfin provided no control on the
shad population in Newnan’s Lake. Scott (1938), on the basis of
a food study in which he found that bowfin ate mostly game fish
in Indiana lakes, reported that bowfin are destructive and useless.
It is important to note that 88 per cent of the volume of fish found
in the stomach contents of bowfin taken from Lake Griffin contained
shad (Fig. 1). From a fish management standpoint, it appears
that the bowfin in Lake Griffin may exert a considerable control
on the shad population. The bowfin from Lake Griffin contained
only about 1 per cent game fish by volume. Shad were not evident
in the stomach contents from Lake Harris and Helena Run, and
there was an increase in game fish and catfish consumption repre-
senting greater pressure on desirable species in these waters (Fig.
1).

CONCLUSION

From the great variety of food items contained in the bowfin
stomachs, it seems that this species will eat whatever it can catch
and swallow. The incidence of snakes in the stomach contents
indicates a very aggressive nature, a character that makes the bow-
fin a desirable species for anglers.

Whether or not bowfin can be used as a management tool de-
pends on the individual circumstances of the particular habitat in
question. A bowfin population which is harmful in one lake might
prove an effective control on rough fish in another lake.

The data show that during the months of January, February,
and March the main food item of bowfin in central Florida is fish.
Crayfish and prawns represent the second and third most valuable
food items. Snakes are present in high volume, but have a low
frequency of occurrence. Perhaps as more snakes become active
with the warming weather their value in the bowfin diet will in-
crease. The rather high frequency of insects in the diet may be
contrasted to their low volume.

LITERATURE CITED

Berry, FREDERICK H. 1955. Food of the Mudfish (Amia calva) in Lake New-
nan, Florida, in Relation to its Management. Quart. Jour. Florida Acad.
Sci., vol. 18, pp. 69-75.

284 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Lacuer, K. F., AND V. C. APPLEGATE. 1942. Further studies on the food of
bowfin (Amia calva) in southern Michigan, with notes on the inad-
visability of using trapped fish in food analyses. Copeia, no. 3, pp.
190-191.

LAGLER, Karu F., AND FrANcis V. Husss. 1940. Food of the long-nosed gar
and the bowfin in southern Michigan. Copeia, no. 4, pp. 239-241.

Scotr, Witt. 1938. The food of Amia and Lepisosteus. Invest. Indiana
Lakes and Streams, vol. 1, no. 9, pp. 110-115.

Department of Zoology, University of South Florida, Tampa,
Florida.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

First Gulf of Mexico Record for Lutjanus cyanopterus

Martin A. Mog, Jr.

A LARGE lutjanid fish, Lutjanus cyanopterus (Cuvier), cubera
snapper, was taken on 14 July 1966 with hook and line by the crew
of the R/V Hernan Cortez of the Florida Board of Conservation
Marine Laboratory. The fish was caught west of St. Petersburg,
Florida (27°43’ N, 84°10’ W) in 25 fathoms. The specimen meas-
ured 672 mm standard length and 840 mm total length, and is now
deposited in the ichthyological reference collection of the Marine
Laboratory (FSBC 4001).

This identification is based on the following salient taxonomic
characters (Rivas, 1949): vomerine patch crescent-shaped, without
a distinct background projection; upper and lower canines strong
and well developed; six gill rakers present on the lower limb of the
first arch. Additional morphological features are as follows: head
2.7 in SL; depth 2.2 in SL; maxillary 2.7 in head, reaching just to the
vertical of the anterior orbital margin; snout 2.3 in head; pectoral
1.4 in head; dorsal X-13; anal III-8; pectoral 17; branched caudal
rays 15; scale rows approximately 46.

Rivas (1949) reported on the first record of this species for the
Atlantic coast of the United States. This was a large specimen
(1,000 mm SL) taken off Ft. Pierce, Florida, on 4 May 1948. Rivas
also stated that the small L. cyanopterus are rarely taken, even in
areas where the adults are common.

Professor Luis R. Rivas, University of Miami, graciously pro-
vided the following records of this rare snapper taken from United
States waters since 1949. Monroe County, Florida, Big Pine Key,
88 mm SL, UMIM 5761; Dade County, Florida, Coral Gables Canal,
375 mm SL, UMIM 4838. In addition to the above, Christensen
(1965) reported on two small specimens, 188 mm SL and 44 mm
SL, taken from the northwest branch of the Loxahatchee River,
Florida, on 3 August and 26 November of 1960. Professor Rivas
confirmed the identifications and these specimens are deposited in
the Florida State University ichthyological collections. A few other
small specimens of L. cyanopterus (100 to 224 mm SL) taken from
Caribbean and Bahamian waters since 1949 are now deposited in
the collections of the United States National Museum and the Uni-
versity of Miami Ichthyological Museum (UMIM).

286 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The cubera snapper does not appear on any faunal lists of the
Gulf of Mexico or the Atlantic Ocean north of Ft. Pierce, Florida,
that are available to me, and I believe this is the first record of L.
cyanopterus in the Gulf of Mexico as well as the northernmost record
of occurrence in the western Atlantic. Dr. William D. Anderson,
Jr., University of Chattanooga, and Luis R. Rivas (personal com-
munications ) also know of no other Gulf or north Atlantic records
of this species. However, L. cyanopterus is probably not as rare
in the Gulf of Mexico as the above data indicate. Dr. John C.
Briggs, University of South Florida (personal communication ), in-
forms me that several fish identified to this species have been taken
from Texas waters, but unfortunately there are no records or pre-
served specimens available. It is also possible that L. cyanopterus
may have been frequently taken in the Gulf and mistaken for its
congener, L. griseus, especially since L. griseus is ubiquitous in the
Gulf and in morphology very close to L. cyanopterus.

LITERATURE CITED

CHRISTENSEN, R. F. 1965. An ichthyological survey of Jupiter Inlet and Lox-
ahatchee River, Florida. Unpublished master’s thesis, Florida State
University, viii + 318 pp.

Rivas, L. R. 1949. A record of lutjanid fish (Lutjanus cyanopterus) for the
Atlantic coast of the United States, with note on related species of the
genus. Copeia, 1949, no. 2, pp. 150-152.

Florida Board of Conservation Marine Laboratory, St. Peters-
burg, Florida. Contribution No. 115.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

The Molid Fish Ranzania laevis in the Western Atlantic
C. RicHARD ROBINS

THE occurrence of Ranzania laevis (Pennant) in Florida waters
has been suspected for some time, for anglers’ descriptions of a
strange slender sunfish scarcely could apply to anything else. A
specimen was finally obtained from the stomach of the dolphin,
Coryphaena hippurus Linnaeus, from off Palm Beach, Florida, on
April 1, 1962. This specimen (UMML 10490) was donated to the
Marine Laboratory collection by the late Al Pflueger, Miami taxi-
dermist. It represents the first record from Atlantic waters of the
United States. Data for this specimen follow: length from snout
tip to upper posterior point of body 217 mm, greatest body depth
94 mm, 17 basal elements in the clavus, the attached rays mostly
lost, dorsal rays 20, anal rays 18, vertebrae 19, axil of pectoral fin
slightly below level of center of eye.

Examination of the literature for other western Atlantic records
shows lack of documentation for many of them and confusion con-
cerning the range. The following discussion summarizes past
records for the region. Jordan and Gilbert (1883, p. 967) state
that Ranzania truncatus (=laevis) is occasional off our[United
States] Atlantic coast. There seems to be no published basis for
this statement which is carried through other papers such as that
by Jordan and Evermann (1898, p. 1755) who add, once taken in
the Bermudas, again without discussion or documentation. Bean
(1906, p. 80) records it from Bermuda solely on the basis of the
Jordan and Evermann statement.

Schreiner and Ribeiro (1903, p. 83) report one specimen of
Ranzania (0.53 meters) taken at “punta de Egrepunha, em S.
Christovao” in the Bay or Rio de Janeiro, December 25, 1900. This
record is repeated by Ribeiro (1915, p. 164) and the specimen is
figured.

Pellegrin (1912) reports observations of a school of Ranzania
seen by Foureau, Governor of Martinique, off that island. Three
specimens (0.62-0.64 meters) were sent to the Paris Museum and
were identified by Pellegrin who provided also a good illustration
and color description.

Schmidt (192la, p. 6, footnote) refers to several postlarval
specimens (5-7 mm) collected by the pana in the Sargasso Sea.

288 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Schmidt (1921b, p. 77, fig. 4) refers to Pellegrin and figures larvae
(1.7, 1.8, and 2.4 mm) from the Sargasso Sea. The 1.7 mm speci-
men is perhaps not a Ranzania for it shows none of the spines
so typical of molid larvae. Schmidt (1926, p 80) again refers to
eggs and early larval stages “in great numbers, chiefly those of
Ranzania” from the Sargasso Sea and later Schmidt (1932, pp. 253-
255, figs. 196-197) illustrates the egg and 5 specimens of Ranzania
from 1.7 to 53 mm, but does not attach locality data to them.

Beebe (1934; pls. 4-5) figures adults and larvae of Ranzania.
However, the caption (pl. 4) includes the statement, “These slow-
moving thick-set fish (Ranzania truncata) are fully eight feet long”
which seems inappropriate for Ranzania as it is not known to ap-
proach such size.

Fraser-Brunner (1951) in his review of the family includes no
new information on Ranzania in the western Atlantic. Finally
Briggs (1960, p. 178) includes it in his list of fishes with circum-
tropical distributions.

Ranzania laevis, may now be added to the known fish fauna of
Florida and thus of the Atlantic coast of the United States. The
few reports of Ranzania compared to the countless sightings of
other molids suggest that it is quite rare in the western Atlantic.
It may prove more common in equatorial waters between Africa
and Brazil.

James C. Tyler, The Academy of Natural Sciences, Philadelphia,
assisted with the literature and reviewed the manuscript.

LITERATURE CITED

BEAN, TARLETON H. 1906. A catalogue of the fishes of Bermuda, with notes
on a collection made in 1905 for the Field Museum. Field Mus., Zool.,
vol. 7, no. 2, pp. 21-84, 14 figs.

BEEBE, WILLIAM. 1934. A half mile down. Nat. Geogr. Soc., vol. 66, no. 6,
pp. 661-704, 29 figs., 16 cold. pls.

Briccs, Joun C. 1960. Fishes of worldwide (circumtropical) distribution.
Copeia, 1960, no. 3, pp. 171-180.

FRASER-BRUNNER, A. 1951. The ocean sunfishes (family Molidae). Bull.
Brit. Mus. (Nat. Hist.), Zool., vol. 1, no. 6, pp. 89-121, 18 figs.

Rosins: The Molid Fish 289

JoRDAN, Davi STARR, AND BARTON W. EVERMANN. 1898. The fishes of
North and Middle America. ... Part II. Bull. U. S. Nat. Mus., no. 47,
pp. i-xxx, 1241-2183.

JoRDAN, Davin STaRR, AND CHARLES H. GILBERT. 1883. Synopsis of the fishes
of North America. Bull. U. S. Nat. Mus., vol. 3, no. 16, pp. i-lvi, 1-
1018.

PELLEGRIN, JACQUES. 1912. Sur la présence d'un banc de Ranzania truncata
Retzius a la Martinique. Bull. Soc. Zool. France, vol. 37, pp. 228-231,
1 fig.

RIBEIRO, ALEPIO DE MirANDA. 1915. Fauna Brasiliense. Peixes. V. (Eleu-
therobranchios aspirophoros). Physoclisti. Arch. Mus. Nac. Rio de
Janeiro, vol. 17, pp. [1-765], many figs. (not continuously paginated ).

SCHMIDT, JOHANNES. 1921la. Contributions to the knowledge of the young of
the sun-fishes (Mola and Ranzania). Medd. Komm. Havunders¢gelser,
ser. Fiskerei, vol. 6, no. 6, pp. 1-16, 15 figs., 1 pl.

——. 192lb. New studies of sun-fishes made during the “Dana” Expedi-
tion, 1920. Nature, London, vol. 107, no. 2681, pp. 76-79, 6 figs.

1926. Further studies of sun-fishes made during the Dana Expedi-
tion, 1921-1922. Nature, London, vol. 117, no. 2933, pp. 80-81.

——. 1932. Dana’a Togt omkring Jorden 1928-1930. Nordisk Forlag,
K¢benhavn, pp. 1-368, frontsp., 278 figs., 1 map.

SCHREINER, CARLOS, AND ALIPIO DE MIRANDA RIBEIRO. 1903. A colleccao de
peixes do Museu Nacional do Rio de Janeiro. Arch. Mus. Nac. Rio de
Janeiro, vol. 12, pp. 67-109.

Institute of Marine Science, University of Miami, Miami, Florida
33149. Contribution No. 707.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

Iodine Toxicity in Relation to Hormones

L. R. Arrincron, R. N. Taytor, C. B. AMMERMAN,
AND A. C. WARNICK

Excess dietary iodine has been shown to result in lactation
failure in rats and high mortality of the young (Ammerman et al.,
1964; Arrington et al., 1965). Levels from 500 to 2500 ppm caused
increasing mortality which approached 100 per cent at the higher
level. Ovulation rate, implantation, litter size, and birth weight
were not affected. Gestation time was not affected, but pro-
longed and incomplete parturition and lack of mothering instinct
were observed. High mortality of newborn rabbits from females
fed 250 to 1000 ppm iodine has been observed, but hamsters and
swine were not affected by 2500 ppm dietary iodine (Arrington
et al., 1965). Other studies of the effects of excess of iodine
(Correa and Welsh, 1960; Galton and Pitt-Rivers, 1959; Wolff et al.,
1949), have not measured the effects upon lactation and survival
of the young. Egg production was reduced, per cent hatchability
of eggs decreased, and hatching of eggs delayed in poultry fed
excessive iodine (Perdomo et al., 1966).

These effects of excess iodine suggested an interference with
hormone production or action. The present study was designed to
compare the effects of dietary thyroid and thiouracil with excess
iodine and to study the effects of lactogenic hormones and proges-
terone upon rats fed excess iodine.

PROCEDURE

Virgin female Long-Evans rats, fed a stock diet (Purina Labora-
tory Chow) were bred to normal males at 110-130 days. Dietary
treatments were provided by adding the thyroid hormones or
potassium iodine to the ground stock diet. Treatments started
10-12 days after mating and continued for 21 days after parturition.
Control diets contained K,CO,; in amounts to provide K equal to
that in iodine diets. Experimental diets and tap water were pro-
vided ad libitum. Females were permitted to litter in individual
cages and observations were made for number of young born alive,
lactation of the females, and survival of the young. |

Experiment 1. Dietary treatments were: (1) control, (2) con-
trol plus 2500 ppm iodine as KI, (3) control plus 28 or 56 mg
thyroid (desiccated, defatted, porcine orgin, Nutritional Biochem-

ARRINGTON ET AL.: Iodine Toxicity and Hormones 291

icals Corp.) per 100 gm diet and (4) control plus 100, 300, or 500
mg thiouracil (2-Mercapto-4-hydroxy pyrimidine) per 100 gm diet.
All diets were fed ad libitum and intake measured for the first
three weeks.

Experiment 2. Prolactin and oxytocin were administered sep-
arately and in combination to pregnant rats fed 2000 and 2500 ppm
iodine as KI. Prolactin was administered subcutaneously in daily
doses of 4 or 8 IU per rat beginning at littering in trial 1, and 8 or 16
IU beginning 10-12 days prior to littering in trial 2. Oxytocin (20
USP units per ml, Armour Veterinary Laboratories) was admin-
istered intramuscularly in daily doses of 5 or 10 USP units at 10-12
days before littering. A combination of prolactin (16 IU) and
oxytocin (5 USP units) was given to 8 rats in trial 2. An equivalent
volume of physiological saline was administered to rats not receiv-
ing prolactin or oxytocin. Hormone treatments were continued
for 6 days post littering or until young died.

Experiment 3. Progesterone (Henley and Co.) was dissolved
in corn oil and administered subcutaneously in alternate daily doses
of 2 and 6 mg per rat. Administration was started 10 days prior to
littering and was continued for 10 days after littering.

RESULTS AND DISCUSSION

Prolonged parturition, failure of lactation and high mortality of
young were observed in rats fed excessive iodine (table 1). Sur-
vival of the young from females fed 2500 ppm iodine was signifi-
cantly less (P < 0.01) than controls.

Dietary thyroid (28 and 56 mg per 100 gm feed) had no effect
upon lactation or survival. There was no incidence of lactation
failure nor was there an indication of increased lactation as indi-
cated by weaning weight of the young.

The two higher dietary levels of thiouracil (300 and 500 mg
per 100 gm diet) resulted in decreased feed intake, prolonged

parturition, lactation failure and mortality of the young which
were similar to and approximately equivalent to the effects of 2500

ppm iodine. Feed intake and survival of the young at 10 days
were significantly lower (P < 0.01) than controls. Approximately
15% of the females fed iodine or the higher levels of thiouracil
failed to complete parturition in the normal time. These were
sacrificed at 48 hours after parturition started and live and dead
fetuses were found in utero.

292 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 1

Reproduction and survival of rats fed excess iodine, thyroid, and thiouracil

% Surviving

Treatment No. No. Av. daily % Born

Litters Lactating feed, gm. alive 3 da. 10 da.
Control 21 20 20.4 98 98 91
2500 ppm I, 13 9 Maye 90 59 Ane
Thyroid, 28* Ml tt 18.3 100 97 93
Thyroid, 56* 9 9 18.9 96 97 97
Thiouracil, 100* 13 2 iy! 91 94 88
Thiouracil, 300* ES 8 Ae UP 68 Ales
Thiouracil, 500* 9 6 IPAS 75+ 69 625

*Mg per 100 gm diet.
** Significantly less than control (P<0.01).
{Calculated as young delivered; some females failed to deliver all young.

TABLE 2

Lactation of rats and survival of young females fed excess iodine and
treated with oxytocin and prolactin

% Surviving

Dietary Prolactin Oxytocin No. % Born
I2, ppm IU USP units litters alive 3 da. 10 da.
Trial 1
0 — — 8 98 98 98
2500 = — 18 99 30 26
2500 4 — 5 97 8 3
2500 8 — a 89 23 23
2500 — 5 18 92 18 155
2500 — 10 21 98 9 a
Trial 2
0 — — a 96 90 90
2000 — — 7 96 42, 35D
2.000 8 — 7 93 40 30
2000 16 — 6 82 48 48
2000 — 5 o 95 63 63
2000 16 5 8 89 39 31

ARRINGTON ET AL.: Iodine Toxicity and Hormones 293

The lactation failure produced by high levels of dietary iodine
was not alleviated by the lactogenic hormones (table 2.) Prolactin
(4, 8, and 16 IU and oxytocin (5 and 10 USP unit) administered
separately or in combination to rats fed iodine were not effective
in inducing lactation. Oxytocin appeared to increase survival in
the second trial, but a summary of both trials indicated no effect.

Weaning weights (21 days) of the young surviving from females
fed iodine or iodine plus hormones were significantly lower

(P < 0.05) than controls.

Progesterone was not effective in improving lactation of females
fed 2000 ppm iodine or the survival of their young (table 3). The
incidence of lactation failure and mortality rate was equal to or
higher among those iodine females treated with progesterone than
in those without progesterone.

TABLE 3

Effect of progesterone upon lactation and survival of young from rats
fed excessive iodine

Nee Nol % Survival
Treatment Litters Lactating 3 days 10 days

Control 3 3 96 96
Control + 2 mg

Progesterone 4 4 95 95
2000 ppm Iodine 10 5 26 25
2000 ppm I, + 2 mg

Progesterone 5 2; 27 24
2000 ppm I, + 6 mg

Progesterone 6 0 0 0

The values listed in all tables for percentage survival of young
are based on number of young born alive in each litter. Subse-
quent mortality represents, in most cases, mortality of the entire
litter. Some females lactated and a portion of them raised all
young born through the normal suckling period.

The effects of thiouracil which were similar to those of excess
iodine suggest that the excessive iodine may act as a thyroid in-
hibitor. Further experiments are required, however, to identify
the mode of action of iodine in producing the effects observed.

294 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SUMMARY

Pregnant female rats were fed 2000 and 2500 ppm iodine or thy-
roid and thiouracil to compare the effects of iodine with the hor-
mones. Additional rats fed iodine were given prolactin, oxytocin
and progesterone. Lactation failure and high mortality of the
young were observed in rats fed excess iodine and thiouracil but
thyroid was without effect. Prolactin, oxytocin, and progesterone,
at the levels studied, were not effective in inducing lactation in rats
fed excess iodine.

LITERATURE CITED

AMMERMAN, C. B., L. R. ARRINGTON, A. C. WARNICK, J. L. EDwaArps, R. L.
SHIRLEY, AND G. K. Davis. 1964. Reproduction and lactation in
rats fed excessive iodine. Jour. Nutr., vol. 84, pp. 107-112.

ARRINGTON, L. R., R. N. Taytor, C. B. AMMERMAN, AND R. L. SHIRLEY.
1965. Effects of excess dietary iodine upon rabbits, hamsters, rats and
swine. Jour. Nutr., vol. 87, pp. 394-398.

Correa, P., AND R. A. Wetsu. 1960. The effects of excess iodine intake
on the thyroid of the rat. Arch. Path., vol. 70, pp. 247-251.

Gatton, V. A., AND R. Pirr-Rivers. 1959. The effect of excessive iodine
on the thyroid gland of the rat. Endocrinology, vol. 64, pp. 835-839.

PERDOMO, J. T., R. H. Harms, AND L. R. ARRINGTON. 1966. Effect of dietary
iodine upon egg production, fertility and hatchability. Proc. Soc. Exptl.
Biol. Med., vol. 122, pp. 758-760.

Wo rr, J., I. L. Crarkorr, R. C. GoLpBERG, AND J. R. Meter. 1949. The
temporary nature of inhibitory action of excess iodine on organic iodide
synthesis of the normal thyroid. Endocrinology, vol. 45, pp. 504-513.

Department of Animal Science, University of Florida, Gaines-
ville, Florida. Florida Agricultural Experiment Stations, Journal
Series No. 2432. Supported by U.S. P.H.S. Grant No. 08760.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

Variation in some Snakes from the Florida Keys
Dennis R. PAULSON

In their study of the herpetofauna of southern Florida, Duell-
man and Schwartz (1958) discussed the variation in specimens
examined by them from different parts of that area. Among species
from the lower Florida Keys especially singled out for comment
were the snakes Diadophis punctatus Linnaeus, Storeria dekayi
Holbrook, and Thamophis sauritus Linnaeus. Recent acquisition
of additional material of these species has made it possible to
elaborate further on their divergence from mainland populations.

Specimens utilized for this study are now in the following col-
lections: United States National Museum (usnm), University of
Michigan Museum of Zoology (ummz), University of Kansas
Museum of Natural History (xu), University of Illinois Museum
of Natural History (urmNH), University of Miami Reference
Collection (umrc), and the personal collections of Albert Schwartz
(as), Richard Thomas (rr), Donald W. Buden (pws), and the
author (pre). With the exception of five specimens from the
Lower Keys in the University of Michigan collection, none of these
specimens has been discussed heretofore in print. Ventral scale
counts were made as suggested by Dowling (1951). Measure-
ments are expressed in millimeters.

ACKNOWLEDGMENTS

I wish to thank Dr. Charles F. Walker and George R. Zug for
permission to borrow University of Michigan specimens, and Dr.
Albert Schwartz, Richard Thomas, and Donald W. Buden for the
use of specimens in their collections. Dr. Schwartz, Mr. Thomas,
and Dr. Fred G. Thompson kindly read the manuscript and offered
pertinent suggestions. My wife, Mary Lynn, assisted with scale
counts, and she and Mr. Buden provided valuable companionship
and assistance in the mosquito-laden pineland of Big Pine Key.

Diadophis punctatus Linnaeus

Duellman and Schwartz (1958) discussed two specimens of this
species from Big Pine Key, noting differences from mainland speci-
mens in ventral and subcaudal counts and cephalic pattern. Their
comment, “Additional material from the lower keys may show that
the population there is worthy of subspecific designation,” prompt-

296 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ed a special effort to collect Diadophis on Big Pine Key. With the
acquisition of four more specimens, it is clear that the population
on that island indeed deserves nomenclatorial recognition. This
form may be called

Diadophis punctatus acricus, new subspecies

Holotype. usNM 151831, an adult female from 3.9 miles north
of U. S. 1, on Florida 940, on Big Pine Key, Monroe County,
Florida, collected 14 June 1964 by Dennis R. Paulson. Original
number prpP 3034.

Paratypes. All from Big Pine Key, Monroe County, Florida:
KU 79851 (original number pre 3035), UImMNH 55626 (original no.
pRP 3036), and prp 3039, two males and a female collected 14 June
1964 by Dennis R. Paulson and Donald W. Buden; ummz 107186,
a male collected 23 July 1952 by John Dickson; and ummz 111365,
a female collected 1 September 1954 by L. Neil Bell.

Diagnosis. A race of Diadophis punctatus like the nominate
race in having fifteen scale rows throughout, eight supralabials,
and a single row of large midventral spots, but differing from it and
the other named races by its pale grayish-brown head, obscurely
spotted and little contrasting chin and labials, and virtual lack of a
muchal ring.

Range. Known only from Big Pine Key, Monroe County, Florida.

Fig. 1. Holotype of Diadophis punctatus acricus, USNM 151831.
Description of type (Fig. 1). Total length 289, tail length 55.
With the normal head scalation of the species (Blanchard, 1942),

Pautson: Variation in Some Snakes 297

including paired internasals, prefrontals, supraoculars, parietals, and
loreals and a single median frontal; nostrils in the border between
the anterior and posterior nasals, but more in the former; two
preoculars and two postoculars, one anterior and two posterior
temporals, and eight supralabials and eight infralabials on each
side; eye bordered ventrally by fourth and fifth supralabials; dorsal
scales in fifteen rows throughout the length of the body, with pre-
apical pits from 0.2 of the scale length from the posterior end;
135 ventrals and 44 subcaudals; anal plate divided.

Coloration in life: dorsum of head pale grayish-brown, dorsal
ground color darkening posteriorly to become blackish by halfway
to vent, black beyond that; a faintly indicated darker vertebral
stripe in the nuchal area disappearing posteriorly in the increasingly
darker ground color; supralabials whitish, speckled with fine gray
dots; a pale yellowish-orange triangle on each side of the neck ex-
tending upward as a faintly indicated line about one scale wide,
separated from its opposite by two mid-dorsal scale rows; chin
whitish, faintly flecked with gray; belly darkening from white at
level of third or fourth ventral through pale yellow, dark yellow,
and orange to anal plate; ventrolateral markings and midventral
spots same color as dorsum at equivalent level; in about one-eighth
of the ventrals, dorsal ground color confluent with the mid-ventral
spot on one side or the other (on both sides in only a few scales );
the ventral spots transversely oriented, most approximately tri-
angular, with the apex directed anteriad; anal plate black with
orange at anterior border and white at posterior border; underside
of tail orange, the dorsal ground color extending well onto the
subcaudals in the form of bars or triangles, the latter being sep-
arated from their opposites by only about one-fifth of the width
of the subcaudal surface; iris orange.

Associated specimen. vrp 617, a male with the data “western
part of Miami,’ is a problematical specimen. Its ventral count
(128) is two below that of any mainland south Florida specimen
and is only one above the mean of Lower Keys specimens. Its
subcaudal count (47) could place it with either the Lower Keys
or mainland populations. Its coloration and pattern, however,
definitely allocate it with D. p. acricus, and I am so considering it
in further discussions. The specimen was given to me as part of
a collection made in south Florida by an individual who visited

298 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Big Pine Key on frequent occasions and is, I suspect, mislabelled.

Variation. One paratype, ku 79851, is of interest in having an
entire anal plate. Among the 1,932 specimens of D. punctatus
which Blanchard (1942) examined, entire anals occurred in only
a few specimens of the race arnyji.

Seven specimens were utilized for scale counts. Supralabials
8-8 (five specimens) and 8-7 (two). Infralabials 8-8 (four), 8-7
(one), and 7-7 (two). Preoculars 2-2 (four) and 2-1 (three).
Postoculars 2-2 (five), 2-1 (one), and 1-1 (one). Anterior tem-
porals 1-1 (seven). Posterior temporals 2-2 (two), 2-1 (one), and
1-1 (four). Ventrals 127-128 (mean, 127.3) in males and 134-136
(135.0) in females. Subcaudals 47-53 (mean, 49.5) in males and
42-44 (49.5) in females.

Total length 208-227 in males and 237-289 in females. One
male is smaller but is unmeasurable. Tail length 45-55 in males
and 46-55 in females; thus females have proportionally shorter tails.

The three specimens collected alive on 14 June 1964 were
compared with one another with regard to coloration. One, the
type, has been described above. xu 79851 was colored like the
type, but the canthus rostralis was slightly paler than the cephalic
ground color. UIMNH 55626 differed from the type in its more
grayish (less brownish) head and paler belly; the latter was almost
entirely yellow, with orange coloration appearing just before the
vent. It also had less contrasting supralabials.

The dorsal ground color of all is similar in preservative. The
heads of all are pale to dark gray dorsally with paler supralabials,
the latter contrasting with the ground color. All have a faintly
indicated nuchal ring, restricted to the lower scale rows in some
and extending most of the way around in others. At its maximum,
the ring is a paler zone, less than one scale wide and interrupted
mid-dorsally by a single scale row. In a few specimens the ring
is almost indiscernable.

The infralabials and chins of the specimens vary from virtually
immaculate to faintly spotted with gray. None is conspicuously
spotted with black as in D. p. punctatus. Ventral spotting varies
from that of the type to small semicircular spots. Only in the type
are any spots confluent with the dorsal ground color. In the other
specimens, dorsal color extends onto the ventrals as short triangles.
Subcaudal dark markings extend from about one-eighth to two-
fifths the width of the subcaudal surface on either side.

PAuLson: Variation in Some Snakes 299

Comparisons. D. p. acricus differs from all other subspecies in
the extreme reduction of the nuchal ring and the paleness of the
head (the other races have distinct muchal rings from one to three
scale rows wide and heads as dark as or darker than the dorsal
ground color). It lacks the strong contrast between the dorsal
head color and the labials and chin which the other races display.
It also differs in its obscurely spotted chin and labials, unspotted in
D. p. edwardsi Merrem and contrastingly spotted in the other races.
D. p. acricus agrees with D. p. punctatus in usually having eight
supralabials and a belly regularly marked with a single mid-ventral
row of large spots.

A paratype was paler anteriorly and ventrally than a specimen
of D. p. punctatus from the south Florida mainland, with which it
was compared in life. Its iris was entirely orange, whereas the
mainland snake had a gold iris, darker and heavily overlain with
blackish ventrally and laterally.

Individual D. p. acricus taken so far are smaller than adults of
all other races except stictogenys Cope. Blanchard (1942) ex-
amined many specimens of D. p. punctatus larger than those of
acricus listed here, and about a third of specimens of both sexes of
punctatus which I examined are larger than any acricus.

The Lower Keys race can also be differentiated on the basis of
its ventral counts. These completely separate it from D. p. ed-
wardsi, arnyi Kennicott, and docilis Baird and Girard, and cluster at
the lower edge of the variation of D. p. punctatus and stictogenys.
However, ventral counts of D. p. acricus do not overlap those of
south Florida D. p. punctatus examined by me, which varied from
131-141 (mean 134.3) in males and 137-145 (mean 141.7) in fe-
males. The low counts of D. p. acricus may represent the end of
the clinal variation shown by the race punctatus from north to south
(Blanchard, 1942).

Subcaudal counts are intermediate between the high counts of
D. p. edwardsi and docilis and the low counts of D. p. arnyi,
stictogenys, and punctatus. They overlap those of all races, includ-
ing south Florida D. p. punctatus. Specimens of the latter which
I examined showed a variation in subcaudal count of 46-51 (mean
48.4) in males and 37-44 (mean 40.6) in females.

An index figure was derived by subtracting the subcaudal from
the ventral count in each specimen. This figure ranged from 74-81

300 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

in Big Pine Key males and 83-91 in mainland males, 91-94 in Big
Pine Key females and 93-111 in mainland females. By this method,
complete separation of males and almost complete separation of
females was obtained.

A single specimen from Key Largo was discussed by Duellman
and Schwartz (1958). The ventral count (137) of this female is
at the low extreme for south Florida specimens but above counts
of D. p. acricus, larger series of which may, however, be found to
encompass that figure. The subcaudal count (40) is below the
minimum of acricus and near the mean for D. p. punctatus from
south Florida. I have not examined this specimen, but from its
scale counts and the description of its head color and pattern it is
clearly D. p. punctatus.

Remarks. Although D. p. acricus is known only from Big Pine
Key at present, it is likely to be found on other Lower Keys suffi-
ciently large to support pineland and scrub vegetation. Among
these are No Name, Little Torch, Middle Torch, Ramrod, Cudjoe,
Summerland, and Sugarloaf Keys. Most individuals of the type
series were collected when they crossed the road through the key’s
extensive pineland at night. Three specimens collected alive on
14 June 1964 were taken on the same short stretch of highway,
from 2.6 to 3.9 miles north of U.S. 1, and between 1943 and 2010
hours. The evening was warm and dry with little wind. The only
other vertebrates taken on the road were two Storeria dekayi Hol-
brook and three Bufo terrestris Bonnaterre. The habitat in which
these were taken is well illustrated by Duellman and Schwartz
(1958, fig. 6, p. 193). The substrate is almost completely rocky
with many solution holes and a few scattered loose rocks. No
snakes were found in several hours of turning rocks in the pine-
land during the day.

Etymology. From the Greek a (not) and krikos (a ring), in
allusion to the “ringlessness” of this subspecies. D. 1. regalis is the
only other form in the genus without a conspicuous nuchal ring.

Specimens examined. D. p. punctatus, total 22. SourH Caroxtna: Dor-
chester County, 7.9 mi. SW Summerville (as 304). FLorma: Jackson County,
8 mi. N. Alta (as 210); Alachua County, GAINESVILLE (DRP 1274); Putnam
County, Drayton Island (umrc 55-183); Martin or Palm Beach County, “east
side, Lake Okeechobee” (umrc 55-182); Glades County, 4.8 mi. NE Fla. 721
on Fla. 78 (pre 1289); 2.6 mi. SW Fla. 721 on Fla. 78 (pre 1524); Dade
County, no other data (umrRc 55-595); 4 mi. E. Fla. 94 on U.S. 41 (pre 3040);

PauLson: Variation in Some Snakes 301

0.5 mi. S U.S. 41 on Fla. 27 (pre 1490); Miami area (including Allapattah,
Coconut Grove, and Coral Gables) (umrc 55-180, 55-184, 55-186, 55-468; pre
1, 1292; pws 18); 4 mi. S South Miami (pre 3055); Matheson Hammock (As
249); Monroe County, 2 mi. W Dade County line on Fla. 94 (pws 94); near
Coot Bay (rr 866); East Cape Sable (umrc 55-179).

D. p. acricus, total 7. FLorma: Monroe County, Big Pine Key (type and
paratypes ); “Dade County, western part of Miami” (probably equals Big Pine
Key) (pre 617).

Storeria dekayi Holbrook

In Trapido’s monographic study of this genus (1944), no speci-
mens from the Florida Keys were listed. Duellman and Schwartz
(1958) recorded four specimens of S. d. victa Hay from the Keys
(Big Pine, No Name, and Sugarloaf) and commented upon differ-
ences in scale counts and coloration between those individuals and
others from the southern mainland. Two additional specimens
have been recently collected, furnishing additional data on the
population of the Lower Keys.

Ventral counts for two males from the Lower Keys are 129 and
131. The same figures for three females from that area are 128,
136, and 140. Subcaudal counts of these specimens are 55 for a
single male and 49, 49, and 51 for the females.

Ventral counts of nine males from the southern Florida main-
land range from 132-144 (mean 139.6). Ventral counts of five fe-
males from the same area range from 136-147 (mean 143.4). Sub-
caudal counts of eight mainland males range from 65-71 (mean
67.8), and those of four mainland females range from 54-62 (mean
58.0). These counts agree well with those presented by Trapido
(1944) and Duellman and Schwartz (1958), with the exception of
the subcaudal counts of males. Both the range (53-73) and the
mean (62.4) given by Duellman and Schwartz indicate counts
lower than those recorded by Trapido and the present author.

The Lower Keys population is similar to that in northern
Florida in ventral and subcaudal counts, as given by Trapido
(1944) and by Duellman and Schwartz (1958), but differs sub-
stantially from that in south Florida, especially in its much lower
subcaudal counts.

As pointed out by Trapido (1944), specimens of S. d. victa from
southern and northern Florida differ in head markings. Specimens
from Gainesville, for example, are characterized by the contrast be-

302 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

tween the pale occipital area and the conspicuous black nuchal
markings. Young individuals from southern Florida are similar, but
most of the adults from that region lose this contrast and possess
virtually concolored heads and necks. Adults from the Lower Keys
are intermediate between typical north and south Florida victa.
All have a pale occipital area and dark neck markings, but these
are not so contrasting as in Gainesville specimens.

It is of interest that Storeria dekayi is distinctly different on the
Lower Keys from the population of the geographically annectant
southern mainland. That it is more similar to populations farther
north may be of significance. This may be a case of the isolation
of a formerly continuous “highland” population and its increasing
separation from the parent stock by the spread of a “lowland” popu-
lation differentiating in the Everglades area. This has been postu-
lated for Coluber constrictor Linnaeus (Auffenberg, 1955) and is
likely for Elaphe obsoleta Say as well. Further study, with atten-
tion to color differences (I have seen no living specimens from
north Florida), may indicate the advisability of recognizing three
races of S. dekayi in peninsular Florida.

A single specimen from Key Largo furnishes the only record for
the species from the Upper Keys. It is a young female (snout-vent
length 125) with a conspicuous head pattern, 144 ventrals, and 50
subcaudals. From these characteristics it is impossible to relate it
definitely to either the Lower Keys or the mainland populations.
S. dekayi may follow the pattern of Diadophis punctatus, Coluber
constrictor, and Elaphe guttata Linnaeus, in which the Upper Keys
populations are more similar to mainland populations than they are
to those on the Lower Keys.

As in Diadophis, individuals of Storeria from the Lower Keys
average smaller than those from the mainland. One male from the
Lower Keys has a total length of 255, and three females from that
area vary from 224-293 in total length. The three largest males and
females examined from the mainland have total lengths of 310-324
and 312-391, respectively.

Specimens examined. Total 22. Fiorma: Alachua County,
Payne’s Prairie (pre 3310; as 90); Broward County, about 4 mi.
NNW Andytown (prp 826, 827, 908); Dade County, 7.6 mi. N
U. S. 41 on Fla. 27 (prep 2465); Miami area (including Coral
Gables) (umrc 55-410, 55-411, 55-414, 55-415, 55-466, 55-469,

PAvuLson: Variation in Some Snakes 303

50-616; pre 1293; pwB 17); west of Princeton (pre 1448); Monroe
County, Key Largo (pre 1427); Big Pine Key (ummz 108211-
108212; pre 3037-3038 ); Sugarloaf Key (umRc 55-409).

Thamnophis sauritus Linnaeus

Duellman and Schwartz (1958) examined a single specimen of
this species from Big Pine Key, the only record for the Keys. Ross-
man (1963), in his revision of the sauritus group of Thamnophis, in-
cluded the Florida Keys in the range of T. s. sackeni Kennicott but
did not further discuss the Keys population. Eight specimens are
now available from Big Pine Key. These include an adult male,
two adult females, and the young of one of the females. The young
were removed from the female in a fully developed state (on 2
July). The female measured 430 in snout-vent length and 258
in tail length. Five of a total of eight young were saved, and four
of these ranged from 134-143 (mean 138.5) in snout-vent length.
The total lengths of three were 214-224 (mean 218.7). These
snakes are considerably smaller than most newborn young examined
by Rossman (1963) from Florida.

Sixteen male and eleven female T. sauritus were examined from
the southern Florida mainland. Ventral counts for these specimens
ranged from 162-172 (mean 166.6) in males and from 155-166
(mean 159.5) in females. Ventral counts of Big Pine Key speci-
mens varied from 162-166 (mean 163.6) in five males and from
156-165 (mean 159.3) in three females. From these data, it can
be seen that little difference exists between these two populations
in this character.

Many specimens of this species lack the tail tip when collected.
Of 35 specimens examined for this study, only 21 could be used
for subcaudal counts, which were made only when the tail ended
in an obvious spine. Subcaudal counts of southern mainland speci-
mens ranged from 123-140 (mean 129.8) in eight males and from
115-134 (mean 122.8) in six females. The same counts for Big Pine
Key specimens ranged from 138-145 (mean 138.6) in five males and
from 127-128 (mean 127.5) in two females.

The population of T. sauritus on Big Pine Key differs from that
of the southern mainland (and those of the remainder of the range
of the species) in its higher subcaudal counts. Rossman (1963)
discussed the somewhat clinal nature of the variation in this char-

304 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

acter, and his illustration of this (Fig. 10, p. 154) shows that sub-
caudals tend to increase toward the south. The condition on Big
Pine Key may be considered the terminus of this cline, although
admittedly disjunct geographically and of a greater magnitude of
difference than that between other annectant populations.

Unfortunately, none of the material from Big Pine Key is ac-
companied by information on color in life. As coloration is of the
greatest significance in diagnosing subspecies of this group, further
discussion of the possible nomenclatorial recognition of the Lower
Keys population must await the collection of additional specimens.
Ribbon snakes are apparently not uncommon on those keys with
permanent fresh water, as I have observed others on Big Pine Key
and found two individuals (both badly smashed road kills) on
Cudjoe Key.

Specimens examined. Total 35. FLorwa: Collier County, 2.3 mi.
W. Carnestown (pRP 513); Deep Lake (umrc 55-541, 55-548);
Broward County, 12.1 mi. NNW Andytown (pre 669); 8.5 mi.
NNW Andytown (pre 668); 6.9 mi. NNW Andytown (prp 667);
4.9 mi. NNW Andytown (pre 671); 4 mi. NNW Andytown (pRP
828-829); Dade County, Hialeah (umrRc 55-524) Coral Gables
(umRC 55-522); Franjo (prep 1221); “Krome Avenue, 10 miles
west of Miami” (umrc 55-523); 19 mi. W and 5.0 mi. S. Miami
(pRP 847); Monroe County, Pinecrest (UMRC 55-508-517, 55-542,
595-556-557 ); Big Pine Key (ummz 11372; pre 640, 1325 and five
young from it. )

DISCUSSION

Fourteen species of snakes have been recorded from the Florida
Keys (Duellman and Schwartz, 1958). Of these, four are restricted
to the Upper Keys and one to the Lower Keys, the rest extending
through both groups of islands. The fourteen include four sub-
species described from the Keys. These are Diadophis punctatus
acricus, Coluber constrictor haasti Bell, Elaphe guttata rosacea
Cope, and Elaphe obsoleta deckerti Brady. Of these, C. c. haasti
has been synonymized with C. c. priapus Dunn and Wood (Auffen-
berg, 1955) and E. g. rosacea and E. o. deckerti with E. g. guttata
Linnaeus and E. o. quadrivittata Holbrook respectively (Duellman
and Schwartz, 1958). Although there is no question that specimens
of Coluber from the Lower Keys can be matched by specimens from
northern Florida and elsewhere in the range of priapus, “haasti” is

PAuLson: Variation in Some Snakes 305

a very homogeneous form, consisting entirely of shiny black snakes
with or without white chins and labials, and never with the paler
bellies and more extensive labial and gular white areas often found
on peninsular priapus. It is also a smaller form, as Bell (1952)
pointed out.

The situation with Elaphe is even more open to debate. Most
specimens of E. guttata from the Lower Keys are easily separable
from most mainland specimens. Even though there is some indi-
cation of a clinal trend in the color variation of this species, the
population heretofore considered an endemic Keys subspecies is a
relatively homogeneous one, and its nomenclatorial status should
not be threatened by the occasional mainland specimens with its
diagnostic characters. The Lower Keys population differs at least
in coloration of adults from all others of the species, and students
of variation and zoogeography should be cognizant of this differ-
ence, even if the name rosacea is not utilized.

The same comments are in order for E. obsoleta. Adult speci-
mens from the Keys are easily distinguishable in life from 90 per
cent or more of specimens of the mainland races quadrivittata or
rossalleni Neill. Apparently the young of all three south Florida
nominate forms are indistinguishable from one another, a condition
which has parallels in other species (e.g., Coluber constrictor).
However, typical adults of each form are recognizable as such, and
these “typical” specimens not only represent the vast majority of
their respective populations but occur over well-defined geographic
and ecologic areas and seem to satisfy the criteria utilized in dis-
tinguishing subspecies. One of the reasons given by Duellman and
Schwartz (1958) against recognizing a race deckerti is that indi-
viduals from the eastern mainland of southern Florida often show
faint dorsal blotches. If the Lower Keys have a recognizable popu-
lation, it might be expected that intergrades between it and rossal-
leni (which occupies the adjacent mainland) would be found iv
that area, as Neill (1949) originally stated.

The controversy about Florida races of these two species of
Elaphe has persisted long enough (Neill, 1949, 1954; Dowling,
1952; Duellman and Schwartz, 1958) to indicate any solution as
subjective. Considering some or all of the named forms invalid
should not obscure the recognition of their value to the student of
biogeography. The latter must depend upon the systematist for

306 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

his basic tools, i.e., morphologically and geographically defined
populations. My subjective interpretation of the situation is that
more is to be gained by the maintenance of the names rosacea and
deckerti for the endemic Keys populations of Elaphe than by their
suppression. Typical rosacea is restricted to the Lower Keys and
intergrades with guttata in the Upper Keys. The fact that rosacea-
like individuals occur on the southern mainland is to me indicative
of an extension of the gene pool of the Lower Keys form northward
after its original isolation in that area. The range of E. o. deckerti
extends at least from Elliott Key (pre 3068) to Grassy Key (not
preserved ).

Of the remaining Keys species, Storeria dekayi, Thamnophis
sauritus, and Micrurus fulvius Linnaeus differ from their nearby
mainland relatives in scale counts. The first two are discussed in
the present paper, and counts listed by Duellman and Schwartz
(1958) for Micrurus from Key Largo show the rather pronounced
difference in ventral counts between specimens from there and the
mainland. Another specimen more recently collected on Key Largo
agrees with the previously cited examples in its low ventral count.

The two specimens of Drymarchon corais Daudin I have seen
from the Lower Keys (both from Big Pine Key) were entirely
black, without any indication of pale gray or pinkish on the head,
as is usual on mainland specimens. Again, more material of this
species from the Keys may show the presence of a recognizable
entity there, but I know of only a few preserved specimens from
that area.

The specimen of Heterodon platyrhinos Latreille from Key
Largo which was discussed by Duellman and Schwartz (1958) has
a low ventral count, but an insufficient series of mainland females
was at hand for comparison. The validity of this record may be
questionable, as Key Largo is a very unlikely locality for a Het-
erodon. The next farthest south definite record for the species is
Homestead, although the indefinite citation “Cape Sable road” may
be still farther south. This species is generally found in sandy
areas, in which it burrows, and I doubt if it would become estab-
lished on Key Largo, where its usual prey (Bufo) is absent. Toads
do occur on the Lower Keys, and Heterodon could survive there,
short of the difficulties encountered in burrowing in limestone
rock.

PAuLson: Variation in Some Snakes 307

The other snakes occurring in the Keys (Opheodrys aestivus
Linnaeus, Tantilla coronata Baird and Girard, Natrix fasciata Lin-
naeus, Agkistrodon piscivorus Lacépéede, and Crotalus adamanteus
Beauvois ) have not been studied adequately to determine whether
they too are differentiated from their mainland relatives. Roger
Conant is studying Natrix fasciata, and Tantilla from Florida have
been examined by Sam R. Telford, Jr., but neither work has been
published as yet. Only a single specimen of T. coronata is avail-
able from the Keys (Key Largo), but adequate material is present
in collections of the other species. Natrix Agkistrodon, and Cro-
talus, with their superior powers of dispersal through coastal
habitats, may avoid differentiation by continuous gene flow. The
first two are aquatic and the last semi-aquatic, often seen swimming
in mangrove-lined canals and even in Florida Bay. Opheodrys
would be worth further study, as it may not disperse so rapidly and
is known from throughout the Keys.

The Keys support endemic races of Kinosternon bauri Garman,
Malaclemys terrapin Schoepff, and Eumeces egregius Baird. As
well, populations of Scaphiopus holbrooki Harlan, Bufo terrestris,
Hyla cinerea Schneider, Gastrophryne carolinensis Holbrook, Rana
pipiens Schreber, Anolis carolinensis Voigt, and Lygosoma laterale
Say differ to some degree from mainland populations (Duellman
and Schwartz, 1958; writer’s observations). Eight of the 14 species
of snakes occurring on the Keys exhibit variation from the norm of
populations on the mainland. It is clear that the isolated nature
of the Keys, especially the lower ones, has served as a mechanism
to promote at least a low order of speciation.

The degree of endemism shown by the entire herpetofauna of
the Keys is striking. Of 30 indigenous species (excluding the
aquatic sea turtles, Malaclemys, Crocodylus, and Natrix), at least
18 (60 per cent) exhibit marked population differences between
the Keys and mainland or between the Upper and Lower Keys.
This high percentage appears to be unique in temperate North
America but may be parallelled or exceeded by some of the
Mexican coastal island groups. In some of the latter cases, how-
ever, the inhabitants of the islands are isolated by considerable
distances of deep water. Florida Bay and most of the channels
between the Keys are shallow, usually protected waters which may
allow relatively rapid dispersal of reptiles and amphibians with a

308 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

lowering of sea level. Additional collecting and further study of
the herpetofauna is imperative before booming development de-
stroys the already limited habitats.

ADDENDA

The proposed union of Diadophis regalis with D. punctatus (Mecham,
1956, Copeia, pp. 51-52; Gehlbach, 1965, Proc. U. S. Nat. Mus., vol. 16, pp.
300-307) was questioned by Myers (1965, Bull. Florida State Mus., vol. 16,
pp. 43-90). This problem does not affect the conclusion of the present paper.

Telford (1966, Bull. Florida State Mus., vol. 10, pp. 261-304) described
the Tantilla from southeastern Florida as a new species, T. oolitica. The single
Key Largo specimen differed in head pattern from all Dade County specimens
examined.

LITERATURE CITED

AUFFENBERG, WALTER. 1955. A reconsideration of the racer, Coluber con-
strictor, in eastern United States. Tulane Stud. Zool., vol. 2, pp. 89-155.

Bett, L. Nem. 1952. A new subspecies of the racer Coluber constrictor.
Herpetologica, vol. 8, p. 21.

BLANCHARD, FRANK N. 1942. The ring-neck snakes, genus Diadophis. Bull.
Chicago Acad. Sci., vol. 7, pp. 1-144.

Dow.Linc, HERNDON G. 1951. A proposed standard system of counting
ventrals in snakes. Brit. Jour. Herpet., vol. 1, pp. 97-99.

— —. 1952. A taxonomic study of the ratsnakes, genus Elaphe Fitzinger.
IV. A check list of the American forms. Occ. Pap. Mus. Zool. Univ.
Michigan, no. 541, pp. 1-12.

DUELLMAN, WILLIAM E., AND ALBERT SCHWaRTZ. 1958. Amphibians and
reptiles of southern Florida. Bull. Florida State Mus., vol 3, pp.
181-324.

NemLL, WitFrepD T. 1949. A newsubspecies of rat snake (genus Elaphe),
and notes on related forms. Herpetologica, vol. 5, 2nd suppl., pp. 1-12.

. 1954. Ranges and taxonomic allocations of amphibians and reptiles
in the southeastern United States. Publ. Res. Div. Ross Allen’s Reptile
Inst., vol. 1, pp. 75-96.

RossMAN, Dovuctas A. 1963. The colubrid snake genus Thamnophis: a
revision of the sauritus group. Bull. Florida State Mus., vol. 7, pp.
99-178.

Trapipo, Haroxtp. 1944. The snakes of the genus Storeria. Amer. Midland
Nat., vol. 31, pp. 1-84.

7450 S. W. 102 Street, Miami, Florida 33156.

Quart. Jour. Florida Acad. Sci. 29(4) 1966 (1968 )

FLORIDA ACADEMY OF SCIENCES

COUNCIL FOR 1966

President: Marcaret L. GILBERT
President Elect: Jackson P. SICcKELS
Secretary: JOHN D. Kixpy
Treasurer: JAMES B. FLEEK
Past President: O. E. Fryr
Past President: Grorcr K. RE
Chairman, Charter and By-Laws Committee: ELMER C. PRICHARD
Finance Committee: JOHN S. Ross
Honors Committee: CLARENCE C. CLARK
Quarterly Journal Committee: I. G. Foster
Talent Search Committee: ALFRED P. MILs
Program Committee: JACKSON P. SICKELS
Editor, Quarterly Journal: Pierce BRODKORB
Chairman, Biological Sciences Section: ELMER C. PRICHARD
Physical Sciences Section: RicHARD E. GARRETT
Social Sciences Section: MERLIN G. Cox
Medical Sciences Section: Mauricrt A. BARTON
Science Teaching Section: ALBERT A. LATINA
Conservation Section: ELBERT A. ScHory, SR.
AAAS Council and Conference Representative: JoHn D. McCRoNE
State Coordinator of the Junior Academy: LoutseE V. AsH
Councilor at Large, Elected: FREDERICK H. BERRY
Councilor at Large, Elected: Witpur F. BLock
Councilor at Large, Appointed: SAMUEL F. CLARK
Councilor at Large, Appointed: Howarp R. BissLAND

ADDITIONAL COMMITTEE CHAIRMEN

Auditing: Joun S. Ross

Awards and Grants: H. K. WALLACE

Future Annual Meetings: MARGARET GILBERT
Local Arrangements: JAMEs D. Ray, Jr.
Membership: JAckson P. SICKELS
Resolutions: O. E. FRYE

310 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Necrology: D.C. Swanson

Nominating: ALFRED P. MILLs

Visiting Scientist Program: CLARENCE C. CLARK
News Letter: ALFRED P. MiLus

MEMBERS OF THE ACADEMY
September 14, 1966

Sectional membership is indicated as follows: B, Biological
Sciences; C, Conservation; M, Medical Sciences; P, Physical Sci-
ences; S, Social Sciences. Members are requested to inform the
Editor of their sectional preferences.

Agens, Frederick F., 406 Mission Hills Avenue, Tampa 10, Florida P
Akin, Benjamin, 1019 N.W. 11th Court, Miami 36, Florida B
Alam, Dr. Syed Qamar, Dept. Home Economics, Florida A & M Univ., Talla-

hassee, Florida S
Alexander, Dr. Taylor R., Dept. Botany, Univ. Miami, Coral Gables, Florida B
Allabough, Edwin D., Jr., 2114 Howard Drive, Orlando, Florida B
Allen, Ross, Ross Allen’s Reptile Inst., P. O. Box 367, Silver Springs, Florida

32688 B
Allen, Dr. Ted T., Jacksonville Univ., Jacksonville, Florida B
Almodovar, Dr. Luis R., Box 286, San German, Puerto Rico B
Alt, David, Dept. Geology, Univ. Montana, Missoula, Montana P
Anderson, William D., Jr., Dept. Biology, Univ. Chattanooga, Chattanooga,

Tennessee 37403 B
Andrews, Donald H., 750 N.E. 33rd Street, Boca Raton, Florida
Arean, Dr. Victor M., Dept. Pathology, Univ. Florida, Gainesville, Florida M
Arnold, Dr. Luther A., 143 Norman Hall, Univ. Florida, Gainesville, Florida B
Ash, Mrs. Louise V., 2405 N.W. 18th Place, Gainesville, Florida T
Ash, Dr. Willard O., 2405 N.W. 18th Place, Gainesville, Florida P
Ashford, Dr. Theodore A., Univ. South Florida, Tampa, Florida B
Auffenberg, Walter, Florida State Museum, Gainesville, Florida B
Averre, Dr. Charles W., III, Sub-Tropical Experiment Station, 18901 S.W. 300

St., Homestead, Florida 33030 B
Baker, Harry J., 6202 Gulf Drive, Panama City, Florida 32402
Baker, Robert A., III, 2004 Romeria, Austin, Texas 78757
Ballard, Dr. Stanley S., Dept. Physics, Univ. Florida, Gainesville, Florida P
Banerjee, Shib Das, 818 W. Pensacola St., Tallahassee, Florida
Banks, Joseph E., 1411 Marion Ave., Tallahassee, Florida
Barkuloo, James M., Fishery Biologist, Game and Fresh Water Fish Com.,
P. O. Box 576, Panama City, Florida 32402 B
Barton, Dr. Maurice A., 1900 Almeria Way S., St. Petersburg, Florida M
Barton, Dr. William K., 861 Sixth Ave. S., St. Petersburg, Florida M
Beck, William M., Jr., 1621 River Bluff Road, Jacksonville 11, Florida B

Membership List 311

Becknell, Dr. G. G., 6900 Dixon Avenue, Tampa 4, Florida P

Beery, Dr. John R., Dean, School of Education, Univ. Miami, Coral Gables,
Florida S

Bellamy, Dr. R. E., 2311 B Street, Bakersfield, California B

Bellamy, Dr. Raymond F., 326 DeSoto, Tallahassee, Florida S

Beller, H. E., 743 duPont Bldg., Miami 32, Florida

Bellomy, Mildred D., 80 E. 53rd Terrace, Hialeah, Florida

Bennett, Frank A., P. O. Box 3, Olustee, Florida

Beohm, E. E., 324 W. 7th St., Jacksonville, Florida 32206

Berne-Allen, Allan, 144 Washington Drive N., St. Armands Key, Sarasota,
Florida 33577

Berry, Frederick H., Bureau of Commercial Fisheries, Tropical Atlantic Biology,
75 Virginia Beach Ave., Miami, Florida 33149 B

Betts, Joseph M., Jr., 406142 Bronough, Tallahassee, Florida 32303

Biber, David, 725 S.W. 4th Ave., Gainesville, Florida 32601

Bieber, Theodore I., Dept. Chemistry, Florida Atlantic Univ., Boca Raton,
Florida P

Bird, L. C., 303 S. 6th Street, Richmond, Virginia B

Birdsey, M. R., Dept. Biology, Miami-Dade Junior College, Miami, Florida B

Bissland, Howard R., 810 N.W. 19th Terrace, Gainesville, Florida C

Blanchard, Dr. Frank N., Dept. Geology, Univ. Florida, Gainesville, Florida P

Block, W. F., 2616 Fairway Ave., S., St. Petersburg, Florida P

Bodman, John W., 18 Wedgemere Ave., Winchester, Massachusetts 01890

Bonninghausen, Russel A., 313 E. Lakeshore Dr., Tallahassee, Florida

Bonniwell, Bernard, Univ. Villanova, Villanova, Penn.

Borders, Huey I., 3400 S.W. 26th St., Fort Lauderdale, Florida

Boss, Dr. Manley L., Florida Atlantic Univ., Boca Raton, Florida B

Boulware, Joe W., Univ. South Florida, Tampa, Florida

Breder, Dr. C. M., Jr., R.F.D. 1, Box 452, Englewood, Florida 33533

Breen, Dr. Ruth S., 1806 Croydon Drive, Tallahassee, Florida 32303

Brey, Dr. Wallace S., Jr., 800 N.W. 37th Drive, Gainesville, Florida P

Brodkorb, Dr. Pierce, Dept. Biology, Univ. Florida, Gainesville, Florida B

Brooker, Dr. Ralph H., Dept. Physics, University of South Florida, Tampa,
Florida P

Brooks, Dr. Karl M., Dept. Psychology, Florida A & M, Tallahassee, Florida
223000 S

Browder, James S., 1018 N.W. 10th Ave., Gainesville, Florida

Brown, Charles P., 641 N.W. 34th Terrace, Gainesville, Florida B

Brown, Dr. Relis B., Dept. Biol. Sci., Florida State Univ., Tallahassee, Florida
32306 B

Brown, William N., 2006 South Fern Circle, Leesburg, Florida

Broyles, Dr. Arthur A., Dept. Physics, Univ. Florida, Gainesville, Florida P

Buckland, Charlotte B., 2623 Herschel St., Jacksonville, Florida B

Buckwalter, Howard M., P. O. Box 926, Homestead, Florida

Bullen, Mrs. Adelaide K., 103 Seagle Bldg., Gainesville, Florida B

Bullen, Ripley P., 103 Seagle Bldg., Gainesville, Florida B

Burgess, William D., Florida College, Tampa, Florida B

312 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Buri, Dr. Peter, Div. Natural Sci., New College, Sarasota, Florida B
Burlingham, Kenneth, 5106 Memory Lane, El] Paso, Texas
Burns, Russell M., Box 900, Marianna, Florida

Burton, R. H., 1301 N.W. 180th Terrace, Miami, Florida

Byrd, I. B., Bureau of Commercial Fisheries, P. O. Box 6245, St. Petersburg
Beach, Florida 33736 B

Byron, C. W., 1034 Samar Road, Cocoa Beach, Florida

Caldwell, Dr. David K., Los Angeles County Museum, Los Angeles, California
90007 B

Calvin, D. Bailey, Univ. Miami Medical School, 1600 N.W. 10th Avenue,
Miami, Florida

Camacho, Dr. E. Oliver, 740 N.E. 181st St., North Miami Beach, Florida

Campbell, James A., 2707 Bronte Ave., Nashville, Tenn. B

Candlin, A. H. S., College Engineering, Embry-Riddle Aeronautical Inst.,
Daytona Beach, Florida P

Carlisle, Dr. Victor W., 223 McCarty Hall, Univ. Florida, Gainesville, Florida
32601 P

Carmichael, Billy E., Chipola Jr. College, Marianna, Florida

Carr, Dr. A. F., Jr., 14 Flint Hall, Univ. Florida, Gainesville, Florida B

Carr, Joseph A., Jr., 3402 Riverview Drive, Tampa, Florida P

Carr, Rebecca Ann, 375 W. 18th St., Apt. 117, Hialeah, Florida

Carr, Dr. T. D., Dept. Physics, Univ. Florida, Gainesville, Florida P

Carroll, Dr. Edward J., 721 Dupont Plaza Center, Miami, Florida

Carter, William J., P. O. Box 247, Yankeetown, Florida 32698

Chalmers, Dr. E. L., Jr., College Arts & Sci., Florida State Univ., Tallahassee,
Florida 32306

Chamelin, Dr. I. M., 615 Phelps Ave., Winter Park, Florida 32789 B

Chapman, H. L., Range Cattle Experiment Station, Ona, Florida 33865 B

Chester, Richard H., Marine Laboratory, 1 Rickenbacker Causeway, Miami,
Florida 33149 B

Chester, Alphonse A., 674 South Lakemont Ave., Winter Park, Florida

Chew, Victor, 380 Greenway Ave., Satellite Beach, Florida 32935

Christensen, Robert F., 19425 N.W. 24th Ave., Opa Locka, Florida

Clapper, Russell B., 2072 Braman St., Fort Myers, Florida 33901

Clark, Dr. Clarence C., Univ. South Florida, Tampa 4, Florida P

Clark, Fred E., Dept. Biology, Stetson Univ., DeLand, Florida B

Clark, Samuel F., Dept. Chemistry, Florida Atlantic Univ., Boca Raton, Florida
P

Clarke, Walter V., 1195 S.E. 17th St., Fort Lauderdale, Florida

Clary, J. D., 1608 Meadowbrook Ave., Lakeland, Florida 33803

Claus, Dr. Edward P., Ferris State College, Big Rapids, Michigan

Clugston, James P., Bur. Commercial Fisheries Biological Lab., Ft. Crockett,
Galveston, Texas 77552 B

Collier, Prof. Albert W., 1622 Seminole Dr., Tallahassee, Florida

Coleman, Sylvia E., Research Lab. 151, Veterans Administration Center, Bay
Pines, Florida 33504 M

Conard, Dr. Henry S., Lake Hamilton, Florida P

Membership List 313

Cope, Paul T., 415 7th St. S., St. Petersburg, Florida
Coulter, Jay W., 10791 S.W. 46th St., Miami, Florida

Cox, Joseph R., Dept. Physics, Florida Atlantic Univ., Boca Raton, Florida
33432 P

Craig, Dr. Palmer H., Dean of Science and Mathematics, Florida Atlantic
University, P. O. Box 430, Boca Raton, Florida P

Craighead, Dr. Frank C., Box 825, Homestead, Florida B

Creager, Don B., Prof. Biology, Jacksonville Univ., Jacksonville 11, Florida B

Crossman, Roy A., Jr., 106 4th St., Jar-Phyl Village, Winter Park, Florida
33880

Crouch, Dr. G. E., Jr., Physics Dept., Florida State University, Tallahassee,
Florida P

Cunha, Dr. T. J., 252 McCarty Hall, Univ. Florida, Gainesville, Florida B

Dambaugh, Dr. Luella N., Univ. Miami, Coral Gables 46, Florida S

Dana, Allan H., 6606 S.W. 60th St., South Miami, Florida

Davis, Fanny-Fern, P. O. Box 475, Valparaiso, Florida 32580

Davis, Dr. George K., Director Biological Sci., 317 Nuclear Sci. Bldg., Gaines-
ville, Florida 32601 B

Davis, Gordon, Motel Farina, 10600 N.W. 27th Ave., Miami, Florida

Davis, Jefferson C., Jr., Dept. Chemistry, Univ. South Florida, Tampa, Florida
33020). P

Davis, Robert H., Dept. Physics, Florida State Univ., Tallahassee, Florida

Davis, Robert L., 11525 82nd Avenue N., Largo, Florida

Dauer, Dr. Maxwell, Dept. Radiology, Univ. Miami, School of Medicine, Miami,
Florida 33136 M

Deichmann, Dr. Wm. B., School of Medicine, Univ. Miami, Coral Gables 34,
Florida M

Denman, Dr. Sidney B., Box 1364, Stetson Univ., DeLand, Florida S$

Dequine, John F., Southern Fish Culturists, Box 251, Leesburg, Florida B

Derr, Dr. Vernon E., 219 N. Lakeland St., Orlando, Florida P

Dew, Dr. Robert J., Jr., Univ. Tampa, Tampa, Florida P

Dickinson, Dr. J. C., Jr., Florida State Museum, Gainesville, Florida B

Dijkman, Dr. Marinus J., 6767 S.W. 112th St., Miami 56, Florida B

Dobkin, Dr. Sheldon, Dept. Biological Sciences, Florida Atlantic University,
Boca Raton, Florida B

Doyle, Dr. Laura M., 365 Hawthorne Avenue, Palo Alto, California

Driver, Paul J., 1347 West 9th St., Jacksonville 9, Florida

Drysdale, Taylor, 5526 Parkdale Drive, Orlando, Florida

Dubbeldam, Dr. Pieter S., 506 Magnolia Ave., Melbourne Beach, Florida

Dudley, Frank M., Div. Physical Sci., Univ. South Florida, Tampa, Florida P

DuMond, Frank V., P. O. Box 246, Goulds, Florida 33170

Dunning, Dr. Wilhelmina F., Dept. Microbiology, Univ. Miami, Coral Gables
46, Florida B

Dyrenforth, Dr. L. Y., 3885 St. Johns Ave., Jacksonville, Florida M

Eddins, W. T., 800 Wickham Road, Melbourne, Florida 32901

Edwards, Dr. Joshua L., Dept. Pathology, Univ. Florida, Gainesville, Florida M

Edwards, Dr. Richard A., Dept. Geology, Univ. Florida, Gainesville, Florida P

314 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Emmerton, Ernest E., 213 N.W. 34th Terr., Gainesville, Florida

Etheridge, Dr. Richard, Dept. Zoology, San Diego State College, San Diego,
California B

Evans, Dr. Elwyn, 500 E. Colonial Drive, Orlando, Florida M
Ewing, Upton C., 362 Minorca Ave., Coral Gables, Florida B
Farrell, Walter M., Dept. Psychology, Univ. Miami, Coral Gables, Florida S

Ferguson, Dr. John C., Dept. Biology, Florida Presbyterian College, St. Peters-
burg, Florida B

Fernandez, Dr. Mario J., 3521 Price Ave., Tampa, Florida P
Field, Dr. Henry, 3551 Main Highway, Coconut Grove 33, Florida S

Finucane, John H., 1320 58th St. S., Gulfport, Florida B

Fleek, Dr. James B., 518 Patricia Lane, Jacksonville Beach, Florida P

Fly, Prof. Lillian, 4060 Battersea Road, Coconut Grove, Florida B

Foote, Dr. Perry A., Coll. Pharmacy, Univ. Florida, Gainesville, Florida P

Foraker, Dr. Alvan G., 800 Miami Road, Jacksonville 7, Florida M

Ford, Dr. Ernest S., Dept. Botany, Univ. Florida, Gainesville, Florida B

Forman, Guy, Dept. Physics, Univ. South Florida, Tampa, Florida P

Foster, Dr. I. G., Florida Presbyterian College, St. Petersburg, Florida P

Foster, Dr. Virginia, Box 87, Pensacola College, Pensacola, Florida

Fox, Dr. Laurette E., P 504, J. Hillis Miller Health Center, Univ. Florida,
Gainesville, Florida M

Fox, Dr. S. W., Institute of Molecular Evolution, Univ. Miami, 521 Anastasia,
Coral Gables, Florida 33134 B

Foxman, David A., 511 Daroco Ave., Coral Gables, Florida 33146

Friedland, Bernard, 1010 Manati Ave., Coral Gables, Florida 33134

Frye, Dr. O. E., Jr., Game and Fresh Water Fish Comm., Tallahassee, Florida
B

Fuller, Dorothy L., P. O. Box 418, DeLand, Florida B

Fyvolent, Joel D., Suite 610, 1 Davis Blvd., Tampa, Florida 33606

Gardner, Elizabeth Ann, Clin. Res. Unit. Nat’ Children’s Cardiac Hospital,
1475 N.W. 12th Ave., Miami, Florida 33136 M

Garrard, Dr. Leon A., 1617 N.W. 10th Terr., Gainesville, Florida B

Garrett, James R., RCA Service Co., Missile Test Proj., Bldg. 989, Mail Unit
811, Patrick AFB, Florida P

Garrett, Richard E., Dept. Physics and Astronomy, Univ. Florida, Gainesville,
Florida P

Gathman, C. A., 501 21st Avenue N., Lake Worth, Florida B

Gilbert, Dr. Margaret, Dept. Biology, Florida Southern College, Lakeland,
Florida B

Gilman, L. C., Dept. Zoology, Univ. Miami, Coral Gables 46, Florida B

Girard, Murray, 5900 Devanshire Blvd., Coral Gables, Florida B

Goethe, C. M., 3731 Tea Street, Sacramento 16, California B

Goin, Dr. Coleman J., Dept. Biology, Univ. Florida, Gainesville, Florida B

Goldsmith, Victor, Oceanographic Institute, Florida State Univ., Tallahassee,
Florida 32306 B

Golightly, Jacob F., Jacksonville Univ., Jacksonville, Florida 32211

Membership List 315

Gordon, Thomas E., Jr., D.D.S., 2517-A E. Colonial Drive, Orlando, Florida
32803 M

Graff, Miss Mary B., Mandarin, Florida S$
Gramling, Dr. L. G., Coll. Pharmacy, Univ. Florida, Gainesville, Florida M

Gray, Oscar S., Gray Industries, Inc., 948 N.W. 44th Street, Fort Lauderdale,
Florida 33309

Gresham, W. B., Jr., P. O. Box 10019, Tampa, Florida 33609

Guedry, Dr. Fred E., Jr., Naval Aerospace Medical Institute, Naval Air Station,
Pensacola, Florida 32512 M

Guilbert, Edward H., Pensacola Junior College, Pensacola, Florida

Gut, H. J., P. O. Box 700, Sanford, Florida B

Haigh, Paul J., Florida Presbyterian College, St. Petersburg, Florida

Haines, Dr. Charles, P. O. Box Drawer A, Belle Glade, Florida

Hall, Everette E., Jr., P. O. Box 12976, Univ. Station, Gainesville, Florida
32601 M

Hanley, James R., Jr., 6446 Anvers Blvd., Jacksonville 10, Florida

Hansen, Dr. Keith L., Biology Dept., Stetson Univ., DeLand, Florida B

Harlow, Richard F., 938 Millard St., Tallahassee, Florida B

Harms, Dr. R. H., Dept. Poultry Sci., Univ. Florida, Gainesville, Florida B

Harrington, Dr. Robert W., Jr., Florida State Board of Health, P. O. Box 308,
Vero Beach, Florida B

Harris, Herbert, Edward Waters College, Jacksonville, Florida P

Hatala, Dr. Robert J., 2167 Vivian Way S., St. Petersburg, Florida

Hausman, Daniel W., 615 Paul Russell Road, Tallahassee, Florida

Heemstra, Phillip C., Box 70, Univ. Miami, Inst. Marine Sci., 1 Rickenbacker
Causeway, Miami, Florida 33149 B

Heinrich, Dr. Edwin P., Orange Park, Florida 32073

Hellwege, Dr. Herbert, Rollins College, Winter Park, Florida P

Hentges, Dr. James F., Jr., 253 McCarty Hall, Univ. Florida, Gainesville,
Florida B

Heuser, Dr. Gustave F., 608 Hillside Drive, Lakeland, Florida

Hill, William H., 325 Poincianna Drive, Birmingham, Alabama B

Hill, William P., Studio 9, 275 W. Magnolia Ave., Merritt Is., Florida

Hilmon, J. B., P. O. Box F, Fort Myers, Florida

Hobbs, Dr. Horton H., Jr., U. S. National Museum, Washington, D.C. B

Holland, Willis A., Jr., 290 N.E. 150th St., North Miami, Florida

Holman, Dr. J. Alan, Dept. Biol. Sci., Illinois State Univ., Normal, Illinois B

Hopkins, E. F., 1207 Biercliff Drive, Orlando, Florida 32806

Houser, James G., Tech. Director, Martin Co., P. O. Box 5837, Orlando,
Florida P

Houtz, Dr. Phillip, P. O. Box 2525, Miami Beach, Florida 33140

Howard, William E. H., Box 399, Florida A & M Univ., Tallahassee, Florida S

Hsia, S. L., Dept. Dermat., Univ. Miami School of Medicine, 1600 N.W. 10th
Ave., Miami, Florida 33136 M

Hubbell, Dr. T. H., Univ. Michigan, Museum of Zoology, Ann Arbor, Mich-
igan B

316 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Hubbs, Dr. Carl L., Scripps Institution of Oceanography, La Jolla, California
B

Huddleston, Carla J., 857 Ridge Road, Ambridge, Pa. 15003
Hughes, Dr. William E., Box 1337, Stetson Univ., DeLand, Florida

Hull, Robert W., Dept. Biol. Sci., Florida State Univ., Tallahassee, Florida B

Hunt, Burton P., Dept. Zoology, Univ. Miami, Coral Gables 46, Florida B

Hunter, Dr. George W., III, Dept. Microbiology, Univ. Florida, Gainesville,
Florida B

Ingle, Robert M., Florida State Board of Conservation, Tallahassee, Florida B

Ivey, Dr. Marvin L., Dept. Natural Sciences, St. Petersburg Junior College,
St. Petersburg, Florida

Jenkins, George L., Dept. Physics, Stetson Univ., DeLand, Florida P

Jerris, S. R., Roehr Products Co., Inc., P. O. Box 960, DeLand, Florida

Jodrey, Louise H., 1901 S. Ocean Blvd., Boca Raton, Florida 33432

Jones, Dr. E. Ruffin, Jr., Dept. Biology, Univ. Florida, Gainesville, Florida B

Jordon, Dr. Juliane, Florida Southern College, Lakeland, Florida

Joyce, Edwin A., Jr., 2800 4th St. S., St. Petersburg, Florida B

Kane, Howard L., 1264 Cleburne Drive, Fort Myers, Florida 33901

Kaplan, Sherman R., 1680 Meridian Ave., Miami Beach, Florida

Katz, Edith E., R. D. 2, Box 788 E, DeLand, Florida 32720

Kendall, Harry W., Dept. Physics, Univ. South Florida, Tampa, Florida P

Kerman, Herbert D., Halipar District Hospital, Daytona Beach, Florida M

Keuper, Dr. Jerome P., Brevard Engineering College, Melbourne, Florida P

Kilby, Dr. John D., Dept. Biology, Univ. Florida, Gainesville, Florida B

Kinser, B. M., P. O. Box 158, Eustis, Florida P

Kinsey, P. E., 1647 Third Ave., N., Jacksonville Beach, Florida

Klaunberg, Dr. Henry J., 1010 Cordova St., Coral Gables, Florida 33134

Knowles, Robert P., 2101 N.W. 25th Ave., Miami, Florida M

Koczy, Dr. F. F., Institute Marine Science, 1 Rickenbacker Causeway, Miami,
Florida 33149 B

Koger, Dr. Marvin, Dept. Animal Science, Univ. Florida, Gainesville, Florida
32601 B

Kolipinski, Dr. Milton C., U. S. Geological Survey, 51 Southwest First Ave.,
Miami, Florida 33130 P

Kornegay, William F., 1408 N.W. 19th Ave., Ocala, Florida

Krivanek, Dr. Jerome O., Univ. South Florida, Tampa, Florida B

Kronsbein, Dr. John, College Engineering, Univ. Florida, Gainesville, Florida
SPARE 1

LaCava, Dr. Frederick W., 264 South Atlantic Ave., Ormond Beach, Florida

Lacy, Dr. Burritt S., 1523 North Drive, Cherokee Park, Sarasota, Florida

Lackey, Dr. James B., Box 497, Melrose, Florida B

Laessle, Dr. Albert M., Dept. Biology, Univ. Florida, Gainesville, Florida B

Larson, Dr. Edward, Dept. Zoology, Univ. Miami, Coral Gables, Florida B

Latham, Dr. James P., Dept. Geography, Florida Atlantic Univ., Boca Raton,
Florida P

Latina, Albert A., 311A Science Bldg., Univ. South Florida, Tampa, Florida B

Lawrence, Dr. John M., Dept. Zoology, Univ. South Florida, Tampa, Florida B

Membership List 317

Laxson, D. D., 231 W. 41st St., Hialeah, Florida

Layne, Dr. James N., Dept. Conservation, Fernow Hall, Cornell Univ., Ithaca,
Nay. B

Leavitt, Dr. Benjamin B., Dept. Biology, Univ. Florida, Gainesville, Florida B

Lee, Mrs. Annette J., P. O. Box 1233, Lake City, Florida

Lee, Clarence E., 2833 N.E. 26th Ave., Lighthouse Point, Pompano Beach,
Florida 33064

Leigh, Dr. W. Henry, Dept. Zoology, Univ. Miami, Coral Gables 46, Florida B

Leto, Frank P., Jr., 4713 Leila Ave., Tampa 11, Florida T

Liedholz, Gerhard A., 6060 S.W. 114th Street, Miami, Florida 33156

Lilly, Dr. John C., 3430 Main Highway, Miami, Florida 33133

Linton, Thomas L., Univ. Georgia, Marine Institute, Sapelo Is., Georgia B

Little, Dr. William Asa, M-304 Med. Sci. Bldg., Univ. Florida, Gainesville,
Florida 32601 M

Long, Dr. William Hendren, Dept. Meteorology, Florida State Univ., Talla-
hassee, Florida 32306 P

Lorz, Dr. Albert, 409 Newell Hall, Univ. Florida, Gainesville, Florida B

Lovell, William V., Route 2, Box 18, Sanford, Florida P

Lutz, Nancy E., Box 114, Mandarin, Florida

McBee, Ethelyne L., 184 E. Duncan St., Columbus, Ohio 43202

McClure, J. S., Jacksonville Univ., Jacksonville, Florida 32211 P

McCrone, Dr. John D., Florida Presbyterian College, St. Petersburg, Florida B

McDarment, Corley P., Route 1, Box 205, Eau Gallie, Florida 32935

Macgowan, Prof. Robert, Florida Southern College, Lakeland, Florida S$

McKenzie, Doris, 1475 N.W. 12th Ave., Miami, Florida 33136

McMahan, Mary Ruth, 6070 6th Ave., North, St. Petersburg, Florida

Man, Eugene H., P. O. Box 8293, Univ. Miami, Coral Gables, Florida

Mann, Dr. Mary Lee, 29 Green Lane, Jamestown, Rhode Island 02835

Marks, Dr. Meyer B., 1680 Meridian Beach, Miami Beach, Florida B

Marlatt, Dr. Robert B., 29985 S.W. 184th Court, Homestead, Florida

Martin, George T., 94 Murray Blvd., Charleston, South Carolina

Martz, Roger A., P. O. Box 1840, Vero Beach, Florida 32960

Meck, Mervin E., M. D., 225 N. Causeway, New Smyrna Beach, Florida M

Meisel, Max, 4444 Post Ave., Miami Beach, Florida 33140

Menzel, Dr. R. W., Oceanographic Institute, Florida State Univ., Tallahassee,
Florida B

Miles, E. P., Computing Center, Florida State Univ., Tallahassee, Florida P

Miller, Dr. E. Morton, 1212 Manati Ave., Coral Gables, Florida

Mills, Alfred P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida P

Miner, Dr. Helen I., 770 Lake Road, Bay Point, Miami, Florida 33137

Minto, Wallace L., 1242 North Palm Ave., Sarasota, Florida

Moe, Martin A., Jr., Board of Conservation, Marine Lab., P. O. Drawer F, St.

Petersburg, Florida B

Montgomery, Harley A., 1700 N.E. 105th St., Miami Shores, Florida

Montgomery, Joseph G., Dept. Biology, Manatee Jr. College, Bradenton, Flor-
ida B

Moore, McDonald, Hampton Jr. College, Ocala, Florida

318 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Morrison, Thomas F., 1491 Summerland Ave., Winter Park, Florida 32789

Morton, Richard K., Jacksonville Univ., Contract Branch, Jacksonville, Florida
SPALL 8)

Moya, Frank, Dept. Anesthesiology, Jackson Memorial Hosp., Miami, Florida
M

Mulson, Joseph F., Dept. Physics, Rollins College, Winter Park, Florida P
Murray, Mary Ruth, 1326 S.W. Ist St., Miami, Florida S

Mustian, William R., Jr., 529 Bonnie Drive, Lakeland, Florida

Nation, Dr. James L., 537 N.W. 35th Terrace, Gainesville, Florida
Neithamer, Richard W., Florida Presbyterian College, St. Petersburg, Florida

Nelson, Dr. Gid E., Jr., Univ. South Florida, Tampa, Florida B

Niven, Dr. Jorma I., 1803 E. Lakeview Ave., Pensacola, Florida

Noble, Dr. Nancy L., 1550 N.W. 10th Ave., Miami, Florida

Nordlie, Dr. Frank G., Dept. Zoology, Univ. Florida, Gainesville, Florida 32601
B

Ober, Lewis D., 1235 N.E. 204th St., North Miami Beach, Florida B

O’Brien, Robert E., P. O. Box 39, Rollins College, Winter Park, Florida

Olle, Miss Esther W., 700 90th St., Surfside 54, Florida B

Olson, Dr. James Allen, Dept. Biochemistry, J. Hillis Miller Health Center,
Univ. Florida, Gainesville, Florida B

Oren, Benjamin G., 1640 Tigertail Ave., Miami, Florida

Orgell, Wallace H., Dept. Biol. Sci., Florida Atlantic Univ., Boca Raton, Flor-
idayoo4o2) B

Orr, Robert K., 725 E. Palmetto St., Lakeland, Florida 33801

Owens, Clarence B., Box 8, Florida A & M Univ., Tallahassee, Florida S

Page, Melvin E., P. O. Box 7068, 2810 First St., North, St. Petersburg, Florida
33734

Palmer, Dr. A. Z., Dept. Animal Husbandry & Nutrition, Univ. Florida, Gaines-
ville, Florida 32601 B

Pan, Huo-Ping, U. S. Dept. Interior, 3700 E. University Ave., Gainesville,
Florida 32601

Park, Dr. Mary Cathryne, 450 Norwood St., Merritt Is., Florida S

Paulson, Dr. Dennis R., 7450 S.W. 102nd St., Miami, Florida

Penner, Dr. Lawrence R., Dept. Zoology, Univ. Connecticut, Storrs, Connecti-
Cut

Pepinsky, Prof. Ray, Physical Science Center, Nova Univ., 440 East Las Olas
Blvd., Fort Lauderdale, Florida P

Phillippy, Clayton, 2202 Lakeland Hills Blvd., Lakeland, Florida

Phipps, Dr. Cecil G., Box 181 A, Tennessee Tech, Cookeville, Tennessee P

Pierce, Dr. E. Lowe, Dept. Biology, Univ. Florida, Gainesville, Florida B

Pirkle, Dr. E. C., 611 N.W. 35th St., Gainesville, Florida P

Plendl, Dr. Hans S., Dept. Physics, Florida State Univ., Tallahassee, Florida P

Plyler, Earle K., Dept. Physics, Florida State Univ., Tallahassee, Florida P

Poitras, Dr. Adrian W., Dept. Biology, Dade County Junior College, Miami,
Florida B

Polskin, Dr. Louis J., 1401 S. Florida Ave., Lakeland, Florida M

Membership List 319

Powell, Dr. Howard B., Dept. Chemistry, Univ. Miami, Coral Gables, Florida
Soloed) /P

Prichard, Elmer C., 605 N. Amelia, DeLand, Florida B

Provost, Dr. Maurice W., Box 308 Vero Beach, Florida B

Puryear, Dr. R. W., Florida Normal and Industrial Memorial College, St.
Augustine, Florida S

Rader, Dr. Luke, 7900 S.W. 104th St., Miami, Florida

Rappenecker, Dr. Caspar, Dept. Geology, Univ. Florida, Gainesville, Florida
S60 e

Ray, Dr. James D., Jr., Univ. South Florida, Tampa, Florida B

Reid, Dr. George K., Dept. Biology, Florida Presbyterian College, St. Peters-
burg, Florida B

Reitz, Dr. J. Wayne, President, Univ. Florida, Gainesville, Florida S

Rhodes, Richard A., I, Dept. Physics and Astronomy, Univ. Florida, Gaines-
ville, Florida P

Rice, Dr. Laurence B., 43 W. Bay Drive, Cocoa Beach, Florida

Rich, Earl R., Dept. Zoology, Univ. Miami, Coral Gables, Florida 33134 B

Riemer, Florence R., 9130 S.W. 100 St., Miami, Florida

Rivas, Luis Rene, Dept. Zoology, Univ. Miami, Coral Gables, Florida B

Roberts, Dr. Leonidas H., Benton 202, Univ. Florida, Gainesville, Florida P

Robins, Dr. C. Richard, Institute Marine Science, Univ. Miami, 1 Rickenbacker
Causeway, Va. Key, Miami 49, Florida B

Rolfs, Herman E., 5513 Merrick Drive, Univ. Health Center, Coral Gables,
Florida 33134 M

Ross, Arnold, American Museum of Natural History, Central Park West at
79th St., New York, N. Y. 10024 P

Ross, Dr. John S., Dept. Physics, Rollins College, Winter Park, Florida P

Rowand, Tom, Route 3, Box 5A8, Lake City, Florida

Ryser, Dr. Phillip W., 1315 N.E. 4th Ave., Fort Lauderdale, Florida 33304

Sachs, K. N., Jr., U. S. Geological Survey, E-214 U. S. Nat. Museum, Wash-
ington 25, D. C. 20242 P

Sall, Walter G., 605 Lincoln Road, Miami Beach, Florida

Sandberg, Dr. Douglas H., Dept. Pediatrics, School of Medicine, 1600 N.W.
10th Ave., Miami, Florida M

Sanders, Dr. Murray, Gray Industries, Inc., 948 N.W. 44th St., Fort Lauder-
dale, Florida 33309 M

Sandstrom, Carl J., Dept. Biology, Rollins College, Box 53, Winter Park, Flor-
ida 32789 B

Saslaw, Dr. Milton S., 1350 N.W. 14th St., Miami, Florida 33125 M

Sauer, Dr. E. G. Franz, Dept. Zoology, Univ. Florida, Gainesville, Florida
32601

Sawyer, Dr. Earl M., Dept. Physics, Univ. Florida, Gainesville, Florida P

Schneider, George H., 2292 S.W. 36th Ave., Miami 45, Florida

Schory, Elbert A., Sr., P. O. Box 1468, Fort Myers, Florida

Schrader, H. W., 105 B Williamson Hall, Univ. Florida, Gainesville, Florida
32601 P

Schricker, John Adams, 1760 Oakhurst Ave., Winter Park, Florida 32789

320 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Schultz, Dr. Harry P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida P
Schwartz, Dr. Albert 10000 S.W. 84th St., Miami, Florida 33143 B
Schwarz, Guenter, Dept. Physics, Florida State Univ., Tallahassee, Florida P

Scolaro, Reginald J., P. O. Box 13374, Broadmoor Sta., New Orleans, La. P

Scott, Bruce Von G., 6201 Chapman Field Drive, Miami, Florida 33156

Scruggs, R. M., North Florida Junior College, Madison, Florida

Seaman, Dr. Irvin, 470 Biltmore Way, Coral Gables, Florida M

Shah, N. S., Research Division, State Research Hosp., Galesburg, Illinois M

Shanor, Leland, Dept. Botany, Univ. Florida, Gainesville, Florida 32601 B

Sherman, Dr. H. B., 410 Howry Ave., DeLand, Florida B

Shirley, Dr. Ray L., Nutrition Lab., Univ. Florida, Gainesville, Florida B

Shor, Prof. Bernice C., Rollins College, Winter Park, Florida 32789

Shuster, Dr. Carl N., 2035—26th Ave., N., St. Petersburg 13, Florida P

Sickels, Dr. Jackson P., 541 San Esteban Ave., Coral Gables 46, Florida P

Sigel, Dr. Michael, Dept. Microbiology, Univ. Miami, School Medicine, Coral
Gables, Florida 33134 M

Simon, Dr. Joseph L., Marine Biol. Lab., Woods Hole, Massachusetts

Simons, Dr. Joseph H., 1122 S.W. 11th Ave., Gainesville, Florida P

Sims, Harold W., Jr., 7542—18th Ave. N., St. Petersburg, Florida B

Sisler, Dr. Harry H., Dept. Chemistry, Univ. Florida, Gainesville, Florida P

Six, Dr. N. F., Jr., Scientific Research Laboratories, Brown Engineering Co.,
P. O. Drawer 917, Huntsville, Alabama P

Slack, Dr. Francis, P. O. Box 818, Hobe Sound, Florida M

Smith, Dr. Alex G., Dept. Physics and Astronomy, Univ. Florida, Gainesville,
Florida P

Smith, Earl D., 2309 Coventry Ave., Lakeland, Florida P

Smith, Dr. Fredrick B., Dept. Soils, Univ. Florida, Gainesville, Florida B

Smith, Marshall E., 418 W. Platt St., Tampa 6, Florida M

Smith, Cmdr. Nathan L., 631 N.W. 34th Dr., Gainesville, Florida P

Soldo, Dr. Anthony T., V. A. Hosp., 1200 Anastasia Ave., Coral Gables, Flor-
ida 33134 M

Sokoloff, Dr. Boris Th., Dept. Biology, Florida Southern College, Lakeland,
Florida B

Soule, Dr. James, 104 McCarty, Univ. Florida, Gainesville, Florida B

South, Dr. D. D., Florida Presbyterian College, St. Petersburg, Florida P

Starn, Charles H., 740 N.W. 65th Ave., Fort Lauderdale, Florida

Steinberg, Dr. Roy H., 615 Bayshore Drive, Warrington, Florida 32507

Stelz, William, 604 Westover St., Lakeland, Florida

Stevens, Marion, 3924 Cleveland St., Hollywood, Florida B

Stevenson, Dr. Henry M., Dept. Biological Sciences, Florida State University,
Tallahassee, Florida B

Stewart, Mrs. Violet N., 445—S8th Ave. N.E., St. Petersburg, Florida

Stubbs, Sidney A., P. O. Box 2066, Houston, Texas

Swann, Maurice E., 3101 W. 13th St. Panama City, Florida P

Swanson, Dr. D. C., Dept. Physics, Univ. Florida, Gainesville, Florida P

Sweigert, Ray L., 2761 E. Vina Del Mar Blvd., St. Petersburg, Beach, Florida
33706

Membership List o2i

Swift, Camm C., Dept. Biol. Sci., Florida State Univ., Tallahassee, Florida
32306 B

Swindell, David E., Jr., 905 E. Park Ave., Tallahassee, Florida B
Taffel, Charles, 826 Evernia St., W. Palm Beach, Florida

Taft, Dr. William H., Univ. South Florida, Tampa, Florida

Tanner, W. Lee, Box 38, Lake Panasoffkee, Florida

Tanner, William F., Dept. Geology, Florida State Univ., Tallahassee, Florida P

Taylor, James R., 10922 52nd Ave. North, St. Petersburg, Florida 33708 B

Taylor, John L., 527 New York Ave., Dunedin, Florida B

Tebeau, C. W., Univ. Miami, 307 Aledo Ave., Coral Gables 46, Florida S$

Thomas, Dr. Dan A., Dean of the Faculty, Jacksonville Univ., Jacksonville 11,
Florida P

Thomas, Lowell P., Marine Laboratory, 1 Rickenbacker Causeway, Virginia
Key, Miami 49, Florida B

Thomas, Richard, 10000 S.W. 84th St., Miami, Florida 33143 B

Ting, Dr. S. V., Citrus Experiment Station, Univ. Florida, Lake Alfred, Florida
33850 B

Tinner, J. C., 1305 N. Montford Avenue, Baltimore, Maryland 21213 P

Tissot, Dr. A. N., Agricultural Exp. Sta., Univ. Florida, Gainesville, Florida B

Tocci, Dr. Paul M., 123 Zamora Ave., Coral Gables, Florida 33134

Torre, Dr. Alberto de la, 2100 S. Miami Ave., Miami, Florida 33129

Totten, Henry R., Dept. Botany, Univ. North Carolina, Box 247, Chapel Hill,
North Carolina B

Toulmin, Dr. Lymon D., Dept. Geology, Florida State Univ., Tallahassee,
Florida P

Tyrone, Victor, Box 1438, St. Petersburg, Florida 33731

Van Vleck, David B., Dept. Zoology, Univ. Miami, Coral Gables, Florida B

Van Wagtendonk, Dr. W. J., Research-Veterans Admin. Hosp., Coral Gables,
Florida 33134 M

Vernon, Dr. Robert O., Box 631, Tallahassee, Florida P

Vestal, Dr. Paul A., Dept. Botany, Rollins College, Winter Park, Florida B

Wadell, Dr. Glenn H., 14130 S.W. 99th Ave., Miami, Florida 33158

Wade, Richard A., Institute of Marine Science, Univ. Miami, Miami, Florida B

Wallace, Dr. H. K., Dept. Biology, Univ. Florida, Gainesville, Florida B

Ward, Dr. Daniel B., 733 S.W. 27th St., Gainesville, Florida B

Warnick, Dr. Alvin C., McCarty Hall, Univ. Florida, Gainesville, Florida B

Warnke, D. A., Oceanographic Institute, Florida State Univ., Tallahassee,
Florida B

Warren, Joseph F., Jr., 6835 9th Ave. North, St. Petersburg, Florida

Webb, Dr. S. David, Florida State Museum, Gainesville, Florida B

Weber, Dr. George F., 1122 S.W. 3rd Ave., Gainesville, Florida B

Webster, W. C., Box 211, Gonzalez, Florida

Weigel, Dr. Robert D., Dept. Biology, Illinois State Univ., Normal, Illinois B

Weisbord, Norman E., Dept. Geology, Florida State University, Tallahassee,
Florida P

Weise, Dr. Emil H., 1360 Avondale Ave., Jacksonville, Florida B

Weise, Gilbert N., 8601 Emerald Isle Circle N., Jacksonville 16, Florida

322 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Weiser, Dr. Josejf, Florida Memorial College, St. Augustine, Florida

Wellman, Wayne E., 967 S.W. 5th St., Miami 36, Florida P and S

Wells, Dr. Harry W., Dept. Biological Sciences, Florida State Univ., Talla-
hassee, Florida B

West, Felicia E., 750 N.W. 20Ist St., Miami, Florida

Westfall, Dr. Minter J., Jr., Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Westgate, Dr. Philip J., Central Florida Experiment Station, P. O. Box 909,
Sanford, Florida B

Wheat, Dr. Myron W., Jr., Dept. Surgery, College Medicine, Univ. Florida,
Gainesville, Florida 32601 M

Whittaker, Edward, 1320 N.W. 14th St., #503, Miami, Florida

Wilcox, Dr. Charles J., Dept. Dairy Sci., Univ. Florida, Gainesville, Florida
SVACO I 183

William, Dr. James H., Route 1, Box 312, Avon Park, Florida S

Williams, Louise Ione, Lakeland High School, Lakeland, Florida B

Williams, Dr. Robert H., Univ. Miami, P. O. Box 8233, Coral Gables 46, Flor-
iday Bb

Wisner, Carl V., Jr., P. O. Box 260, Fort Lauderdale, Florida

Wilson, Druid, E-506, U. S. National Museum, Washington 25, D. C. B

Wilson, Dr. John L., 495 Savannah State College, Savannah, Georgia P

Wiltse, Dr. James C., Route 3, Box 180, Orlando, Florida

Witham, Ross, 10 Lake Point, North River Shores, Stuart, Florida B

Woodburn, Kenneth D., Fla. State Board of Conservation, Marine Lab., P. O.
Drawer F, St. Petersburg, Florida C

Wolfenbarger, Dr. D. O., Sub.-Trop. Exp. Sta., Route 2, Box 508, Homestead,
Florida B

Woolfenden, Dr. Glen E., Dept. Zoology, Univ. South Florida, Tampa, Flor-
ida B

Wright, Shirley Jean, P. O. Box 4552, Miami Beach, Florida

Yaffa, Harold, 1466 Bradley Ave., Camden 3, New Jersey B

Yakaitis, Dr. Albina, Dept. Anatomy, Univ. Miami, Coral Gables 34, Florida

Yerger, Dr. Ralph W., Dept. Zoology, Florida State University, Tallahassee,
Florida B

Young, Dr. Harold, P. O. Box 539, Monticello, Florida

Zeppa, Robert, 1700 N.W. 10th Ave., Miami, Florida 33136

Zinner, Dr. Doran D., 2017 Alhambra Circle, Coral Gables, Florida B

Zinober, Dr. M. R., 2105 Friley Road, Ames, Iowa 50012

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1966

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Presbyterian College
Florida Southern College
Florida State University
Jacksonville University
Marymount College
Miami-Dade Junior College
Mound Park Hospital Foundation
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida

University of Tampa

Pal
‘e*

FLORIDA ACADEMY OF SCIENCES _
Founded 1936

OFFICERS FOR 1966

President: MaArcARET GILBERT
Department of Biology, Florida Southern College
Lakeland, Florida

President Elect: JAcKson P. SICKELS
Department of Chemistry, University of Miami
Coral Gables, Florida

Secretary: Joun D. Kirtsy
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: JAMES B. FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: PrerceE BRODKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges
should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

iT%

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

VOLUME 30

Editor
PIERCE BRODKORB

Published by the

FLortipA ACADEMY OF SCIENCES
Gainesville, Florida
1967

PUBLICATION DATES OF VOLUME 30

NumBer 1: April 16, 1968
NuMBER 2: July 24, 1968
NuMBER 3: September 20, 1968
NuMBER 4: December 24, 1968

New TAxa PROPOSED IN VOLUME 30

Cleidodiscus georgiensis Price (Trematoda: Dactylogyridae)
Dactylogyrus jaini Price (Trematoda: Dactylogyridae )
Centrolenella taylori Goin (Amphibia: Centrolenidae )
tAcris barbouri Holman (Amphibia: Hylidae)

tHyla miofloridana Holman (Amphibia: Hylidae)

tF ossil

64
Vat
115
128
132

CONTENTS OF VOLUME 30

NUMBER 1

Interferometry of Jupiter at 18 Mc/s
Jorge May and Thomas D. Carr

Effect of Hurricane Betsy on the southeastern Everglades
is Taylor R. Alexander

Fishes of the St. Johns River, Florida Marlin E. Tagatz

Composition and feed value of shrimp meal W. G. Kirk,
R. L. Shirley, J. F. Easley, and F. M. Peacock

Animal remains from a midden at Fort Walton Beach
Elizabeth S. Wing

Fossil vertebrates from Navassa Island, W.I. Thomas H. Patton

Notes on the trematode genera Cleidodiscus and Urocleidus

C. E. Price

Aerial respiration in the Florida spotted gar Brian McCormack

Invertebrates found in water hyacinth mats James O'Hara
NUMBER 2

Rainfall distribution in the Miami area Louis C. Sass

Two ancient Florida dugout canoes
Ripley P. Bullen and Harold K. Brooks

Variation in plantar tubercles in Peromyscus polionotus
Michael H. Smith

A new gill trematode from Georgia Charles E. Price
A new centrolenid frog from Guyana Coleman J. Goin
Behavior of iguana during rain storms Coleman J. Goin
Additional Miocene anurans from Florida J. Alan Holman

Variation in bovine cholinesterase levels
C. J. Wilcox and H. H. Head

Officers and members of the Academy

iii

81

97

108
et
115
119
121

141
145

NUMBER 3

Differential thermal analysis of wavellite Frank N. Blanchard
Effects of gamma irradiation on prickly pear cactus Carl D. Monk
Notes on Balanus humilis Conrad, 1846 Arnold Ross
Spawning season and sex ratio of echinoids John W. Brookbank

Sleep behavior in frogs
J. Allan Hobson, Coleman J. Goin, and Olive B. Goin

New records of fishes from the western Caribbean
Ray S. Birdsong and Alan R. Emery

Cuban lizards of the genus Chamaeleolis
Orlando H. Garrido and Albert Schwartz

Natural factors affecting deer movements
Richard F. Harlow and William F. Oliver, Jr.

Round-tailed muskrat in west-central Florida John R. Paul

Mating behavior of Peromyscus polionotus Michael H. Smith
NuMBER 4

Effectiveness of family psychotherapy Sidney B. Denman

Morphological variations of Gymnodinium breve in situ
Alexander Dragovich

Chinsegut Hill-McCarty Woods, Hernando County, Florida
Stephen L. Beckwith

Relationship of sand pine scrub to former shore lines A. M. Laessle
Sea turtle nest survey of Hutchinson Island, Florida Robert A. Routa

Yolk pigmenting value of dried kenaf tops
Jack L. Fry, George M. Herrick, and R. H. Harms

Persistent viremia in EIA infected horses
J. H. Flynn, G. H. Waddell, and V. R. Saurino

Temperature and humidity effects on survival of chickens
R. W. Dorminey, H. R. Wilson, I. J. Ross, and J. E. Jones

iv

161
168
173
WET

184

187

197

221
227

230

295

301

316

Quarterly Journal
of the

Florida Academy of Sciences

Vol. 30 December, 1967 No. 4

CONTENTS

Effectiveness of family psychotherapy Sidney B. Denman

Morphological variations of Gymnodinium breve in situ
Alexander Dragovich

Chinsegut Hill-McCarty Woods, Hernando County, Florida
Stephen L. Beckwith

Relationship of sand pine scrub to former shore lines A. M. Laessle
Sea turtle nest survey of Hutchinson Island, Florida Robert A. Routa

Yolk pigmenting value of dried kenaf tops
Jack L. Fry, George M. Herrick, and R. H. Harms

Persistent viremia in EIA infected horses
J. H. Flynn, G. H. Waddell, and V. R. Saurino

Temperature and humidity effects on survival of chickens
R. W. Dorminey, H. R. Wilson, I. J. Ross, and J. E. Jones

Mailed December 24, 1968

241

245

250
269
287

295

301

316

QUARTERLY JOURNAL OF THE FLORA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Mares should be 1% inches all around.

TirLes should not exceed 55 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and
AvpREss in paragraph form following Literature Cited.

LITERATURE CrTeED follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLEs are charged to authors at $19.95 per page or fraction. Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

LEcENDs for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($17.30 per page, $15.80 per half
page. Drawincs should be in India ink, or good board or drafting paper, and
lettered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. Do not submit illustrations needing reduction by more than one-half.
PHotocrapPus should be of good contrast, on glossy paper. Do not write
heavily on the backs of photographs.

ProoF must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FRORIDA ACADEMY OF SCIENCES

Vol. 30 December, 1967 No. 4

Effectiveness of Family Psychotherapy
SIDNEY B. DENMAN

THE literature dealing with the etiology of mental illness gen-
erally affirms that in many psychiatric disorders the family rela-
tionships of the patient are of major significance. As a conse-
quence of this theoretical position, family psychotherapy has been
extensively utilized as one of the methods of treatment for psy-
chiatric patients.

If it is true that family relationships are significant factors
in the development of psychiatric disorders, and that family psy-
chotherapy is an effective therapeutic agent, one might expect to
find a significant difference in the level of post-hospital adjustment
between those patients who receive extensive family psychotherapy
and those who receive little or no family psychotherapy. A search
of the literature resulted in finding only two references, (Sampson,
Messinger, and Towne, 1962) and (Glasser, 1963), which were
directly related to studying the empirical results of family therapy
efforts. In an effort to learn more about this significant area, this
research attempts to learn about the relationship between the
quantity of family psychotherapy and the reported level of adjust-
ment of discharged psychiatric patients.

SUBJECT GROUP

The subject population for this study consisted of 251 patients
who were consecutively discharged between January 1, 1965, and
December 31, 1966, from a small voluntary psychiatric inpatient
unit with a capacity of 32 beds and an average daily census of 27
patients. Inasmuch as this psychiatric unit is a part of a univer-
sity medical center, no claim is made that patients discharged

242, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

from this unit are typical of the American psychiatric patient popu-
lation.

Data SOURCES

Data for this study were obtained from two major sources.
Hospital records on all 251 patients provided information on age,
sex, referral source, financial status, diagnosis, and length of hos-
pitalization. Other data were obtained by a questionnaire mailed
to each discharged patient. At the time the questionnaire was
sent, these patients had been discharged for an average of ap-
proximately one year and ranged in post-discharge time from 3 to
24 months. Of the 251 questionnaires mailed, 166 or 66 per cent
were completed.

A word of caution must be inserted about the use of a ques-
tionnaire in obtaining follow-up data on psychiatric patients.
Ideally, one would like to have a more systematic evaluation of
post-hospital adjustment of patients made by a team of psychia-
trists, psychologists, and psychiatric social workers to avoid possible
errors in patient self-report on level of post-hospital adjustment.
Such an ideal research situation was impossible under the finan-
cial limitations of this research effort. There was no alternative
other than to supplement hospital data with a mailed question-
naire.

METHODOLOGY

In an effort to learn about the influence of variations in the
quantity of family psychotherapy on reported post-hospital ad-
justment, two groups of discharged patients were selected. One
consisted of 62 patients whose families participated in two or
fewer family therapy sessions, and a second group consisted of 70
patients whose families participated in ten or more sessions of
family therapy during the patient’s hospitalization. The patient
composition of these two groups did not differ at the .05 level of
significance in socio-economic status, sex, referral source, or diag-
nosis.

The level of post-hospital adjustment for these discharged pa-
tients was determined by examining the patients’ responses to
questionnaire items concerning level of personal adjustment, em-

(ce)

DENMAN: Family Psychotherapy 24

ployment capability, and the status of their present family rela-
tionships. The questionnaire consisted of a Likert-type rating
scale of five alternative responses along a continuum extending
from “very much better” to “very much worse”. Patient responses
to the categories of “very much better” and “somewhat better”
were combined to serve as a measure of an improved level of
adjustment. The three categories of “about the same”, “somewhat
worse’, and “very much worse’, were combined to serve as an
indicator of a lack of improvement in post-hospital level of adjust-
ment.

FINDINGS

The majority of the discharged psychiatric patients included in
these two groups reported themselves as functioning at a higher
level in terms of personal adjustment, employment capability,
and family relationships than they were during the three months
prior to hospitalization. While 77 per cent reported themselves
as functioning “very much better” or “somewhat better”, only 23
per cent reported themselves as “about the same”, “somewhat
worse , and “very much worse’.

When the group receiving ten or more sessions of family therapy
was compared with the group receiving two and fewer sessions,
no differences at the .05 level of significance were found in re-
ported level of post-hospital functioning in terms of personal ad-
justment, employment capability, or personal relationships with
their families. Patients in the group receiving more family therapy
did not report themselves as functioning significantly any better
than the group receiving little or no family therapy.

IMPLICATIONS

The failure of this research to find a positive relationship be-
tween the quantity of family therapy and a higher level of re-
ported post-hospital adjustment for discharged psychiatric patients
was unexpected. It had been assumed that a larger quantity of
psychotherapy with the families of patients would make increased
contributions to patient improvement.

This study was limited to seeking to learn of any possible re-
lationship between the number of family therapy sessions and the

244 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

level of reported post-hospital adjustment. No effort was made to
measure any possible relationship between post-hospital adjust-
ment and other variables. All that can be reported here is that
no positive relationship was found between the number of sessions
of family therapy and an improved level of reported post-hospital
adjustment for these discharged psychiatric patients.

LITERATURE CITED

GuasserR, P. H. 1963. “Changes in Family Equilibrium During Psycho-
therapy.” Family Process, vol. 2, pp. 245-264.

SAMPSON, H., S. I. MESSINGER, AND R. D. Towner. 1962. “The Mental
Hospital and Family Adaptations.” Psychiatric Quarterly, vol. 36,
pp. 704-719.

Department of Psychiatry, University of Florida College of
Medicine, Gainesville, Florida 32601.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968)

Morphological Variations of Gymnodinium breve Davis

ALEXANDER [DRAGOVICH

VariABILiTy in dinoflagellates reflects internal processes and
external environment and is, therefore, useful in both taxonomic
and ecological studies (Bursa, 1963). Deformed Gymnodinioideae
in laboratory cultures were noted by Kofoid and: Swezy (1921),
Lebour (1925), and Biecheler (1952). Morphological variations
of dinoflagellates under natural conditions, often induced by chang-
es in salinity and temperature or by hydrostatic pressure, occur
frequently in the vicinity of estuaries (Bursa, 1963). This paper
describes several variant forms of the Florida red-tide organism,
Gymnodinium breve, observed in field samples (Fig. 1).

The fish-killing Florida red tide is a natural phenomenon in the
Gulf of Mexico associated with discolored water containing dense
populations of G. breve. Gymnodinium breve also occurs in Trini-
dad (Lackey, 1956) and is the chief suspect related to fish mortal-
ities that occur frequently in the Orinoco River (Gulf of Paria)
(Rounsefell and Nelson, 1966).

Published material on morphological variations in G. breve is
scarce. Since 1948, when Davis described this naked dinoflagellate
as a new species, few publications have been concerned with its
morphology. Lackey and Hynes (1955) gave descriptions of G.
breve; Steidinger (1964) published two photographs of G. breve
(from living specimens); and Steidinger and Williams (1964) de-
fined a new variation of G. breve and discussed cyst formation.
Variation in cultured forms was reported by Wilson (1967); he
also described the most common encystment stage and the general
features of reproduction.

This study is based on plankton samples collected monthly from
February 1964 through February 1965 at 6 locations in Tampa Bay
and Charlotte Harbor (Florida), and 16 locations in neritic waters
of the Gulf of Mexico (Dragovich and Kelly, 1966). The neritic
stations were positioned at 5, 10, 15, and 20 miles offshore along a
series of 4 transects perpendicular to the coastline. At each station
the water column was sampled at 5-m intervals from surface to
bottom. Station depth varied from 5-20 m.

The five morphological variations most frequently observed dur-
ing this study are described in this paper, under the arbitrary des-

246 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Fig. 1. Sketches of morphological variations of Gymnodinium breve.

ignations of forms A, B, DF, C, and D (Fig. 1). All drawings were
made from living specimens.

Form A is a wide, flat cell; breadth varies from 20u to 40u. The
cell is several times broader than long. The breadth referred to
throughout this paper is the breadth across the girdle, ventral view.

DracovicH: Variation in Gymnodinium 247

Breadth is used instead of length because it is more clearly defined
in G. breve than the length.

The cytoplasm is transparent, finely granulated, and without
distinct chromatophores; in transmitted light appears pale green in
color. A large nucleus is usually positioned near the center, all cells
have a small apical protuberance of epicone, and a transverse
groove, or girdle, is centrally located. The sulcus extends into the
epicone, and both girdle and sulcus appear smooth, though slightly
impressed. The organism is equipped with a transverse and a lon-
gitudinal flagellum. The transverse flagellum is within the girdle and
circumscribes the cell in this form and in all other forms. No in-
gested food was noted.

The shape of form B is similar to that of form A but has the
appearance of being slightly deformed and inflated. The breadth
of the cell varies from 25, to 60, and is several times greater than
the length. Numerous round and discoid chromatophores are pres-
ent. The clarity of the cell varies from translucent to opaque, and
the appearance of the cytoplasm in transmitted light varies from
light green to yellowish brown. The nucleus, epicone, sulcus, and
girdle are more pronounced in form B than in form A; the flagella
are the same as in A. No ingested food was observed.

Forms classified as DF are probably predivision stages of form
B, but the actual division process was not observed. Form DF is
generally larger than form B and measures up to 70» wide. The
cell is butterfly-shaped, and some cells have a split along the sulcus
up to the epicone.

The shape of form C is suboval, and the cell breadth varies
from 18 to 40u. The cell may contain numerous oval, suboval, and
discoid chromatophores or may be without distinct chromatophores.
Cell color usually is pale green or yellowish green, and the cell may
be transparent, translucent, or opaque. The distinct nucleus is
located centrally; in some cells, it is contained entirely in the hypo-
cone. The epicone is smaller than the hypocone in some speci-
mens, and the apical protuberance of the epicone was not observed
in all cells. The sulcus (which extends into the epicone and at
times branches) and girdle are pronounced and impressed. Cells
have flagella as do forms A and B. No ingested food was noted.
Form C was similar to the typical mature G. breve cell in cultures
described by Wilson (1967).

248 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Form D is intermediate between forms B and C, but is neither
as wide as form B nor as flat as form C. The apical protuberance
of the epicone and the nucleus, chromatophores, sulcus, girdle,
flagellae, and color are similar to form B.

The described forms occurred at all depths; at times, all forms
were present in single samples. Although similar forms were ob-
served in earlier investigations, records of their occurrence in the
field samples were made for the first time in the present study.

Form C occurred more frequently than the others and was the
only one found in Charlotte Harbor (Florida). Studies have dis-
closed occurrences of form B during bloom and nonbloom periods
of red tide. Form A occurred in the offshore waters through most
of the year, but was never seen in bloom proportions. Form D was
present throughout the area, but was less frequent than forms A, B,
and C.

From an ecological point of view, form A may indicate unfav-
orable conditions for the development of red tide, and form B (and
DF) favorable conditions. Forms C and D may be regarded as
ecologically intermediate forms between A and B (and DF).

Culturing of these forms from single cell isolates will be re-
quired to determine whether they are the same species. Observa-
tions of unialgal cultures of G. breve have shown that forms B and
DF may transform into type C cells (John H. Finucane, personal
communication ). Thus it is possible that these forms belong to the
same species. Wilson (1967) reported that some of the irregularly-
shaped G. breve cells reverted to normal cells when transferred
individually with a micropipette to fresh medium.

The facts that forms observed under natural conditions and in
culture differed from the original description (Davis, 1948) sug-
gest that further and more detailed research on taxonomy and
ecology of G. breve is needed.

LITERATURE CITED

BIrECHELER, B. 1952. Recherches sur les peridiniens. Bull. Biol. France et
Belgique, vol. 36, pp. 2-149.

Bursa, A. S. 1963. Some morphogenetic factors in taxonomy of dinoflagellates.
Fish. Res. Board Canada Studies, No. 798, pp. 55-66.

Davis, C. C. 1948. Gymnodinium brevis sp. nov., a cause of discolored water
and animal mortality in the Gulf of Mexico. Bot. Gaz., vol. 109 pp.
358-360.

DracovicH: Variation in Gymnodinium 249

DracovicH. A., AND J. A. Ketuiy, Jr. 1966. Distribution and occurrence of
Gymnodinium breve on the west coast of Florida, 1964-65. U.S. Fish
Wildl. Serv., Spec. Sci. Rep. Fish. 541, 15 pp.

Koror, C. A., AND O. Swezy. 1921. The free-living unarmoured dinoflagellata.
Mem. Univ. California, vol. 5, 562 pp.

Lackey, J. B., AND J. A. Hynes. 1955. The Florida Gulf coast red tide. Eng.
Prog. Univ. Florida, no. 9, 24 pp.

Lackey, J. B. 1956. Known geographic range of Gymnodinium brevis Davis.
Quart. Jour. Florida Acad. Sci., vol. 19, 71 pp.

Lresour, M. V. 1925. The dinoflagellates of northern seas. Mar. Biol. Assn.
U.K., Plymouth, 250 pp.

ROUNSEFELL, G. A., AND W. R. NELSON. 1966. Red-tide research summarized
to 1964 including an annotated bibliography. U. S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 535, 85 pp.

STEIpINGER, K. A. 1964. Gymnodinium breve Davis. Florida Board Conserv.
Leafl. Ser. Plankton 1, 2 pp.

STEIDINGER, K. A., AND J. WituiaMs. 1964. Gymnodinium breve Davis. Fla.
Board Conserv. Leafl. Ser. Plankton 1( 1a), 2 pp.

Witson, WiLLiaAM B. 1967. Forms of the dinoflagellate Gymnodinium breve
Davis in cultures. Contr. in Mar. Sci., vol. 12, pp. 120-134.

Bureau of Commercial Fisheries Biological Laboratory, St. Pet-
ersburg Beach, Florida. Contribution No. 26.

Quart. Jour. Florida Acad. Sci. 30(4) 1967( 1968 )

Chinsegut Hill-McCarty Woods, Hernando County, Florida
STEPHEN L, BECKWITH

THE name Chinsegut Hill holds a certain fascination to resi-
dents and travelers alike in the vicinity of Inverness and Brooks-
ville, two towns situated in west-peninsular Florida along Highway
U. S. 41. It is the present home of the Beef Cattle Research
Station, Agricultural Research Service, U. S. Department of Agri-
culture. In the past it was designated as a National Wildlife
Refuge, and before that it was the home of Colonel and Mrs.
Raymond Robins. The present report is concerned with a general
survey from September 1965 through July 1966 of a five-acre tract
owned by The Nature Conservancy in an area known as McCarty
Woods immediately adjacent to the Chinsegut Hill tract itself.

Grateful acknowledgment is made to The Nature Conservancy,
particularly Dr. Walter S. Boardman, for assistance in making this
study possible. The author is also deeply indebted to Dr. Dan
Ward, Mr. John Beckner, and Mr. William G. D’Arcy of the Uni-
versity of Florida Herbarium for aid in identifying plants. Scien-
tific names of plants are in accord with Radford et al. (1964) or
Small (1933); common names of plants are derived from the above
or from Kelsey and Dayton (1942). Common and scientific
names of vertebrates are from Blair et al. (1968).

LOCATION AND HisroricAL BACKGROUND

The Chinsegut Hill-McCarty Woods area is located in Hernan-
do County six miles north of Brooksville, or about 50 miles north
of Tampa. This county was first known as Benton County in 1843,
but its name was changed to Hernando County a few years later,
and its present boundaries were established in 1887. Early home-
steaders came largely from Georgia (Jones and Morrison, 1915).

What is now known as Chinsegut Hill was first owned by
Colonel Pearson of Columbia, South Carolina, in 1842. In 1852
it was acquired by Colonel Francis N. Ederington, and in 1904
Colonel Raymond Robins purchased this tract from the grand-
children of Colonel Ederington. The name “Chinsegut” Hill,
given to it by Colonel Robins, was derived from the Innuit Indian
Tribe of Alaska and means “The Spirit of Lost Things.” Colonel
Robins was a colorful figure in diplomatic and economic circles

BeckwitH: Chinsegut Hill-McCarty Woods 251

while Mrs. Robins (the former Margaret Drier) was noted pri-
marily for her cultural and social activities. The hospitality and
generosity of Colonel and Mrs. Robins became almost legendary
and brought Chinsegut Hill its fame (Treiman, undated).

One of the outstanding features of Chinsegut Hill is a tract
of about 400 acres of old growth, “virgin” longleaf pine, which
represents the only remaining sizeable acreage of such trees in
Florida. (See Figs. 1 and 2.) Colonel and Mrs. Robins’ interest
in preserving this magnificent stand of trees and at the same time
in providing a location for investigating cattle raising in west-
central Florida, which they rightly foresaw as being of great eco-
nomic importance, caused them to transfer ownership of their
lands to the Federal Government in 1932. They were given “For
maintenance of a wildlife refuge, forest preserve and experiment
station’ (Miss Lisa von Borowski, pers. com.). Mrs. Robins
passed on in 1945 and the Colonel in 1954, but the name Chinse-
gut Hill lingers on about the Brooksville area.

In order to perpetuate in its natural state a portion of the virgin
stand of longleaf pine in the Chinsegut Hill area, in 1965 The
Nature Conservancy acquired title to a five-acre tract bordering
the western boundary of Chinsegut Hill itself in what is known as
McCarty Woods. This tract was originally owned by Colonel
Robins, but was omitted in the 1932 deed to the Government be-
cause of some doubt as to its title.

Stupy TECHNIQUES
Vegetation Analyses

Vegetation sampling stations marked with aluminum stakes
were located every 100 feet along two lines 100 feet apart, 10
stations per line (Fig. 3). At each station trees were randomly
selected for study by the point-centered quarter method described
by Cottam and Curtis (1956). This method selects sample trees
in each quadrant about a station depending upon their proximity
to the center point. Since there were 20 grid points and 4 quad-
rants per point, this technique resulted in a random sample of 80
trees. A tree was designated as having a dbh (diameter at breast
height) of 3.0 inches or more. Tree diameters were taken with
a diameter tape, and heights were measured with an Abney level.

252 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

|. Cee —
Fig. 1. Growth form of “virgin” longleaf pine in a roadside park north of

Inverness (Citrus County), Florida. Note the flattened, spreading crown and
the abundance of Spanish moss (Tilandsia usneoides), an air plant common

in this locality.

BeckwitH: Chinsegut Hill-McCarty Woods 253

Fig. 2. Longleaf pine stand with dense understory of sweetgum, water
oak, and similar species resulting from fire exclusion for at least 34 years.

Ree ae “ANE Cor.
& Sec, 3-22-19
20
|
|
|@19 . @2
| A
|
ie Generalized Map of
| CHINSEGUT HILL -MCCARTY WOODS
| @17 é
| Hernando County, Florida
rere Ses Sec. 3 Ie22 Sian 9 E
|
rae Scale — 1: 1800
eee!
| 100 0 100
Beer
L754
| Legend
eae —-—- Boundary
Bis —*— Fence line
Se -156- Contour line

® Observation station

Note: Elevations (feet MSL) measured
by Abney level after estimating
160-foot contour from U. S. Geological
Survey map.

Fig. 3. Map of the Chinsegut Hill-McCarty Woods area showing gen-
eralized contours and locations of stations used in vegetation studies.

BecxwitH: Chinsegut Hill-McCarty Woods 25}

OU

In addition to trees sampled as described above other hard-
woods were also measured as they were encountered. These were
particularly large or tall individuals that were missed in the ran-
dom sample. Such observations provided supplementary data on
trees present on the study area. Nineteen other trees were meas-
ured in this manner.

Because of the interest in the longleaf pine on this tract addi-
tional trees of this species were also measured. Increment borings
were taken from representative trees as they occurred in each
quadrant to obtain information on growth rate. Eighteen trees
were bored for this purpose, and, of these, nine were bored a
second time for determining ages. Seven years were allowed to
correct age at breast height to actual age.

Ground vegetation, up to a height of four and one-half feet,
including grasses, herbs, shrubs, and trees less than 3.0 inches dbh,
was evaluated at 10 of the stations used in the point-centered quar-
ter method described above, five stations being randomly selected
from each of the two lines of points. At each station so chosen
three random points were located within a radius of 50 feet.
Clusters of three plots measuring 0.5 < 2.0 meters were then laid
down near each point, one plot at the point, a second plot two
meters west, and a third plot two meters east. There were thus
nine plots located about each station, or a total of 90 plots on
which measurements were made.

Plant abundances were estimated ocularly as the percentage of
the total area covered by each plant. Indices to express the amount
of area covered were the following:

Index Amount of Area Covered
‘iF Less than 1.0%
1.0 to 20.0%
20.1 to 40.0%
40.1 to 60.0%
60.1 to 80.0%
80.1 to 100%

Ok WW

The final coverage index for a species was obtained by adding
all the indices for it on the various plots and dividing by the num-
ber of plots on which it occurred. In the vegetative analyses, “T”
designated a coverage of less than 1.0 per cent; in the final sum-

256 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

mary “t meant that the average coverage index was less than 0.1.
Frequencies of occurrence were also calculated to show the distri-
bution of each plant over the study area. Abundance data for the
various plants thus included both a coverage index and a frequency
index expressed as a percentage.

Soil Analysis

An extension type auger was used in obtaining samples of the
soil present on the McCarty Woods Tract. Due to the difficulty
in boring through the thick, plastic clay and the coarse pulverized
limestone it was impractical to go much below a foot or two in
depth. However, one boring was made down to the 5-foot level.
Nine soil samples were taken in all.

Bird and Mammal Observations

Rodents were sampled with Sherman live traps measuring
3 x 3 x 9 inches. These were set out at stations one chain apart
along each line of vegetation sampling plots. There were 13 sta-
tions in each line and two traps per station, for a total of 52 traps
on the study area. The trapping period was the nights of Sep-
tember 18 and 19, 1965. A mixture of rolled oats and sunflower
seed was used as bait. All rodents captured were identified, aged
and sexed, and marked by toe clipping prior to release.

Abundance data for birds and mammals other than rodents
were recorded only on an incidental basis. For this reason it is pos-
sible to only list species encountered rather than attempt any
quantitative estimates of relative numbers.

DESCRIPTION OF THE CHINSEGUT HitL-McCarry Woops
Climate

The Chinsegut Hill area is characterized by a subtropical cli-
mate with a mean annual temperature near 72 F and an average
annual precipitation of about 58 inches (U. S. Department of
Commerce, 1966). Temperature extremes range from the low 20's
(usually in January) to the upper 90’s in May or June. November,
with an average of only 1.84 inches of rainfall, is the driest month
of the year. The wettest months are usually June, July, and
August each with about 8 inches or more of precipitation. Oc-

BeckwitH: Chinsegut Hill-McCarty Woods PABST

casionally as much as 15 inches of rain may occur in one of the
summer months.
Geology and Topography

Terrain in the Chinsegut Hill area is quite rolling by contrast
to many sections of Florida. Cooke (1945) describes this region
as part of the Central Highlands, which extends down the middle
of peninsular Florida southward almost to Glades County. In the
Brooksville area these Highlands are underlain by Suwanee Lime-
stone of late Oligocene origin. According to Cooke in Hernando
County this limestone is quite variable in color and_ hardness.
Where it occurs in an unaltered form it usually has a yellowish or
creamy color and generally consists of soft, granular lime-like
particles, some of which are of organic origin. It may occur in
layers up to 100 feet thick.

Elevations in McCarty Woods range from just over 160 feet in
the extreme northwest corner to less than 144 feet in the lower
central portion (Fig. 3). Chinsegut Hill itself, about one and
one-half miles to the northeast, has an elevation of 274 feet and
is one of the highest points in central Florida. By contrast a low
of 70 feet occurs only one and one-quarter miles north of McCarty
Woods. Such marked changes in elevations are hardly character-
istic of Florida where in many places the terrain is so level that
it may drop only a foot or two in several miles.

Soils

Three main soil types are present on this tract: (1) Fellowship
stony clay loam, (2) Hernando fine sandy loam, and (3) Hernando
stony clay loam (Jones and Morrison, 1915). In general surface
soils are grayish or dark brownish-black fine sand and are quite
shallow, usually not over 4 to 6 inches in depth. Below this occurs
dark-brown or blackish, gritty, or more frequently, plastic clay.
In the Fellowship series, which occupies only the northern portions
of the study area, beginning at a depth of 12 to 18 inches there
occurs a creamy, pulverized rocky limestone, obviously derived
from the Suwanee Limestone mentioned above. Other evidence
of this formation was the presence of limestone outcropping in
the vicinity of stations 7 and 8 (Fig. 3). In Hernando fine sandy
loam the gritty or plastic clay extends to at least 30 inches in depth.
This soil type predominates in the southerly three-quarters of the

258 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

tract. Hernando stony clay loam is apparently a graduation be-
tween these two principal soil types and comprises only a small
portion of the area.

The shallow nature of the surface soils and the presence of
clay causes drainage to be extremely poor in nearly level parts of
the tract. This is particularly noticeable following heavy thunder-
storms or prolonged periods of precipitation when water stands in
saucer-like depressions in the north-central or extreme southeastern
portions of the area.

Fire History

McCarty Woods has been protected from fire for many years.
According to Miss Lisa von Borowsky (letter dated November 21,
1965), who worked closely with Colonel Robins and is a long-time
resident of the Brooksville area, these woods have not been burned
since before 1932, or for as long as 34 years. The protection from
burning is apparent in the numerous saplings of white ash, sweet-
gum, American hornbeam and others up to 2 inches dbh, as well
as the density of litter present. A lapse of several years following
burning must be required before conditions become favorable for
some tree species to seed in, for two white ash saplings measuring
1.6 and 1.7 inches, respectively, were each only 13 years old.

The absence of fire is also evident in the lack of any significant
amount of longleaf pine reproduction. The smallest such tree ex-
amined was 11.4 inches dbh, although others as small as 9-10 inches
were observed but not measured. Wahlenberg (1946) regards fire
as virtually a necessity in perpetuating natural stands of this
species.

Vegetation of the
Chinsegut Hill-McCarty Woods

Overstory Vegetation

Quadrat studies and other observations revealed a total of 2]
plant species regarded as trees on the area. Thirteen of these
were recorded by the point-centered quarter method, four by
understory vegetation quadrat studies, and four by general ob-
servations. Tree species encountered were the following (listed
in order of frequency of occurrence according to the point-centered
quarter method):

BecxwitH: Chinsegut Hill-McCarty Woods 259

Sweetgum (Liquidambar styraciflua ) 36.3 per cent
Water oak (Quercus nigra ) 15.0 per cent
Pignut hickory (Carya glabra) 8.8 per cent
American hornbeam (Carpinus carolinianus ) 8.8 per cent
Eastern hophornbeam (Ostrya virginiana) 8.8 per cent
Longleaf pine (Pinus palustris ) 7.5 per cent
Swamp chestnut oak (Quercus michauxii) 5.0 per cent
Live oak (Quercus virginiana ) 2.5 per cent
Florida basswood (Tilia floridana) 2.5 per cent
Hybrid oak (Quercus nigra x Q. virginiana) kD per cent
Winged elm (Ulmus alata) 1.2 per cent
White ash ( Fraxinus americana ) 1.2 per cent
Persimmon (Diospyros virginiana ) 1.2 per cent

Cabbage palm (Sabal palmetto ) —
Redbay (Persea borbonia) _—
Sugarberry (Celtis laevigata ) —
American elm (Ulmus americana ) —
Ground oak (Quercus pumila) —
Sweetbay (Magnolia virginiana) —
Sour orange (Citrus aurantium ) —
Fringetree (Chionanthus americanus ) —

According to the above tabulation sweetgum was by far the
most abundant tree, occurring in a total of 29 (36.3 per cent) of
the 80 possible locations. Most of these were 5 inches or less in
diameter, but a few were over 9.0 inches dbh. This tree was also
the most widely distributed species as it occurred the entire length
of the tract. Water oak was the next most common tree species,
although it occurred in only 12 (15.0 per cent) of the quadrants.
The next most common hardwoods were American hornbeam and
eastern hophornbeam, which occurred only in the central or south-
ern (lower) portions of the study area. Other species encountered
were swamp chestnut oak, live oak, Florida basswood, winged elm,
white ash, and persimmon. One tree observed was definitely a
hybrid between water oak and live oak. Quadrat studies of under-
story vegetation added the following species to the list of trees
present on the tract: Ground oak, sweetbay, sour orange, and
fringetree. The finding of fringetree on this tract is the first record
of this species in Hernando County. In addition to the above trees
the following species were observed but not tallied because they

260 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

failed to occur on the plots: Cabbage palm, redbay, sugarberry,
and American elm.

Exceptionally large or tall specimens of all hardwoods en-
countered were measured whether or not they occurred on the
study plots. Diameters and heights of such trees are listed in the
following tabulation:

Diameter Height

Tree Species (inches ) ( feet )
Pignut hickory 20.6 88
21.9 75
Swamp chestnut oak 27.4 103
30.2 105
Live oak 42.9 110
39.4 87
American elm 16.4 69
Sugarberry 21.6 68

It will be noted that the two largest trees recorded were both
live oaks, one measuring 42.9 inches and 110 feet in height and
the other 39.4 inches dbh and 87 feet in height. Two of the tallest
trees measured were swamp chestnut oaks of 30.2 inches and 105
feet and 27.4 inches and 103 feet.

Longleaf pine.—This species occurred in only the northerly
three tiers of observation plots, or in the north 250 feet of the tract.
This is also the highest portion of the area (Fig. 3). Longleaf pine
occurred in only 6 (7.5 per cent) of the 80 possible locations. The
tallest individual measured had a height of 103 feet and a diameter
of 15.6 inches, but the largest pine recorded was 22.3 inches in
diameter and 87 feet tall.

The maximum age determined from increment borings was 135
years, and this from a tree measuring 19.6 inches dbh. The young-
est tree was 105 years old measuring 15.3 inches in diameter. Based
on the nine borings taken, the average age of longleaf pines on
the study area was 116.6 years.

Based upon measurements made from increment borings the
longleaf pines had a recent growth rate (i.e. during the last inch
of diameter growth) averaging about 12.5 years per inch, but this
ranged from a low of only 5 years (19.6 inch diameter) to a max-
imum of 26 years (21.0 inch diameter). Early growth rate of

BeckwitH: Chinsegut Hill-McCarty Woods 261

these pines was at the rate of from 2 to 4 and occasionally as
many as 7 years per inch of diameter. These data compare with
rates of 10.0 and 5.4 years for recent and early growth, respective-
ly, of longleaf pines in South Carolina according to Wahlenberg
(1946). McCarty Woods trees, therefore, made more rapid growth
during their early years and slower growth in the immediate past
few years than South Carolina trees.

Understory Vegetation

This class of vegetation comprised all major plant species from
the surface to four and one-half feet above the ground. Many tree
species previously encountered by the point-centered quarter meth-
od were also recorded in this quadrat study, and for that reason
they are also mentioned in the following species list. Plants are
grouped according to whether they are trees, shrubs, vines, grasses,
or herbs. The first figure following each species is the average
coverage index and the second is the frequency of occurrence ex-
pressed as a percentage of the total 90 quadrats.

Trees
Longleat pine (Pinus palustris ) t Wal
Pignut hickory (Carya glabra) 0.4 10.0
American hornbeam (Carpinus carolinianus ) 1.4 34.4
Eastern hophornbeam (Ostrya virginiana ) 2 4.4
Swamp chestnut oak (Quercus michauxii) 0.2 8.9
Water oak (Q. nigra) 1.0 53.3
Ground oak (Q. pumila) 0.5 222,
Live oak (Q. virginiana) 0.3 3.3
Sweetbay (Magnolia virginiana) 3.0 3.3
Sweetgum (Liquidambar styraciflua) 2 11.1
Sour orange (Citrus aurantium) 1.0 2.2
Persimmon (Diospyros virginiana) 1.0 ial
White ash (Fraxinus americana) 0.8 10.0
Fringetree (Chionanthus americanus ) 1.0 Jil

Shrubs
Dwarf palmetto (Sabal minor) 0.5 14.4
Saw palmetto (Serenoa repens) 1a lees
Blackberry (Rubus penetrans ) 0.3 lest

Poison sumac (Toxicodendron radicans ) 0.9 73.3

262

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Buckthorn (Sageretia minutiflora)

Stiffcornel dogwood (Cornus stricta)
Devilwood (Osmanthus americanus )
American beautyberry (Callicarpa americana)
Viburnum (Viburnum scabrellum)

Vines

Wild-bamboo (Smilax auriculata )

Bullbriar (S. bono-nox )

Wild-grape (Vitis rotundifolia )

Pepper-vine (Ampelopsis arboreum)

Virginia creeper (Parthenocissus quinquefolia )
Yellow jessamine (Gelesmium sempervirens )
Catsclaw (Doxantha unguis-cati )

Fevervine (Paederia foetida )

Creeping cucumber (Melothria pendula)
Climbing hempweed ( Mikania scandens )

Grasses

Carpetgrass (Axonopus sp. )
Paspalum langei

Panicum anceps

Panicum joori

Panicum sp.

Nutgrass (Cyperus sp.)

Herbs

Ressurrection fern (Polypodium pedioides )
Bracken fern (Pteridium acquilinum )
Ebony spleenwort (Asplenium platyneuron )
Wood-grass (Oplismenus setarius )

Green dragon (Arisaema dracontium )

_ Fringed orchid (Habenaria quinquesta)

Swamp smartweed (Polygonum hydropiperoides )

Smartweed (P. virginianum)
Cock’s comb (Celosia nitida)
Beggar-weed (Desmodium sp.)
Cadillo (Urena lobata)

Frost-weed (Crocanthemum sp.)
Violet (Viola walteri)

Spiny-pod (Cyanchum scoparium)

0.7
0.9
1.0
0.4
0.6

On
1.0
0.8
0.2
0.3
Om
0.5
0.6
0.4
0.1

Beckwith: Chinsegut Hill-McCarty Woods 263

Dichondra (Dichondra carolinensis ) 0.4 5.6
Blue curls (Trichostema dichotomum ) t 3.3
Sage (Salvia lyrata ) 0.2 10.0
Elytraria (Elytraria carolinensis ) 0.6 ela
Dyschoriste (Dyschoriste sp.) O22 14.4
Ruellia (Ruellia caroliniana ) t eh
Justicia (Justicia cooleyi) 0.7 alee
Wild-coftee (Psychotria nervosa) 0.9 Pahl
Twin-berry (Mitchella repens ) 0.7 Pa) al
Bedstraw (Galium floridanum) t 4.4
Birthwort (Aristolochia sp.) t Gn7,
Crownbeard (Verbesina laciniata ) 0.5 ee
Beggar-ticks (Bidens bipinnata) 0.1 10.0

Shrubs.—Of all plants encountered on this five-acre tract the
most numerous one was common poisonivy. This species occupied
as much as 20-40 per cent of some plots and occurred on 73 per
cent of all plots examined. Other common shrubs observed in-
cluded: Dwarf palmetto, saw palmetto, blackberry, buckthorn,
stiffcornel dogwood, devilwood, American beautyberry, and vibur-
num. The noting of dwarf palmetto on this area represents its first
record for Hernando County.

Vines——In proportion to other areas of similar size McCarty
Woods has a relatively large number of vines, some woody in tex-
ture, others of a more or less herbaceous nature. The most com-
mon vines (with frequencies of occurrence of 40 per cent) were
wild-bamboo and catsclaw, which were both characteristic of the
southern or lower and moist three-quarters of the tract, and yellow
jessamine, which was abundant in the dryer, northerly portions of
the tract. Other common vines were Virginia creeper, wild-grape,
and fevervine. Virginia creeper, catsclaw, and fevervine together
frequently formed dense tangles over fallen trees and occasionally
solid mats over extensive portions of the ground surface.

Grasses and Herbs.—The most abundant grass-like plants ob-
served were nutgrass and panic grass, which occurred on 43 and
30 per cent of the quadrats, respectively, and were fairly well dis-
tributed over the entire study area. Among the herbs by far the
most common one was wood-grass, a creeping, perennial herb,
which occurred on almost two-thirds (65 per cent) of the quadrats
and sometimes occupied up to 60 per cent of the area where it

264 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

occurred. It was distributed practically over the entire study area.
Another herb, almost as numerous as wood-grass, was elytraria,
which was present on slightly over one-half (51 per cent) of the
plots examined. Other more frequent herbs included some that
were restricted to the moister portions of the area. These were
justicia, twin-berry, and elephant’s foot. Another common herb,
wild-coftee, was limited to the more level, poorly drained, central
portion of the study area.

Some herbaceous plants collected on this area also represented
county records. These included cadillo and blue-curls, which had
not been previously collected in Hernando County although they
had been reported in neighboring areas. Still another herb, a
smartweed (Polygonum virginianum) was not only a new county
record but established a more southerly extension of its range than
had been previously reported. Earlier records had placed the
southern limit of its range in Marion County, some 40 miles to the
north.

Animal Life

In two nights of trapping with 52 Sherman live traps the only
rodent captured was the cotton mouse (Peromyscus gossypinus ).
A total of at least seven different individuals were present on the
area. One more mouse was captured the first night but it escaped
before being marked, making it impossible to ascertain if it was
recaptured. These rodents are characteristic of subclimax, mesic
conditions that exist on this study area. They are also known to
occur in other environmental conditions, such as in the slash pine
flatwoods type, but this is usually in association with cotton rats
(Sigmodon hispidus). Cotton mice rather than cotton rats are
evidently more closely associated with the climax-type vegetation.
Shrews, particularly the short-tailed shrew (Blarina brevicauda)
might also occur on this tract, but none was captured for definite
proof of their presence.

Other mammals whose presence was determined by sight or
sign were the armadillo (Dasypus novemcinctus), gray squirrels
(Sciurus carolinensis), and raccoon (Procyon lotor). Birds either
seen or heard on this tract included the following:

Red-shouldered hawk (Buteo lineatus )

Bob-white (Colinus virginianus )

BeckwitH: Chinsegut Hill-McCarty Woods 265

Chuck-will’s widow (Caprimulgus carolinensis )

Red-bellied woodpecker (Melanerpes carolinus )

Pileated woodpecker (Dryocopus pileatus )

Blue jay (Cyanocitta cristata)

Tufted titmouse (Bacolphus bicolor )

Carolina wren (Thryothorus ludovicianus )

Ruby-crowned kinglet (Regulus calendula)

Red-eyed vireo (Vireo olivaceus )

Cardinal (Pyrrhuloxia cardinalis )

The above list is far from complete, since it fails to include
numerous warblers and other song birds that obviously pass
through the tract during their migration travels.

DISCUSSION

It is apparent from the foregoing summary that this area repre-
sents a rather unique environment in terms of its vegetative com-
ponents. The northerly, higher one-quarter is characterized by
somewhat xeric conditions resulting in the presence of such plants
as longleaf pine, persimmon, saw palmetto, yellow jessamine, and
blue-curls. The height and diameter of the longleaf pine, the scar-
city of pine reproduction, and the abundance of hardwoods as sub-
dominants clearly indicate the direction of plant succession on this
end of the tract. Hardwoods invading this once longleaf site are
principally sweetgum and water oak.

Over the remainder of the study area the vegetation can be de-
scribed as a mesophytic hammock, indicating that it has certain
plants characteristic of an advanced, subclimax nature. According
to Laessle (1942: 35) “Hammocks are woods dominated by broad-
leaved evergreen trees. They occur on a variety of soils, ranging
from well drained to nearly saturated, but they never occupy areas
that are seasonally or periodically flooded.” Climax-type plants ob-
served here include trees such as pignut hickory, Florida basswood
and sweetgum, a large number of vines, particularly fever-vine, and
a wide variety of herbaceous species. The fact that succession here
is from hydric conditions is suggested by the shallowness of the
surface soils and the heavy, clayey structure of the subsoils. The
moist nature of the site doubtless accounts for the presence of the
smartweed (Polygonum virginianum), definitely a wetland species.

266 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
MANAGEMENT IMPLICATIONS

The handling of this tract of land depends largely upon the
principal objectives which motivated its acquisition by The Nature
Conservancy. If one of these was to take possession of a unique
portion of near-climax vegetation typical of central Florida, then
this objective has certainly been accomplished. The only major
tree species which are characteristic of the true climax and are
absent from this area are southern magnolia (Magnolia grandi-
flora) and American holly (Ilex opaca) (Laessle, 1942).

If another objective was to acquire a stand of old growth long-
leaf pine characteristic of past days in this region of the country,
this has been accomplished. Although comprising only about an
acre, the longleaf pine stand on this tract is doubtless typical of
pine forests that occurred here before the advent of the white man.

There is a definite question, however, about the long-time pres-
ervation of these few longleaf pines, presumably a third major
purpose in the purchase of this area. Doubt arises from the slowed
growth rate of the larger trees and from the lack of pine repro-
duction. In regard to their rate of growth, these trees are prob-
ably in a somewhat critical state that could be easily upset by ex-
treme environmental conditions. If, for example, they were sub-
jected to a protracted drought of 6-8 weeks or longer they could
become so weakened that they would be easily attacked and killed
by various pine bark beetles such as the southern pine beetle
(Dendroctonus frontalis), Ips engraver beetles (Ips spp.) or the
black turpentine beetle (Dendroctonus terebrans). The presence
of hardwoods intermingled with the pines would aggravate the
drough further since they are more resistant to dry conditions and
thus better able to compete for available water supplies.

One of the primary reasons for concern for the welfare of the
longleaf pines is the almost complete lack of pine reproduction to
replace individuals that are lost to natural causes. This is due
largely to the long protection from fire and the resulting abundance
of hardwoods. To open up this stand of pines it would be of great
help to girdle or poison some of the larger gums or oaks and begin
a program of controlled burning. This would gradually clean out
the hardwoods, remove some of the heavy ground litter and pro-
mote the natural reseeding of the pines.

BeckwitH: Chinsegut Hill-McCarty Woods 267

There are certain other factors that have a detrimental effect
on the longleaf pines. One is the high incidence of lightning,
which may occur on as many as 80 days out of the year in the
vicinity of Chinsegut Hill (World Meteorological Organization,
1953-1956. In Komerak, 1964). Another factor is the general
height of the trees relative to surrounding hardwoods. Longleat
pines are somewhat taller than most hardwoods and they also occur
on higher ground elevations. These two factors make these trees
extremely susceptible to lightning strikes which would eliminate
them from the stand. This is another way in which one vegetative
type is replaced by another in the natural process of plant succes-
sion.

In regard to the mesophytic hammock portion of this tract it
could best be preserved by continuing to protect it from fire. Burn-
ing at this stage of development would probably remove some of
the thick debris that has accumulated and perhaps in time would
improve the general appearance of the area, but it seems wiser to
exclude fire and permit the area to develop further toward a cli-
max vegetation type.

SUMMARY

A general survey of the flora and avian and mammalian fauna
was made from September 1965 through July 1966 of a five-acre
tract owned by The Nature Conservancy in McCarty Woods near
Chinsegut Hill, six miles north of Brooksville, Florida. Soils are
generally shallow and underlain by clayey material over Suwanee
Limestone. Fire has been excluded for at least 34 years. About
one-fifth of the area is composed of old-growth longleaf pine up
to 135 years of age. A dense understory of mixed hardwoods such
as sweetgum and water oak and the shortage of pine reproduction
typifies the long absence of fire. The remaining portion of the tract
is described as a mesophytic hammock and contains a wide variety
of near-climax hardwoods and vines. Cotton mice were the only
small mammals collected, but other mammals and birds observed
were typical of the mesic environment. A program of hardwood
control by prescribed burning or other means would be helpful
in perpetuating the few longleaf pines, but fire should be excluded
from the mesophytic hammock portion of the area.

268 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

LITERATURE CITED

Buair, W. F., A. P. Blam, PreRcE BRopkors, F. R. CAGLE, AND G. A. Moore.
1968. Vertebrates of the United States. McGraw-Hill, New York, 2d
edition, 616 pp.

Cooke, C. W. 1945. Geology of Florida. Florida State Dept. Conserv.,
Geol. Surv., Bull. 29, 339 pp.

CottaM, G., AND J. T. Curtis. 1956. The use of distance measures in phyto-
sociological sampling. Ecol., vol. 37, no. 3, pp. 451-460.

Jones, G. B., and T. M. Morrison. 1915. Soils Survey of Hernando County,
Florida. U.S. Dept. Agr., Bur. Soils, 30 pp.

Kesey, H. P., AND W. A. Dayton (editors). 1942. Standardized plant
names. J. Horace McFarland Co., Harrisburg, Pa., 675 pp.

LarEssLE, A. M. 1942. The plant communities of the Welaka area. Univ.
Fla. Publ. Biol. Sci. Ser., vol. 4, no. 1, 143 pp.

RapForp, A. E., H. E. AHLEs, AND C. R. Bett. 1964. Guide to the flora of
the Carolinas. Univ. North Carolina, Chapel Hill, N. C., 383 pp.

SMALL, J. K. 1933. Manual of the southeastern flora. Univ. North Carolina,
Chapel Hill, N. C., 1554 pp.

TREIMAN, M. R. Undated. Chinsegut Hill, 2 pp., lithog.

U. S. DEPARTMENT OF COMMERCE. 1966. Climatological data: Florida—
annual summary 1965. Environ. Scien. Serv. Admin., Weather Bur.,
vol. 69, no. 13, pp. 178-186.

WAHLENBERG, W. C. 1946. Longleaf pine—its use, ecology, regeneration,
production, and management. Chas. Lathrop Pack Foundation, Wash-
ington, 429 pp.

Wor.LD METEOROLOGICAL ORGANIZATION. 1953-1956. World distribution of
thunderstorm days. Tech. Pub. 21, Parts 1 and 2. In Komerak, E. V.
1964. The natural history of lightning. Proc. 3rd Tall Timbers Fire
Ecol. Conf., pp. 139-183.

School of Forestry, University of Florida, Gainesville, Florida
Florida Agricultural Experiment Stations Journal Series, no. 2778.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968 )

Relationship of Sand Pine Scrub to Former Shore Lines

A. M. LAESSLE

No plant community in Florida has stimulated more interest
and near unanimity as to its floristic distinctness and the sharp-
ness of its boundaries with the adjacent sandhill vegetation than
the Florida scrub. But much speculation has taken place as to the
reasons for the floristic differences and the general lack of inter-
gradation between these associations as well as their origins and
successional relationships.

Vignoles (1823) was among the first to comment on the general
similarity in the appearance of scrubs wherever found. Nash
(1895) commented on the almost complete floristic difference be-
tween scrub and adjacent sandhill vegetation and stated that the
“two floras are natural enemies and appear to be constantly fighting
each other.” He also commented, “A bare space of pure white
sand usually separates the two.” He considered the sands of both
communities to have had the same origin and attributed the gen-
erally darker surface of the sandhill soils to be caused by charcoal
deposited there by annual fires. Harper (1940) commented that
fires usually sweep through the scrub about once in a lifetime.
Whitney (1898) remarked that “the boundary between high pine
land (sandhills) and scrub can be located without trouble within
a few feet. . . . It will be shown later that there is no apparent
reason, from the chemical and physical examination, to account
for this difference in the native growth on the scrub as compared
with high pine land—so far as our investigations show there is no
difference in the soil.”

Many have attributed the great vegetational differences be-
tween the communities to lower available nutrients in the scrub
soils (Harper, 1921; Kurz, 1942, with reservations; Laessle 1942,
1958a). Mulvania (1931) in his study in central Florida found
about 6 per cent greater water holding capacity in the sandhill
soils and concluded that most of the evidence indicates that factors
other than fire, such as differences in soil and water relations,
account for the differences between high pine and scrub forests.
Harper (1921) states that scrub soils are poor in humus, clay,
and potash.

Webber (1935) considered the relatively high frequency of

270 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

fire in the sandhill community and the rarity of fire in the scrub to
be the most important factor in the maintainance of the sharp
boundary so generally evident when the two associations abut
one another. He stated that the scrub association is a fire-fighting
machine.

Both Kurz and Laessle (1942, 1958a) realized that most scrubs
occupy ancient dunes formed along shore lines of higher Pleisto-
cene sea levels, some as high as 100 feet (Kurz) or 150 feet (Laessle,
1958a) above present sea level. Kurz also pointed out that with
very few exceptions the plants distinctive of scrub are also found
on relatively recent coastal dunes, and some scrubs might occupy
old wave formed bars or ridges.

Laessle (1958a) hypothesized that wave action along shores of
lakes and even stream action could have washed and sorted sand
deposits which now support scrub.

Kurz thought that the generally yellow soils occupied by sand-
hill vegetation would eventually be lightened in color by leaching
and be succeeded by scrub vegetation. While Laessle (1942,
1958a) thought that the two communities bore no seral relation-
ship, he also found some sandhill communities occupying typical
dune sands.

Such diverse ideas on the reasons why such physionomically
and taxonomically diverse communities can exist side by side with
such sharp boundaries between them, often without any change of
elevation from one to the other, should, I believe, be capable of a
more consistent analysis and interpretation.

METHODS

Vegetational changes across the ecotone between scrub and
sandhill communities were studied by line transects (Buell and
Cantlon, 1950). A 100 meter tape was stretched at approximate
right angles to the ecotone with the 50 meter mark as close to the
middle of the ecotone as could be estimated. From the zero end
in the scrub, all herbs, lichens, and mosses were recorded in the
order intercepted and plotted to the nearest centimeter along a
line in a field notebook. On the same line the intercept to the
nearest decimeter of all shrubs (here defined as woody species
less than 6 feet high) and trees (woody species above 6 feet)

LAEssLE: Scrub and Shore Lines OAT

were also plotted. The results of six of these vegetational transects
without the amount of intercept are shown as plotted from field
notes for each of the three layers in Fig. 1. Some species are
omitted for the sake of simplification.

Soil profiles and soil samples were taken in each community
at least 20-50 meters away from the ecotone. Ten soil samples
near both sides of the transect tape were taken in each com-
munity at depths of 0-2 inches, 2-6 inches, and 6-15 inches. The
10 samples from each depth were then pooled from each com-
munity and analyzed by a graduate student under the direction
of Dr. Hugh Popenoe of the Soils Department of the University
of Florida. Chemical analyses were made from duplicate samples
after being air dried and passed through a 2 mm sieve.

Soil pH was determined using a Beckman Zeromatic pH meter
with glass electrodes. One part soil to 2 parts water was allowed
to stand overnight before testing in one method. In a second
method KCl was added to the above solution to make a normal
solution, which was stirred and allowed to stand for 30 minutes
before testing. The second method is considered more accurate
as it compensates for such elements as aluminum which may be
present.

The exchangeable cations of Ca, Mg, Ka, and Na were extracted
with normal ammonium acetate buffered at pH 4.8. Calcium and
potassium were determined on a Beckman Model B Flame Spec-
trophotometer. Magnesium and sodium were determined on a
Beckman Model DU Flame Spectrophotometer.

Extractable phosphorus was determined colorimetrically on a
Baush and Lomb Spectronic 20 colorimeter using strong Bray ex-
tractant. Nitrogen was determined by titration of a boric acid
solution (Kjeldahl method for total nitrogen). Organic carbon in
the soil was determined by the Walkley-Black wet combustion
method as modified by Walkley. Organic matter was determined
using potassium dichromate as a reducing agent and titrating
with ferrous ammonium sulfate (modified Walkley-Black as modi-
fied by Walkley). Mechanical analysis was done by the method
of Bouyoucos (1951, 1962). The same type of analyses were
made from 15 scrubs frorn widely scattered points as far south as
Deerfield, on the lower Florida east coast and Destin in the Flor-
ida panhandle.

272 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

VEGETATIONAL TRANSECTS

Fig. 1 shows linear distribution of selected species, those that
were considered most characteristic indicators of the two commu-

000 O dom O Oo
@O oO ap

Poo
oo

DOomn
ie)
©

100M.
COLUMNS HEADED BY H=HERBS, S=SHRUBS, T=TREES

4 PANICUM PATENTIFOLIUM GO CLADONIA EVANSII @ QUERCUS MYRTIFOLIA § PINUS CLAUSA

W RHYNCHOSPORA DODEC- © CLADONIA LEPORINA B QUERCUS CHAPMANII 8 QUERCUS LAEVIS »6
ANDRA ap SELAGINELLA SPP. A LYONIA FERRUGINEA 0 PINUS PALUSTRIS

% SOLIDAGO CHAPMANI! #@ ARISTIDA GYRANS f VACCINIUM STAMINEUM

™ LIATRIS LAEVIGATA O ARISTIDA STRICTA ™= CERATIOLA ERICOIDES

@ CLADONIA SUBTENUIS © SPOROBOLUS JUNCEUS #OSMANTHUS AMERICANUS
@ DICRANUM CONDENSATUMO SORGASTRUM SECUNDUM § QUERCUS LAEVIS «6
@ ANDROPOGON FLORIDANUS A ERIOGONIUM TOMENTOSUM

Fig. 1 Line transects through scrub (0-50m) and abutting sandhill vege-
tation (50-100m). Darker symbols represent typically scrub species.

LAESSLE: Scrub and Shore Lines 273

nities. Those species most typical of the scrub are indicated by
the darker symbols, those of the sandhills by the lighter ones.

The most obvious and important difference is in the herbaceous
layer, especially the wire grasses Aristida stricta and Sporopolus
Junceus (S. gracilis (Trin.) Merr). Only one small clump of
Aristida stricta was on the scrub side, and 249 clumps of the com-
bined species were on the sandhill side. While a much larger
number of apparent clumps of wire grasses is shown in transect
4 than in any others, this may well be influenced by two factors.
Shortly after a fire has burned off an area, the blades are too
short to assume a horizontal position and are less apt to be inter-
cepted, as was the case in transect 3. When fire has not occurred
for many years, closely spaced clumps may be so enmeshed with
both dead and living blades that they appear as a single clump.
In transect 4 fire had been recent enough for the separate entities
to be distinguished, but the blades were long enough to have
spread horizontally and thus were more readily intercepted. The
other four transects had not burned for at least 10 years, and
their intercepts no doubt often coalesced to form compound
clumps.

Woody species typical of the scrub such as myrtle and Chap-
man’s oaks, Lyonia ferruginea, and sand pine are not infrequently
encountered on the sandhill side but tend to become less common
with increasing distance from the scrub. Quercus chapmanii, per-
haps because of its thicker more fire resistant bark, tends to show
proportionally greater survival in the sandhill side than does Q.
munciolg tor example, on the scrub side 171 clumps of Q.
myrtifolia occurred to 59 on the sandhill side. Q. chapmanii in-
tercepts were 14 in the former and 16 in the latter.

The absence of fire permits a rapid invasion of the sandhills
by sand pine where a seed source is available. This is best shown
in transect 2. In transect 3 many 2-3 year old sand pines had been
killed by the ground fire only a month previous to the sampling.
This fire, incidentally, burned right up to the edge of the scrub
where it apparently died out. Of the non-key species saw pal-
metto, Serenoa repens, was about equally distributed in each com-
munity (scrub side 27, sandhill side 35).

274 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Soi. TRANSECTS

Results from soil analyses were disappointing to me since I
had long felt the sites of most scrubs are associated with marine
features such as old dunes, bars, and beaches, where maximum
washing and sorting of sands had occurred (Laessle, 1958a). How-
ever, except for five sites all on the “ridge” where the red, clayey
“Citronelle” formation closely underlies the sandhill vegetation no
significant soil difference between the soils of sandhill and scrub
vegetation was apparent. Here, composite results are presented
of comparisons from the two communities located near Kingsley
Lake and Goldhead State Park in Clay County, Mogelvang’s Scrub
and the adjacent sandhill near Rock Springs in Seminole County,
and the Clarcona area just southeast of Clarcona, Orange County.
At these sites a very appreciable increase of phosphorus was en-
countered with increasing depth in the sandhill portions of the
transects. The average extractable phosphorus in the scrub and
sandhills, respectively, was as follows: 0-2 inch horizon, 4.97 vs.
14.32 ppm; 2-8 inch horizon, 1.35 vs. 25.08 ppm; 8-15 inch horizon,
0.92 vs. 26.97 ppm.

Samples from the two communities near Weekiwachee Springs
in Hernando County, and at Inverness-Lecanto and near Citro-
nelle in Citrus County, however showed average extractable phos-
phorus in the scrub and sandhills, respectively, as follows: 0-2
inch horizon, 4.54 vs. 5.05 ppm; 2-8 inch horizon, 3.32 vs. 2.23 ppm;
8-15 inch horizon, 2.30 vs. 1.32 ppm.

Neither the phosphorus nor mechanical analyses showed any
overall consistent differences.

Slightly less yellow 8-15 inch samples from the scrub sides
were often observed and may well be due to the higher amounts
of organic acids leached from the greater accumulation of un-
burned litter encountered there.

Scrub unburned for 20 years or more had soil profiles with
exchangeable cations of K, Ca, and Mg frequently 2-3 times higher
than in the recently burned sandhills. For example, in the Citro-
nelle transect the following analysis was obtained: averages from
the pooling of the 3 horizons samples gave 28.8 ppm vs. 11.5 ppm
for K, 290.0 ppm vs. 82.5 ppm for Ca, and 109.5 ppm vs. 36.25
ppm in scrub and sandhills, respectively. Garren (1943) stated
that burning decreases the acidity, increases the Mg and replace-

LarEssLE: Scrub and Shore Lines Di

able Ca and organic matter in long leaf pine soils, including that
of the sandhills.

On the other hand, if both communities had been unburned for
over 20 years such differences were negligible except for Ca and
Mg which were twice as abundant on the sandhill side: K, 14.0
womlooratnd-2) mech Ca, 87.0 vs. 182.0 at 2-8 inch; Mg, 42.1 vs.
84.1 at 8-15 inch horizons.

Where appreciable difference in these elements is present in
the upper horizon in the soils of the two communities it is doubt-
less due to the burning of the organic particles on the sandhill
soils and their rapid loss through leaching. At the 8-15 inch hori-
zon no consistent difference in these elements was evident.

Recent fires on the sandhill soils also tended to lower the C/N
ratio and raise the pH. In general the scrub soils proved slightly
more acid but comparisons between the soils in some transects
such as the Mogelvant scrub and adjacent sandhill soils, neither
of which had been burned for at least 20 years, show the reverse,
even when run by the N-KCI method for the three horizons
sampled. The pH was 4.5 vs. 3.6, 0-2 inch, 4.65 vs. 4.30, 2-8 inch,
and 4.60 vs. 4.65, 8-15 inch with the scrub again first at each
level.

DIscussION

With no substantiating evidence for any general chemical or
physical differences in the soils of the two communities it has be-
come necessary to reevaluate some of the ideas expressed many
years ago. According to Webber, the nature and relations of the
scrub associations cannot be discussed without comparison with
the high pine association, as the two associations are so commonly
found in juxtaposition and occupy so nearly the same types of
soil in the same region and at the same altitude. He also men-
tions that the two communities may be at the same level or one
uphill or downhill from the other. The change from one flora to
the other is very abrupt between the tangled vegetation of the
scrub and the open forest of long leaf pines with a ground cover
mainly of grasses. The high pine lands burn over almost every
year and this condition has probably existed for many centuries.
The scrub association, however, is a fire-fighting machine.

Webber attributes the fire-fighting equipment to a number of

276 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

distinct factors. Nearly all scrub plants are evergreen and drop
old leaves throughout the year. Scrub bushes and trees so fully
occupy the soil that they absorb available moisture and nutrition
to the extent that the flammable grasses and other herbs are ex-
cluded. The soil surface is thus relatively bare of flammable ma-
terial except for gradually falling leaves and old twigs which
rapidly rot and disappear. Furthermore, a natural fire break is
maintained between the scrub and adjacent sandhill vegetation,
as the extensive root systems of both communities probably limit
the growth of flammable grasses and herbs between these com-
munities.

Fire frequently approaches the scrub and dies out without
gaining entrance. After a number of uninterrupted fire-free years,
however, the scrub accumulates dead limbs and branches, and the
fire-break margin becomes partially covered with grass. Under
these conditions, a fire may ignite the scrub.

Webber's ideas of a natural firebreak between the two com-
munities are often evident to the eye but are not obvious from
the vegetational transects (Fig. 1), where the clumps of wire gras-
ses and other more flammable herbs do not show an appreciable
decrease in their numbers near the scrub border. If the size of
the clumps of wire grasses is ignored there are in the six transects
41 such clumps within 10 m of the scrub borders, 50, 51, 56, and 52
in each successive 10 m interval from this boundary. But, indi-
vidually, only transect 2 shows an obvious trend supporting this
concept while in transect 6 the reverse is evident.

To Webber's explanation as to the occasional spreading of fire
from the sandhills to the scrub, there should be added the fact
that lightning-caused fires may arise in the scrub. The author
has observed such an event when an old sand pine, heavily laden
with Spanish moss, burst into flame when struck by a bolt of
lightning. Old stands of sand pine are frequently festooned with
dense growths of this epiphyte (Fig. 2) which is highly flammable
when dry. Crown fires in the scrub could readily be started in
this manner in only relatively dense stands of sand pine.

Spanish moss could also be an important factor in allowing
creeping ground fires, which not infrequently gain access to the
tops of sand pines and thus start crown fires in scrubs. I have
observed many instances of multiple aged stands of sand pine

LAESSLE: Scrub and Shore Lines iil

Fig. 2. A 50 year old stand of sand pine, 1/2 mile north of Cocoa City
Limit on the E. side of U.S. 1, showing an abundant growth of Spanish
Moss.

which no doubt developed after ground fires had partially opened
scrubs by killing back much of the shrubby understory and even
some of the sand pine. But, as mentioned by Webber, rapid
sprout growth from the crowns of such burned scrub shrubs, soon
develop enough shade to prevent the entrance of the heliotrophs
from abutting sandhill vegetation, at the same time acting as
nurse plants to any young sand pines started from seeds released
by such fires. Cooper et al. (1959) showed that young sand pine
seedlings cannot survive soil temperature above about 125 F.

ORIGIN AND SUCCESSIONAL RELATIONSHIPS

Whitney likewise could find no apparent difference from chemi-
cal and physical analyses of the soils of these communities to ac-
count for their great vegetational differences and thought that it
was chance that determined what area became scrub and sand-

278 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

hill associations and said that the two kinds of vegetation were
not capable of growing together.

According to Webber, the origin of the large scrub areas is
probably traceable in part to the poor, arid character of the soil
on which the associated plants of the scrub can succeed better
than any other group of plants. It would seem probable that in
the early formative period of plant distribution in this part of
Florida, the scrub oaks, because of their mode of spreading by
sprouts from the roots and their adaptability to succeed in such
arid soils, found in these areas an opportunity to dominate the
other vegetation and finally to extend their territory as natural
conditions permitted. This does not explain why scrub vegeta-
tion should become established instead of the sandhill type unless
it can be proved that the scrub soils are actually poorer and drier
than those occupied by sandhill vegetation.

I think it more than chance that scrubs for the most part
occupy well drained positions on ancient marine shoreline fea-
tures as dunes, bars, and beaches and even old fresh water shore-
lines and stream deposits (Laessle, 1958a). Even recent dunes
are colonized by scrub species. It is relatively rare that scrubs
cannot be tied to any of ancient marine or freshwater features.
Yet some sandhill communities do occupy definitely dune topog-
raphy, for example, the previously reported area east of Frost-
proof. This makes the problem difficult to deal with in a simple
and satisfactory way. Evidence of greater tolerance of scrub spe-
cies to salt spray is abundant. It is especially striking on the
dunes just before entering St. Andrews State Park, Bay County,
Florida (Fig. 3). No examples of the sandhill communities in
such a severe environmental situation have been observed. The
sites originally occupied by scrub vegetation fit most situations
and the persistence of sandhill vegetation becomes more plausible
in the light of the scrub’s resistance to frequent fire as pointed
out by Webber.

The present existence of scrub along the shoreline about Red
Hill (see the Childs topographic Quadrangle, U. S. Geological
Survey, 1953), mentioned by Laessle (1958a) as being mid-Pleisto-
cene according to MacNeil (1950), is at least 1,000,000 years old
and may be as much as 10,000,000, and thus Pliocene or even
late Miocene (H. K. Brooks, 1966), a most striking example of

LaEssLE: Scrub and Shore Lines 279

Fig. 3. A highly modified scrub on the second dune ridge 1/4 mile W.
of St. Andrews State Park. This scrub not more than 150 yards from the
Gulf of Mexico shows the effect of wind and salt spray. The sand pine
though at least 60 years old scarcely reached 3 feet in height but had
branches extending at least 25 feet to the leeward.

the scrub’s persistence. Brooks further states that the maximum
Pleistocene sea level did not exceed 90 feet above present and
believes that the climate at many times in the Pliocene was as dry
as that of Corpus Christi, Texas. Perhaps both the sandhill and
scrub vegetation have existed and maintained their same positions
without any great floristic change through all this time or even
longer.

Possible successional relationships between the two communities
have been most varied and speculative. The following from Gar-
ren summarizes some of the ideas concerning scrub forests at that
time. According to Pessin scrub on dunes with no fire would be
succeeded by long leaf pine, oak, hickory and on dunes with in-
frequent fire would be followed by long leaf pine. Harper (1940)
thought that there would be no change with fire once in a life-
time. Webber thought that fire would destroy the scrub if fre-
quent and it would be followed by turkey oak and bluejack oak.
Albert thought that scrub fires in young, non-cone bearing pine

280 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

would be followed by scrub and myrtle oak, and that crown fires
in older sand pine stands would result in no change.

To these ideas must be added those of Kurz who stated that
high pine land vegetation was succeeded by scrub. Laessle (1942)
considered that these communities had no seral relationship. Mil-
ler (1950) thought that sand pine scrub was succeeded by high
pine-turkey oak. Laessle (1958a) stated: “Scrub and_ sandhill
vegetation represent the two most prevalent well-drained sub- or
fire-climax communities in peninsular Florida. Each in the ab-
sence of fire would eventually, and often very rapidly, be succeeded
by predominately evergreen hardwood tree communities, known
in much of the Southeast, as hammocks.” Webber considered
that even with the destruction of the high pine forest (sand hill
community ) scrub would very rarely replace it.

After extensive observation throughout most of Florida and
southeastern Alabama, my thinking on the matter unfortunately
cannot be summed up succinctly. Since the time when most of
the above opinions were expressed many areas of sandhills which
have close proximity to scrubs have been protected from fire for
twenty or more years and the very rapid spread of sand pine
from nearby scrub is very striking. One such area extends for
many miles along U. S. 98 in the area included in the Eglin
Air Force area, between Pensacola and Fort Walton. Here, with
no sign of fire for a very long period, sand pines of all ages
were present from 1.5 ft. d.b.h. to small seedlings but very little
other typical scrub vegetation was present. Myrtle oak was sparse
with scattered Ceratiola present. The other tree vegetation was
predominantly sandhill species such as turkey oak and long leat
pine. Both wire grasses Aristida stricta and Sporobolus junceus
were thin and scattered. Most of the other herbs were typical
sandhill forms, the exception being Rhynchospora dodecandra
which was frequent. The rapid spread of sand pines to the sand-
hill islands in the Ocala National Forest is nearly everywhere evi-
dent since fires have been prevented. Its spread in the sandhills
at the University of Florida Conservation Reserve, Welaka, Put-
nam County, Florida, is continuing rapidly from a nearby scrub,
since observed by Laessle (1958b). The same trend was also
marked in the Camp Blanding area, particularly in the Clay
County portion where at least two scrubs are known to occur.

LAEssLE: Scrub and Shore Lines 281

The invasion of sandhills by the typically scrub oaks, Q. myrtifolia
and Q. chapmanii is apparently much slower but their occurrence
there with decreasing frequency as one moves further from adja-
cent scrubs is evident from the transects (Fig. 1). It would thus
appear that the scrub species will differentially invade the sand-
hills but that a much longer fire free period must occur for com-
plete replacement to be accomplished. It is also of interest to
note that in spite of the supposed racial difference in the pan-
handle form and the peninsular form described by Little and
Dorman (1952) in which the former was described as having less
serotinous cones, seed release by both subspecies without fire is
sufficient to cause their rapid invasion of adjacent sandhill com-
munities when fire is eliminated.

Of interest in this regard is Webber's interpretation that the
present limits of the large scrub areas became established centuries
ago and that a state of near equilibrium has existed for many
years. If Webber considers fires to be a natural condition in the
sense that lightning frequently causes them in this region, then I
agree with him but would extend the time of relative stability of
the two communities to millions of years rather than hundreds.

The Florida scrub is the ecological equivalent of the California
chaparral and according to Axelrod (1958) extensive paleobotani-
cal studies over the past 40 years have shown that chaparral has
been a major plant formation in California since mid-Pliocene.

The fate of scrub kept fire-free for extended periods of time
is not as clear as I once thought (Laessle, 1942, 1958a). One
scrub on Bear Point, an arm like projection into Lake Placid’s
eastern shore, just north of Childs, Highlands Country, Florida, has
been protected from fire for an extended period and is composed
of typical scrub flora except for a complete lack of sand pine.
This scrub has the aspect of a hammock but lacks any characteris-
tic arboreal hammock species. Part of this peculiarity may be the
lack of a seed source in this region of such typically hammock
species such as Magnolia grandiflora, Quercus laurifolia, Carya
glabra, Ilex opaca, Persea borbonia, Cornus florida, etc., which are
rare and far from this site. Such typical scrub species as Carya
floridana with a d.b-h 13.1 inch and Quercus myrtifolta with a
d.b.h. of 11.0 inch were observed as also were such characteristic
hammock species as Mitchella repens and Callicarpa americana.

282 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Though no signs of fire were detected on the tree trunks here,
charcoal was detected in a soil sample at a depth of 7-8 inches.
The nearest sand pines were at least a mile from this scrub.

It seems very likely that sand pine formerly occurred here but
the lack of fire for an extended period has not permitted its re-
generation. Sand pines over 75 years old are very rare and on
this basis a fire free period of at least this duration is suggested.

Another very old scrub with a few large and widely scattered
72 year old sand pines was studied on U.S. 1, about 1 mile south
of the Rosedale Road, Indian River County, Florida. Though no
sign of fire was observed on the oldest trees, charcoal was also
abundant in the soil here. There were a few 8-10 foot sand pines
and a few small seedlings but the ground vegetation was very
sparse consisting mostly of lichens with extensive areas of bare
white sand. Lichens, Cladonia evansii, C. leporina, C. subtenuis,
and C. prostrata comprised 11.9 per cent of the ground cover.
The per cent of shrub cover, mostly Ceratiola ericoides, Quercus
myrtifolia, and Serenoa repens was only 15.6 per cent. The
tree canopy composed almost exclusively of sand pine was only
23.19 per cent. It is difficult to imagine how this sparse vegeta-
tion could possibly support a fire either of ground or crown type
which could initiate a rejuvenation of the scrub vegetation. No
signs of any invasion of hammock species were observed. It thus
appears that it is possible for scrubs to run out, or at least, reach
a very depauperate condition in which state they may remain in-
definitely.

A similar and almost as sparsely vegetated scrub was studied
at Destin, Okaloosa County, Florida. Here no fire was evident on
the bark of sand pines, some of which were over 105 years old.
Ground vegetation composed only 9.9 per cent cover which was
mostly Cladonia evansii, C. subtenuis, C. leporina, and Dicranum
condensatum. Shrubs formed 33.8 per cent cover largely of Quer-
cus myrtifolia and Chrysobalanus oblongifolius. The tree canopy
was 40.1 per cent, 33.36 of it being scattered sand pine which in
spots contained some juvenile trees. No signs of invading ham-
mock species nor species typical of any other community were de-
tected. Here charcoal was also found in the soil. Parts of this
scrub contained enough sand pine reproduction and ground vege-

LAESSLE: Scrub and Shore Lines 283

tation to make rejuvenation possible either by means of ground
or crown fire, but this scrub’s fate seems dubious.

It is worth noting that the three scrubs described above are
small and isolated from abutting sandhill vegetation or any other
flammable communities.

On the other hand scrubs showing evidence of succession
toward hammock are not infrequently encountered. One _ such
example was studied on the offshore bar and dune system just
West of U.S. A1A, about a mile north of Ormond Beach, Volusia
County, Florida. Here fire protection has been in effect for a
considerable period due to its close proximity to many dwellings.
Many laurel oaks, red bays, and a few Magnolia grandiflora were
thoroughly mixed with scrub species. Although the sand pines
were quite mature, many in the 8-10 inch d.b.h. class, the larger
laurel oaks were only about 2 inch d.b.h. Soil analysis in the
exchangeable cations, K, Ca, Mg, and P while higher than in
most scrubs studied were not appreciably different from others
which showed no signs of invasion by hammock species. Young
scrub soils, as the one here, may not have been subject to the
leaching of elements for a long period as were many of the very
old inland ones and this factor may well be of significance.

I have observed no evidence of succession toward sandhill
vegetation from scrub. There are, however, some typically sand-
hill plants as Quercus laevis found in extensive otherwise typically
scrub areas, notably in many portions of the Ocala National Forest.
Here they are always scattered and not generally accompanied
by any appreciable number of young trees. Here, also, are oc-
casional sandhill herbs such as Stillingia spathulata and stinging
neetle, Cnidoscolus stimulosus but never in large numbers and
only when a more clayey subsoil is found within a foot or two
from the surface or when there is evidence of mixture of dune
sand with soils from nearby areas supporting sandhill vegetation,
(Laessle, 1958a). I have never observed the invasion of Pinus
palustris in the scrub. This supports the observations of Webber
that large long leaf pine trees extend entirely to the margin [of
the scrub]. The complete absence of the remains of the extremely
decay-resistant heart of longleaf stumps even in the periphery of
the scrub was given by Laessle (1958a) as supporting evidence of
the past spatial stability of the two communities.

284 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

However, the finding of sandhill vegetation on definitely dune
topography as a shoreline feature of the Sangamon interglacial stage
as reported by the author (1958a, p. 375), does hint at the idea that
scrub vegetation may have formerly existed there and that an
unusually high frequence of fire may have eliminated it.

CONCLUSION

Webber's paper was based on his observations on inland scrubs,
many of which lack the higher phosphorus content observed in
abutting sandhills. He believes the lack of frequent fires in the
scrub, and their almost yearly occurrence in the sandhills, to be
the most important single factor responsible for the maintainence
of the very sharp boundary between the communities and to
account for their spatial stability for the very extended periods.

Where scrub is not isolated from higher sandhill vegetation the
former occupies the more coastal position probably because of the
greater salt spray tolerance of scrub vegetation, the factor that
ground fires burn down hill with difficulty, and the fact that on-
shore winds would tend to keep fires less frequent in the areas
occupied by scrub. In the light of well established Cenozoic
shorelines away from present coastlines, similar origins for most
of the more ancient scrubs is postulated.

There remains the problem of explaining the scrubs which can-
not be tied to the definitely ancient shore lines. Part of this
difficulty may be due to the great changes in topography that
have been wrought by the karst nature of Florida which has ex-
tensively changed the original topography through solution of the
soft Ocala limestone. Furthermore, many of such problematic
scrubs are partially or completely bordered by vegetation much
less subject to frequent fires than in the sandhills. Provided the
soil is poor and excessively drained, even subtle differences in
fire frequency may be enough to permit the development of such
partially fire-protected scrubs.

While scrubs are remarkably similar with regard to their major
vegetative components their apparent isolation from each other is
not as great as one might at first believe. Such important scrub
elements as myrtle oak (Quercus myrtifolia), Chapman’s oak (Q.
chapmanii), live oak (Q. virginiana, with its many variations),

LaEssLE: Scrub and Shore Lines 285

and tree lyonia (Lyonia ferruginea) frequently occur in the drier
portions of intervening “scrubby flatwoods” of Laessle (1942).
These species also are often widely scattered in the sandhills and
drier hammocks. The major gap in the range of typical scrubs
containing sand pine is between the inland peninsular scrubs in
Clay County and the panhandle scrubs the nearest of which are
in Franklin County, a distance of approximately 150 miles. The
nearest coastal peninsular scrub at Cedar Key, is over a hundred
air line miles from the Franklin County scrub. If the range of
sand pine were once more continuous, it is most likely that very
considerable erosion of the limestone plain in the region of the
Suwannee River Valley is the most plausable cause for the large
gap in the present range of the sand pine (H. K. Brooks, personal
communication). If the separation of sand pine into two races, a
panhandle form and a peninsular form is valid, as proposed by
Little and Dorman, this gap in the present range must have
existed for a considerable period.

The present sandhill vegetation presents no such spatial prob-
lem, as this community is nearly continuous from at least south
central North Carolina, and Hale County in central Alabama, to
its southern limit in Desoto and Highlands counties, Florida.

In conclusion, there is little doubt that these most distinct
communities have existed essentially unchanged, for the most part
from at least the Miocene, and that the differential frequency of
fire in them has played the major role in maintaining their most
contrasting floristic and structural characteristics. Soil difference
between scrub and sandhill areas, at least as far as the macro-
elements and physical characteristics present are concerned, are
not consistently great enough to account either for the generally
sharp boundary between them, nor for their more general spatial
pattern.

AKNOWLEDGMENT
Financial support for this study was obtained through a grant
from the National Science Foundation.

LITERATURE CITED
AxELRopD, D. I. 1958. Evolution of the Madro-Tertiary geoflora. Bot.
Rev., vol. 24, pp. 433-509.
Bovyoucos, G. J. 1951. A recalibration of the hydrometer method for
making mechanical analysis of soils. Agron. Jour., vol. 43, pp. 434-
438.

286 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

1962. Hydrometer method improved for making particle size analysis
of soils. Agron. Jour., vol. 54, pp. 464-465.

Brooks, H. K. 1966. Geological history of the Suwannee River. In Geology
of the Miocene-Pliocene series of the Georgia-Florida area (N. K.
Olson, ed.), pp. 37-45. Atlantic Coastal Plain Geological Association
and Southeastern Geological Society.

BuELL, M. F., anp J. E. Cantton. 1950. A study of two communities
of the New Jersey pine barrens and a comparison of methods.
Ecology, vol. 31, pp. 567-586.

Cooper, R. W., C. S. ScHOPMEYER, AND W. H. D. McGrecor. 1959. Sand
pine regeneration on the Ocala National Forest. U.S.F. Serv. Prod.
Res. Rep. No. 30, pp. 1-37.

GarrEN, kK. H. 1943. The effect of fire on vegetation of southeastern
United States. Bot. Rev., vol. 9, pp. 617-654.

Harper, R. M. 1921. Geography of central Florida. Florida Geol. Surv.,

7th Ann. Rept., pp. 71-307.

1940. Fire and forests. Am. Bot., vol. 46, pp. 5-7.

Kurz, HERMAN. 1942. Florida dunes and scrub, vegetation and geology.
Florida Geol. Surv., Bull. No. 23, pp. 1-154.

LArEssLE, A. M. 1942. The plant communities of the Welaka area. Uni-
versity of Florida Press, Biol. Sci. Series, vol. 4, no. 1, pp. 1-143.

———. 1958a. The origin and successional relationships of sandhill vegeta-
tion and sand pine scrub. Ecol. Monog., vol. 28, pp. 361-387.

———. 1958b. A report on succession studies of selected plant communities
on the University of Florida Conservation Reserve, Welaka, Florida.
Quart. Jour. Florida Acad. Sci., vol. 21, no. 1, pp. 101-112.

LirtLe, E. L., Jn., anp K. W. Dorman. 1952. Geographic differences in
cone opening in sand pine. Jour. Forestry, vol. 50, pp. 204-205.

MacNem, F. S. 1950. Pleistocene shore lines in Florida and Georgia.
U. S. Geol. Surv., Prof. Paper 221-F, pp. 95-107.

Miniter, R. 1950. Ecological comparisons of plant communities of the
xeric pine type on sand ridges in central Florida. Unpublished M. S.
thesis, University of Florida.

Mutvania, M. 1931. Ecological survey of a Florida scrub. Ecology, vol.
12, pp. 528-540.

Nasu, G. V. 1895. Notes on some Florida plants. Bull. of the Torrey
Bots Clube wooly 22.n04 appeals

VIGNOLES, CHARLES. 1823. Observations upon the Floridas. Bliss, New
Vorkencia Of pp:

WespseER, H. J. 1935. The Florida scrub, a fire fighting association. Amer.
Jour. Bot., vol. 22, pp. 344-361.

Wuitney, M. 1898. The soils of Florida. U. S. Dept. Agri., Bull. 13,
pp. 14-27.

Department of Zoology, University of Florida, Gainesville, Flor-
ida 32601.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968)

Sea Turtle Nest Survey of Hutchinson Island, Florida
Rosert A. Roura

EACH summer a considerable number of sea_ turtles come
ashore on the beach of Hutchinson Island, near Stuart, Florida,
to lay their eggs (Fig. 1). The bulk of the nesting population
consists of loggerhead turtles, Caretta caretta caretta (Linne); how-

Fig. 1. A loggerhead turtle nest.
ever, a few green turtles, Chelonia mydas mydas (Linne), have
also been reported as nesting here.

Raccoons frequently dig up the nests and local residents have
estimated that between 50 and 75 per cent are destroyed. This
study was undertaken to obtain an estimate of the total number
of nests and eggs and also the extent of nest damage by preda-
tion.

METHODS AND MATERIALS

Three one-mile stretches of beach were designated as study
areas (Fig. 2) and were checked during the day on Monday,

288 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

CaN
- FT. PIERCE INLET

2

——>2

ATLANTIC

OO Oo?

OCEAN

HUTCHINSON
ISLAND

Ooo 0.0

O70 e- Oo
a) oe eee

5017006 6 O

COTO oO O65 3.6508

MILES

ST. LUCIE
INLET

Fig. 2. Hutchinson Island, Florida, showing study areas.

Wednesday, and Friday of each week in May, June, July, and
August of 1967. A beach buggy was used for transportation and
was driven up each nest trail to count and mark the nest. The
new nests in each area were counted as well as those disturbed by

Routa: Sea Turtle Nest Survey 289

raccoons. The moon phase and the position of the nest relative to
the dune front were recorded. The dune front was defined as the
seaward face of the dunes where erosion takes place during
periods of extreme high tide.

RESULTS AND CONCLUSIONS

The turtles began nesting during the first week in May and
continued through the last week in August. The peak of activity
was reached during the last week in June, shortly after the sum-
mer solstice. Figure 3 illustrates that nesting activity appeared to

50

—— NO. OF NESTS
---- NO. DESTROYED

40

NESTS
wi
°o

NO. OF
N
°o

A
tm)

4
-
Foor Linas PA,

(0) eos. eos x / . AN
MAY | JUNE 2 JULY 3 AUG. 2 AUG. 30

Fig. 3. Sea turtle nesting and nest predation.

be directly related to increasing and decreasing day length. The
greatest nesting activity occurred in the last two weeks of June
and the first two weeks of July. Nesting activity during the
month of June was slightly higher than in July (Fig. 4).

The total number of nests which occurred in area A was found
to be 209. The height of the dune front of area A ranged from
0 to 5 ft. Tall Australian pines, growing continuously along the
beach behind the dune front, presented a high, dark skyline.
Area B had a total of 308 nests. The dune front here averaged

290 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

0 to 3 ft. with a predominant vegetation of sea grapes which
presented a lower and brighter skyline than area A. A total of
188 nests were observed in area C (Fig. 5). The height of the

300 300
Ww
- o
tw 7200
W200 a
2
irs
°o uw
3 (o)
2 2
100 100
0 0
MAY JUNE JULY ~— AUGUST AREA A AREA B AREA C
Fig. 4. Nesting by month. Fig. 5. Nesting by area.

dune front of this area ranged from 5 to 10 ft. The predominant
sea grapes were considerably taller than those in area B and
presented a high, dark skyline. The only discernible difference in
the areas was the varying skyline and this may indicate that the
turtles are attracted to a relatively bright beach on which to nest.

No apparent correlation between nesting activity and moon
phase could be established.

A definite nest site preference relative to the dune front was
noticed since 689 nests occurred below and 16 nests occurred above
the dune front. It must be noted that in area A the dune front
was apparently too high to permit any turtles to climb over it.
The other areas have a lower dune front and the turtles were able
to crawl over without apparent difficulty.

An effort was made to determine the total numbers of nests
and eggs for the entire 22.4 mile beach on Hutchinson Island.
Table 1 gives the results obtained by using the formula, estimated
no.: no. counted: miles surveyed: miles of beach. Carr (1952)

Routa: Sea Turtle Nest Survey 291

lists an average of 120 to 130 eggs per nest for Atlantic loggerhead
turtles. Since that species is the predominant one on the island,
the egg number estimates were based on an average of 125 per
nest.

Predation by raccoons was easily recognized from the egg shells
scattered around the nest area (Fig. 6). During the month of

Fig. 6. A turtle nest destroyed by raccoons.

May, this predation was almost nonexistent since only one nest
was destroyed in the study areas. In June, eight nests were de-
stroyed and July had the greatest incidence of predation with 40
nests being damaged. August showed a decline with only six
nests destroyed (Fig. 7). Area A had the greatest incidence of
predation for the entire nesting season with 40 disturbed nests.
Area B followed with 13 nests damaged and area C showed only
two destroyed nests (Fig. 8).

As area A is the least used by fishermen and bathers, this
might account for the heavy predation by raccoons. Area B is
used more heavily and area C is used to the highest degree and

292 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Sy
{o)
b
oO

aQ (a)
: :
© ©
- -
230 30
Qa a
2) wn
E bE
72) n
W WwW
2 2
520 520
° °
2 2
lo 10
0 fe)
MAY JUNE JULY AUGUST AREA A AREA B AREA C
Fig. 7. Nest predation by month. Fig. 8. Nest predation by area.

contains a public beach. The little-used areas probably have a
higher raccoon population. The fact that predation did not occur
to any great extent until July may reflect the possibility that the
raccoons require some time to become oriented to the presence of
the eggs on the beach. It is possible that animals other than
raccoons are responsible for some of the predation but only the
tracks of raccoons were seen around the nests examined. Some
poaching was noticed outside of the study areas but this was kept
to a minimum by conservation officers. Total predation was low
since only 7.8 per cent of the counted nests were destroyed. It
was also noted that the raccoons did not necessarily dig up only
freshly made nests. Some of the nests were several days old when
they were destroyed.

Several unusual nests were noticed during the course of this
project. The largest nest of the loggerhead type was observed on
May 12, with the tracks measuring 5 ft. 9 in. across. The smallest
was found on July 12, with the tracks measuring 24 in. across. In
area B, on May 22, an unusual nest was noted in that the flipper
marks were much closer together on the tracks than is the case

Roura: Sea Turtle Nest Survey 293

with a loggerhead track. The nest area also contained a pit which
is not characteristic of loggerhead nests. Another nest of this same
type was found in area C on August 12, and both closely resembled
Carrs (1952) description of a green turtle nest. A green turtle
nest had been reported from this area (Carr and Ingle, 1959)
and to determine if these nests might also be those of green
turtles, the 74 eggs from the August 12 nest were dug up and
reburied in an enclosure. The first of the eggs hatched after 71
days and verified the tentative identification. If the earlier nest
was also that of a green turtle, it can be estimated that approxi-
mately 15 green turtles nested along the entire length of Hutchin-
son Island. The validity of this estimate is enhanced by the obser-
vation of several of these green turtle nests outside the study
areas.

TABLE 1
Estimates for sea turtle nests and eggs
May June July August Total
Total nests counted# 94 285 262 64 705
Destroyed nests counted I) 8 AO 6 55
Estimated nests» 702 2,128 1,957 478 D260
Estimated eggs 87,750 266,000 244,625 59,750 658,125
Estimated nests destroyed 8 60 299 45 412
Estimated eggs destroyed 1,000 7,500 31,370 5,625 51,500

a Total counted in 3 miles of beach.
» Estimated for 22.4 miles of beach.

SUMMARY

An estimated 5,265 nests containing about 658,125 eggs oc-
curred on Hutchinson Island from May through August, 1967.
Through similar projection of the sample data, a total of 51,500
eggs were believed to have been destroyed by raccoons (7.8 per
cent). Peak nesting activity occurred during the last week of
June. The nesting turtles appeared to prefer a relatively bright
beach. No significant relationship between nesting activity and
moon phases was found. Most of the nests occurred below the
dune face and only a few turtles climbed over the dunes. Signi-
ficantly, two green turtle nests were found in the study areas and
an estimated 15 green turtles probably nested on Hutchinson Is-

land.

294 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

LITERATURE CITED

Carr, Arcuie. 1952. Handbook of turtles. Comstock Publ. Assoc., Ithaca,
New York, xv + 542 pp., 82 pls.

Carr, ARCHIE, AND Ropert M. INGLE. 1959. The green turtle (Chelonia

mydas mydas) in Florida. Bull. Mar. Sci. Gulf & Carib., vol. 9, no.
3, pp. 315-320.

Florida Board of Conservation Marine Laboratory, St. Peters-
burg, Florida. Contribution No. 114.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968)

Yolk Pigmenting Value of Dried Kenaf Tops
Jack L. Fry, GEorcE M. Herrick, AND R. H. Harms

KenaFr (Hibiscus cannibus L.) is a tall, woody, annual plant
which produces a long, unbranched stem when grown in dense
stands. It is native to Africa where it is used for both food and
fiber. It is widely cultivated for fiber production in the tropics
and subtropics, including many Latin American countries.

There has recently been much interest in the use of kenaf in
animal feeds and as a pulp source for paper manufacture. New
varieties of kenaf have been developed by Wilson et al. (1965).
When Everglades-41 variety was at about the 6 foot growth stage,
it was found by Wing (1967) to make a silage comparable to
that of most high quality roughages.

In paper manufacturing it is probable that the leaves will be
recovered, dried, and made available to the feed industry as kenat
leaf meal. It has been suggested that this meal might replace
alfalfa as a xanthophyll source in the diets of laying hens and
broilers. The study herein reported is an investigation of the po-
tential of kenaf as a pigmenter of egg yolks.

MATERIALS AND METHODS

Dehydrated kenaf tops were used in two trials to replace
alfalfa in the diets of egg production type hens. The kenaf meal
was prepared from plants approximately 125 days old. The top

TABLE 1
Basal diet composition
Ingredient Percent
Degerminated white corn meal 68.00
Soybean meal (50 percent protein ) 20.80
Ground limestone 5.90
Defluorinated phosphate 1.95
Salt 25
Vitamin and trace mineral mix? 50
Sand 2.60

1 Supplied per kilogram of diet: 6,600 I.U. vitamin A, 2,200 I.C.U. vitamin
Ds, 500 mg. choline chloride, 40 mg. niacin, 4.4 mg. riboflavin, 13 mg. panto-
thenic acid, 22 mcg. vitamin Biz, 125 mg. ethoxyquin, 20 mg. iron, 2 mg. cop-
per, 200 mcg. cobalt, 1.1 mg. iodine, 100 mcg. zinc, 71 mg. manganese, and
2.2 mg. menadione sodium bisulfite.

296 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

36 inches of the plants (Everglades-41 variety) were dried and
then ground in a Wiley mill. The composition of the pigment-free
basal diet is shown in Table 1. In the first trial the kenaf and
alfalfa leaf meal (20 per cent protein) were incorporated into the
diet at 2% per cent and 5 per cent each. When the kenaf and
alfalfa were added to the diet, adjustments were made in the
level of corn, soybean meal and sand in order to keep the diets
iso-caloric and iso-nitrogeneous. The nutrient analysis of ingredi-
ents according to Maddy et al. (1963) was used as a basis for
these formulations. A synthetic pigmenter, beta-apo-8’-carotenal,
was used as a “control” at levels of 30, 60, 90, and 120 grams per
ton of basal.

Eighty individually caged hens which had been in production
approximately 10 months were depleted of pigment by feeding
the basal diet (Table 1) for 35 days. At this time yolk color was
evaluated visually using a color rotor (Heiman and Carver, 1935)
and found to have an average score of 5.5. The hens were then
randomized into groups of ten and one group placed on each of
the treatments. Visual color rotor scores were obtained on all
eggs collected during a 14-day repletion feeding period.

In the second trial the alfalfa and kenaf were included in the
test diets on the basis of analyzed xanthophyll content. Kenat
meal (71 mg xanthophyll per pound) was used at 5 per cent of
the diet whereas alfalfa leaf meal (105 mg xanthophyll per pound )
was used at 3.38 per cent of the diet. Levels of 30, 60, and 90
grams of beta-apo-8’-carotenal were fed; twenty pigment-depleted
hens were on each of the five test diets. Yolk color scores were
obtained as previously described.

RESULTS AND DISCUSSION

Maximum yolk pigmentation occurred between the 10th and
14th day after repletion feeding began. The maximum average
scores for each test diet in both trials are presented graphically
in Figures 1 and 2.

Color rotor scores of eggs from hens fed the diet containing
2% per cent alfalfa equalled that of the group fed 5 per cent
kenaf in the first trial (Fig. 1). Color scores of eggs from hens
fed 5 per cent alfalfa were two units higher than these groups.
When comparing the values obtained when increasing levels of

Fry er Au.: Yolk Pigment and Kenaf 297

iS Trial #|
oe e

5 % Alfalfa

5% Kenaf
‘Ss * and
24 % Alfalfa

2+ %Kenaf >

Maximum Yolk Color Scores
wo

CuemiOmmrzone 30) 40 «50 60'° 70° 80’ 90° 100 I10 120
gM/ton Beta apo-8-carotenal

Taye 1. Yolk pigmentation by beta-apo-8’-carotenal, alfalfa, and kenat
trial 1).

beta-apo-8’-carotenal were fed it appears that utilization of pig-
ment in the feed does not result in a linear relationship with the
amount of pigment present in the feed.

A comparison of the values obtained for the two different
levels of kenaf and alfalfa with their equivalents in pigmenting
value by beta-apo-8’-carotenal suggests a similar non-linear rela-
tionship of pigmentation and ingredient level. Higher levels of

298 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Trial #2

€ 3.38 % Alfalfa

5% Kenaf >

ro) =

o

Maximum Yolk Color Scores
@

~s
wee ae ae

0 10 20 30 .e) 50 60 70 80 90 100—s 110 120

gM/ton Beta -apo-8- carotenal

Toe 2. Yolk pigmentation by beta-apo-8’-carotenal, alfalfa, and kenaf
trial 2).

these ingredients would no doubt be less effective per xanthophyll
unit.

Although kenaf was only one-half as effective as alfalfa when
used on an equivalent weight basis, xanthophyll determinations
indicated that 3.38 per cent alfalfa should be equivalent to 5 per
cent kenaf in the diet. When these levels were used in trial 2
(Fig. 2) yolk color scores of 11.8 and 11.3 were obtained for

Fry er AL.: Yolk Pigment and Kenaf 299

alfalfa and kenaf, respectively, indicating a less efficient utiliza-
tion of the xanthophyll in kenaf.

It is apparent that the laying hen can utilize the xanthophyll
in kenaf; however, the xanthophyll level in kenaf is not as high
as in alfalfa leaf meal and it appears that the biological activity is
not as high. It should be pointed out that the kenaf used in these
studies was from the top 36 inches of plants of 125 days maturity.
It has been suggested that a better source for this purpose might
be (1) the top 24 inches of the plant, (2) stripped leaves of
plants of this maturity, or (3) the entire plant at about the 6 foot
stage of maturity. It has been estimated (Killinger, personal com-
munication) that approximately two tons of oven dry kenaf meal
could be produced per acre from kenaf at the 6 to 7 foot growth
stage.

All of these three sources would be expected to have higher
xanthophyll and protein content and less fiber than the sample
used in this study. Also, the sample was stored (55-60F.) for
approximately 3 months prior to use; since no anti-oxidant was
added there may have been a significant loss of xanthophyll.

SUMMARY

Two experiments were conducted with egg production type
hens to evaluate dehydrated kenaf tops as a source of xanthophyll
in poultry feeds. Kenaf at 5 per cent of the diet produced
acceptable yolk pigmentation but was not as effective as alfalfa
on either a weight basis or a xanthophyll content basis.

Further study of other kenaf sources is warranted since there
is the possibility of kenaf being readily available as a feed ingredi-
ent for yolk pigmentation.

ACKNOWLEDGMENTS

The kenaf for this study was supplied by Dr. R. V. Allison,
Everglades Experiment Station, Belle Glade, Florida, 33430, and
the beta-apo-8’-carotenal by Hoffman-LaRoche, Inc., Nutley, New
Jersey. Appreciation is expressed to Mr. T. P. Moffses for his
assistance and to Mrs. Mary Maud Sharpe for the xanthophyll
determinations.

300 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

LITERATURE CITED

HEIMAN, V., AND J. S. Carver. 1935. The yolk color index. U. S. Egg
Poultry Magazine, vol. 41, no. 8, page 40-41.

Mappy, K. H., R. B. Graincer, W. A. DuDLEY, AND F. Pucnau. 1963. The
application of linear programming to feed formulation. Feedstuffs,
vol. 35, no. 15, pages 28-30, 70-73.

Witson, F. D., T. E. Summers, J. F. JoyNeER, D. W. FISHLER, AND C. C.
SEALE. 1965. “Everglades 41” and “Everglades 71”, two new varie-
ties of kenaf for fiber and seed. Cir. S-168, Florida Agr. Exp. Sta.

Wine, J. M. 1967. Ensilability, acceptability and digestability of kenaf.
Feedstufts, vol. 39, no. 29, page 26.

Department of Poultry Science, University of Florida, Gaines-
ville, Florida 32601. Florida Agr. Exp. Sta. Journal Series No. 2949.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968 )

Persistent Viremia in EIA Infected Horses

J. H. Fiynn, G. H. WappeELL, anv V. R. SAuURINO

In a few virus diseases one of the characteristics associated with
the host-virus relationship is a viremia. This viremia is, however,
short in duration in most cases as exemplified by measles virus and
others. The virus disease of the horse, equine infectious anemia
(EIA), has been considered by the field of veterinary medicine as
unique due to a persistent viremia. Numerous clinical studies have
shown the presence of virus within the peripheral blood stream for
extended periods of time (Dreguss and Lombard, 1954; Mott et al.,
1947). Clinically the disease may establish itself as an acute infec-
tion which terminates fatally or manifests itself as a subacute or
chronic infection. This subacute or chronic infection is the form
in which prolonged viremia occurs, and is termed clinically as a
“carrier state.

In a so-called “carrier” horse, the disease equine infectious
anemia generally manifests itself clinically with repeated febrile
episodes about every 15-30 days. The only method of confirming a
clinical diagnosis of EIA in the chronically infected animal was,
until 1966, by injecting blood from the clinically suspected animal
into a normal, uninfected, recipient horse. Establishment of disease
in the recipient animal was evidence of infection in the suspect
horse.

In 1966, Saurino et al. described a serologic test for the deter-
mination of the existence of virus in the peripheral blood of the
“carrier horse as well as the acutely infected animal. This serologic
test was the immune adherence (JI.A.) test previously described by
Nelson for the detection of Treponema pallidum (1953). The IA
test has proven to be the first reliable and sensitive diagnostic test
for the disease. The test is, however, measuring only the soluble
antigens of viral products after the virus particles have been re-
moved from the serum by absorption with red blood cells.

This study was undertaken to provide a serologic test for the
measurement or viral antigens, both soluble and insoluble in origin,
found in the serum of horses infected with EIA. The test used was
the indirect passive hemagglutination test (IPHA). This serologic
test employed purified gamma globulin coated onto sheep red blood
cells by the method of Stavitsky (1954). In support of the IPHA

302 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

the IA was used to measure only the soluble viral products found
in the sera of infected horses. The IPHA is a sensitive and reliable
test which has been used in various immunologic problems con-
cerning virus infection (Lefkowitz, 1966; Smith, personal commu-
nication ).

All horses used in this study were thoroughbreds. The animals
infected with EIA consisted of three clinically different types. The
virulent, acute type of infection (V) was consistently caused by the
classical Wyoming strain of EIA virus. The virulent carrier (VC)
was a field case which was undetectable clinically except for inter-
mittent fever. This strain of virus, when animal tested, rapidly
killed two test horses. A mild carrier (MC) was also a field case
and was clinically undetectable except for intermittent febrile epi-
sodes. When the MC horse was animal tested one horse failed to
show signs of infection and the second test animal became a
“carrier.” The animals employed for testing the MC strain of virus
were MCT, and MCT,. Sera from twenty-four normal thorough-
bred horses were employed as clinically normal animals and there-
fore served as controls.

ANTIGENS

The antigens used in all serologic tests, hemagglutination, im-
umne adherence and indirect passive hemagglutination, were horse
sera. The sera was removed from clotted bloods by centrifugation
within 24 hours after obtaining the sample. All blood cells were
removed from the sera to avoid hemolysis. The sera were stored at
-20°C, and were heat inactivated at 56°C. for 30 minutes. This
was done to inactivate complement which may have reacted pro-
ducing positive readings.

The antigen used in the IA test was absorbed twice with 0.2 ml
packed human group O red blood cells (RBC’s) per ml of sera.
Each absorption was for 30 minutes at room temperature. The sera
used in the HA and IPHA were heat inactivated but not absorbed.

The sources of antigen were sera from the three clinically dis-
tinct types of EIA; virulent (V) type, virulent carrier (VC) type
and mild carrier (MC) type. Sera were also obtained from the
mild carrier’s test horses (MCT, and MCT.), and a normal group
of horses.

The virulent type of sera was obtained from a horse which had

FLYNN ET AL.: Viremia in Horses 303

been inoculated with the Wyoming strain of virus. This horse was
a thoroughbred gelding, 3 years of age. The horse showed all the
characteristic clinical signs of the disease after injection and at
necropsy the spleen was enlarged approximately eight times normal
size. Based on the clinical signs and symptoms the horse was con-
sidered to be acutely infected with a virulent strain of the virus.

The virulent carrier (VC) type of sera was obtained from a
thoroughbred gelding, approximately 8 years of age. The disease
was diagnosed on the basis of clinical signs, recurrent fever, and
inoculation of test horses. The VC horse, although clinically ap-
pearing normal except for recurrent fever cycles, produced acute
terminal cases of the disease in two test horses following injection
of this sera. Based on the clinical response of the test horses, this
horse was considered a carrier of the virulent strain of the virus.

The mild carrier (MC) type of sera was obtained from a thor-
oughbred stallion, 3 years of age. This type of the disease clinic-
ally showed only intermittent fever with no other significant signs.
Sera from this horse was injected into two thoroughbred test
horses, MCT, and MCT.). One of the test animals (MCT,)
showed acute signs of the disease within two weeks after injection.
Test animal (MCT:) showed no clinical signs throughout the test
period. This animal (MCT:) was subsequently challenged with
the VC strain of virus and developed an acute infection followed
by development of the carrier state. Test animal (MCT,) also
developed a mild type of carrier state with only the recurrent fever
as a clinical sign.

The clinically normal horses used in all these tests were thor-
oughbreds of various ages. All the normal horses were in training
for racing and none exhibited clinical signs under this stress.

ANTIBODY

The antibody used in both the IA and IPHA was obtained from
the sera of infected horses (VC, MC, and MCT,) and from normal
horses (N). The 19S antibody component was obtained from these
horse sera by fractionation. The sera was fractionated by ion ex-
change columns (DEAE), (Allen, et al., 1955). Each serum was
dialized against 0.015 M tris buffer (2-amino-2-hydroxymethyl-l,
3-propanediol) at 4°C overnight. The columns (20 cm long, 0.8
em inside diameter) used for fractionation were packed with

304 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

DEAE-cellulose (N,N-diethylaminoethyl ether cellulose). The di-
alized sera was added to the top of the column. A two step salt
gradient (NaCl) was used to fractionate the sera. The first gradient
was obtained by mixing 0.12 M NaCl in 0.015 M tris buffer with
a 0.00 M salt solution in buffer and then passing through the seed-
ed column. The second step consisted of 0.4 M NaCl in 0.015 M tris
buffer mixing with 0.04 M NaCl in 0.015 M tris buffer solution and
then passing through the same column. The solutions passing
through the column under constant air pressure eluted off the vari-
ous proteins by ion exchange. After passing through the column
the solutions were passed into a continuous flow cell in a Gilford
recording spectrophotometer (model 2000) at a wave length of
280 mp. At this wave length the solutions were measured for pro-
teins and the quantity of absorbance was read continuously and
was recorded in a characteristic pattern. The antibody portion of
the serum being fractionated was collected and stored at -20°C in
5 ml fractions. Storage in 5 ml aliquots was done to reduce degen-
eration of the large gamma-globulins (19S) to smaller gamma-
globulins (7S). The antibody used in the IA and IPHA was both
unconcentrated, and concentrated 5:1 with polyethylene glycol.

The antibody fractions were characterized by sucrose density
gradient ultracentrifugation. The 19S and 7S antibody fractions of
sera were identified by comparison with a standard protein with a
known S value (S refers to Svedberg units). The density gradients
were prepared by layering 6.5 ml each of four sucrose solutions
(30, 25, 20, and 15 per cent) into a 35 ml centrifuge tube, (Beck-
man, polyethylene). The solution with the highest density was
added first followed by the progressively less dense solutions. The
gradients were allowed to equilibrate over-night at 4°C to establish
a linear gradient. After equilibration, 2 ml of the concentrated pro-
tein fractions obtained by serum fractionation were layered on sep-
arate linear gradients. The marker protein bovine serum albumin
(BSA), 90 pg/ml and 2ml of 15 per cent sucrose was mixed with
the antibody proteins. The gradients were centrifuged at 25,000
rpm for 18 hours in an SW-25 swinging bucket rotor (Beckman
model L preparative ultracentrifuge ). ,

The density gradients were removed from the centrifuge after
18 hours, pierced through the bottom of the tube and read by con-
tinuous flow by the Gilford recording spectrophotometer (Gilford

FLYNN ET AL.: Viremia in Horses 305

Instrument Laboratories, Inc., Oberlin, Ohio) at a wave length of
280 mp» for proteins. The S values were determined by a compari-
son with the standard BSA (Wood, 1964).

The amount of protein in the gamma-globulin fractions used
in the IA and IPHA was determined by a phenol test. The pro-
cedure was that of Bradshaw (1966). Four dilutions of each of
the fractions used (1:40, 1:160, 1:640, and 1:2560) was tested.
Quantitation was done with a spectronic 20 (Beckman) colori-
meter at a wavelength of 740 mp. The amount of protein was de-
termined by a standard BSA curve.

SEROLOGY

Hemagglutination: Each of the animal sera was tested by
hemagglutination following heat inactivation. The sera were dilut-
ed initially 1:10 and then 2 fold dilutions to 1:20,480 in normal
saline. The sera were diluted in a total volume of 0.2 ml and in
separate experiments 0.05 ml of 1.5 per cent human group 0 RBC'’s
and sheep RBC’s were added. The test was read after settling at
room temperature for 1-1% hours. The test was read as +, +, or
— agglutination. The controls were 0.2 ml saline and 0.05 ml of
RBC’.

Immune Adherence: The immune adherence test used was that
described by Nelson (1953). The test was used due to its high
sensitivity for antibody 0.001 to 0.003 pg antibody nitrogen/ml
(Nelson, 1953). The test used heat inactivated and twice absorbed
sera as the antigen. The sera were diluted in Verona buffer saline
(VSB). The initial dilution was 1:10 followed by four-fold dilutions
to a dilution of 1:163,840 in a volume of 0.1 ml. The optimal anti-
body dilution for use was determined by a block titration. A total
volume of 0.1 ml of antibody was added to each of the tubes con-
taining 0.1 ml of the serum dilution and the mixture was incubated
for 20 minutes at 37°C. After incubation 0.2 ml of guinea pig
complement (1:100) was added and the mixture was further incu-
bated for 10 minutes at 37°C. After the second incubation 0.05 ml
of human group 0 RBC’s was added, the tubes were shaken and
incubated at 37°C for another 10 minutes with shaking. The re-
action mixture was then removed from incubation and allowed to
settle at room temperature for 24% hours. The patterns on the bot-
tom of the tubes were read on the basis of 4+ agglutination to 0

306 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

for no agglutination. Controls for the IA test included antigen con-
trols with and without complement, antibody controls with and
without complement, complement control and cell control.

Indirect Passive Hemagglutination: The indirect passive hemag-
gluttination test was adapted from the procedures of Boyden
(1951), Smith (personal communication), Cook (1964), Faucon-
nier (1965), and Lefkowitz (1966). This test was also chosen be-
cause of its high sensitivity to antibody nitrogen (0.003-0.005 pg
antibody nitrogen/ml). Preserved sheep red blood cells (Hyland
Laboratories ) were washed four times in 0.15 M phosphate buffered
saline (PBS), pH 7.2 and resuspended to a concentration of 1.5
per cent. The cells were mixed with an equal volume of tannic
acid solution in PBS, pH 7.2 at a concentration of 1:20,000 and
allowed to stand at 4°C for 15 minutes. The sensitized cells were
centrifuged and washed once in PBS at pH 7.2. The cells were
then resuspended to the original volume (1.5 per cent in PBS, pH
6.4. It was of utmost importance that the cells were coated with
antibody (19S or 7S) immediately after sensitization with tannic
acid.

One volume of antibody (concentrated 5 times with polyethyl-
ene glycol and dialized against PBS, pH 6.4 overnight at 4°C) and
an equal volume of sensitized cells (1.5 per cent) were mixed in
one-half the volume PBS, pH 6.4 (earlier experiments employed
19S antibody directly from the column). The mixture was incu-
bated at room temperature for 15 minutes. The coated cells were
washed once in a solution of normal heat inactivated rabbit serum
at a dilution of 1:50 in PBS, pH 7.2 (NRS, Hyland Laboratories ).
The cells were then resuspended to original volume (1.5 per cent)
ma INU gus) 160), qolsl Wo

The test antigen for IPHA was horse sera which had been heat
inactivated but not absorbed. The antigen was diluted 1:10 initial-
ly and then in four-fold dilutions to 1:63,840 in PBS, pH 7.2 con-
taining NRS at a 1:50 dilution. A final volume of 0.5 ml of each
serum dilution was mixed with 0.05 ml of coated sensitized cells.
The mixture was then incubated at 37°C for 30 minutes with shak-
ing. After incubation the tubes were allowed to stand at room
temperature for 2 to 2% hours before reading. The agglutination
pattern on the bottom of the tubes was read and scored on the
basis of 4+ for total agglutination to 0 for lack of agglutination.

FLYNN ET AL.: Viremia in Horses 307

PROTEIN PATTERN OBTAINED
WITH DEAE-CELLULOSE

i \ = 280 mp

ISS

ABSORBENCY

0.00M NaCl /0.12 M NaCl 0.04M NaCl /0.40 M NoCl

Fig. 1. Elution pattern of serum proteins from a horse infected with equine
infectious anemia. The serum proteins were eluted from a column of DEAE-
cellulose with increasing concentrations of NaC]. Proteins were quantitated by
absorbance at a wave length of 280 My.

The controls in these experiments consisted of NRS coated tanned
cells and antigen, NRS coated cells and PBS, pH 7.2; antibody
coated cells and PBS, pH 7.2; tannic acid cells and PBS, pH 7.2,
and uncoated cells in PBS, pH 7.2.

EXPERIMENTAL RESULTS

Serum Fractionation: Fig. 1 depicts a typical protein pattern
acquired by elution of sera from DEAE-cellulose with increasing
concentrations of NaCl. The first portion of the pattern includes
the 7S gamma-globulin proteins, which was eluted by the 0.00 M
NaCl to 0.06 M NaCl gradient. The second portion of the pattern

308 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

A = 280 mp BOVINE SERUM
ALBUMEN

GAMMA GLOBULIN
S= 7.08

ABSORBENCY

MENISCUS

30% SUCROSE GRADIENT IS%

Fig. 2. Characterization of 7S gamma-globulin by sucrose density gradient
centrifugation. Sucrose gradients (15/30 per cent w/v) were centrifuged at
25,000 rpm’s in an SW-25 rotor for 18 hours. Proteins were quantitated by
absorbance at 280 my.

was obtained by the 0.04 M NaCl to 0.2 M NaCl gradient. This por-
tion includes the 19S gamma-globulin which was collected and
utilized herein for serologic tests.

Antibody Characterization: The protein removed from the horse
sera was characterized by both the phenol test and sucrose density
gradient centrifugation. The results of the nitrogen determinations
indicated that the quantity of protein per ml of column fraction
varied with the column. The range of protein gamma-globulin as
indicated by the phenol test was between 24.8 and 30 »g per ml.

The density gradients showed the protein peaks removed by
fractionation to contain 7S gamma-globulins (Fig. 2) and 19S
gamma-globulins (Fig. 3). The 7S fraction was obtained from the
first protein peaks eluted from the DEAE and the 19S fraction was
obtained from the last protein peak to be eluted. In Figs. 2-3, the
large peak is the 4.68S bovine serum albumin (BSA, 90 g/ml)

FLYNN ET AL.: Viremia in Horses 309

which acted as the marker protein. The sedimentation constant of
the two unknown peaks in Figs. 2-3 were calculated by the pro-
portion:

distance from meniscus (A) _ Sedimentation constant (A)
distance from meniscus (B) Sedimentation constant (B)

Figure 2: Sedimentation Constant

Dg = 35 mm 35 mm _ 4.68 S
Dy, = 93 mm 53 mm SE
SF =4.68 S

Figure 3: Sedimentation Constant

D, = 32 mm 32 mm = 4.68 S
Dy 129 mm 129 mm SE

Sg = 4.68 S

Spy = 285 Sia 18 .84

The 19S gamma-globulins were stored in small aliquots to re-
duce the break down of the 19S to 7S. Premininary tests demon-
strated that 7S antibody did not react well in the IA test but did
react in the IPHA.

Hemagglutination: The hemagglutination test (HA) was run
on all serum samples, in separate experiments with both human
group 0 red blood cells and sheep blood cells. The HA indicated
the need for absorption of the serum before using the human cells
in the immune adherence test; however, results of the HA using
the sheep RBC’s indicated no significant agglutination (titer<
1:10). The lack of agglutination by sheep RBC’s allowed the use
of unabsorbed serum in the IPHA.

Immune Adherence and Indirect Passive Hemagglutination: In
previous work with the animal test it had been established that a
horse once infected with the virus of EIA would either succumb
to the infection or become a “carrier”. In earlier work Saurino et
al. showed that both the IA and IPHA tests were capable of de-
tecting single positive sera from horses infected by virus causing
each of the three clinical types of infection.

Immune adherence and indirect passive hemagglutination tests

310 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

A =280mp BOVINE SERUM

ALBUMIN
S=4.68

GAMMA GLOBULIN
S=18.84

ABSORBENCY

MENISCUS

30% SUCROSE GRADIENT IS%

Fig. 3. Characterization of 19S gamma-globulin by sucrose density gradient
centrifugation. Sucrose gradients (15/30 per cent w/v) were centrifuged at
25,000 rpm’s in a SW-25 rotor for 18 hours. Proteins were quantitated by
absorbance at 280 my.

were performed on sera from the mild carrier (MC) horse cellected
over an 18 day period in an attempt to show the fate of the virus
and virus products during one eighteen day fever cycle. Over the
18 day period the temperature was taken twice daily, the MC horse
showed no characteristic clinical signs of disease except the febrile
peaks every 12-13 days. In Fig. 4 the febrile response over an
eighteen day period is compared with levels of virus and virus
products as indicated by IPHA and IA. The figure depicts a com-
paratively low level of antigen present at the onset of the cycle.
A gradual rise leads to a leveling-off of the antigen levels. This
leveling is concomitant with a leveling-off in the temperature of
the animal. The temperature response occurs two days after the
highest virus titer. The onset of fever also coincides with a rapid
drop in the titers as indicated by both tests.

The comparison of virus titers with temperatures of MCT,

FLYNN ET AL.: Viremia in Horses 311

L——\ Temperature 106
Oe==-0 IA

ag ae a
or f

~
ae
&
°F

TEMPERATURE

SEROLOGIC TITERS

Cn mn SuiAniSmmicy ir) 18° 19) 20) 2 22° 23.24 25 26 27 28 1
DATE OF BLEEDING

Fig. 4. Comparison of serologic and clinical responses of a mild carrier
(MC) of equine infectious anemia over an 18 day period. Results of immune
adherence and indirect passive hemagglutination tests in relation to the febrile
response.

2—— Temperature
Oe=--— IA
oa [PHA

TEMPERATURE °F

SEROLOGIC TITERS

cues a noo 8) 9) ION 2) 13) 14) 1S) | 1G) sli%. 18) 19) 120) “2l 22) (23
DATE OF BLEEDING AFTER INJECTION

Fig. 5. Comparison of serologic and clinical responses of a mild carrier test
horse (MCT,) of equine infectious anemia over an 18 day period. Results of
immune adherence and indirect passive hemagglutination tests in relation to
the febrile response.
showed for the first ten days a normal temperature and lack of anti-
gen (Fig. 5). The figure compares the titer of antigen with tem-
perature over a twenty-three day period from preinoculation
through one fever cycle. On the eleventh day following infection,
antigen was detectable by the IPHA and by the thirteenth day,

312 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

both IA and IPHA indicated the presence of antigen. The antigen
level gradually rose until a febrile response occurred on the fif-
teenth and sixteenth days. An antigen peak was reached on the
nineteenth day.

The MCT, horse was compared to the temperatures taken over
the same twenty-three day period as MCT,. However, no tempera-
ture changes were noted and no virus titers were obtained with
either the IA or IPHA.

The twenty-four “normal” sera were run as controls and no sig-
nificant titers were obtained.

DISCUSSION

The virus disease of equine infectious anemia (EIA), like other
virus diseases, establishes a parasitic relationship with the host
which may result in either death of the host or a prolonged carrier
state. Unlike most virus diseases, the virus of EIA is found in the
sera continually during the life of the infected animal. In develop-
ment of the IPHA test for use to detect the EJA virus in the sera,
a new approach was followed. Purified antibody (19S) obtained
from infected horses was employed to detect the presence of virus
or soluble virus products in the sera of infected animals. The re-
sults of the correlation of clinical signs and daily temperatures,
with the levels of virus over an extended period of time, revealed
in the case of the MC horse a_ fluctuation of levels of virus and/or
virus products. The fluctuation can be interpreted as a possible
course of the viral infection; the virus being influenced by the hosts
defenses.

In the MC horse the IPHA and JA test showed a low level of
antigen during the febrile responses. The presence of an antibody-
antigen complex would result in low titer and may also be re-
sponsible for the induction of a febrile response. The levels of
virus in the days following the febrile response indicated a rise
and leveling off.

The equilibrium state at this point is the most important con-
sideration in the disease for this represents a true carrier state. The
antigen level does not rise, therefore antibody or other neutralizers
are probably acting. The presence of this neutralizing substance
is not, however, able to affect a decrease in antigen, and thus a

FLYNN ET AL.: Viremia in Horses 313

steady equilibrium between virus and antibody is established. As
the results indicated, the virus in the MC horse showed a rise and
resulting high titer on the thirteenth day. This rise in titer may be
due to the intracellular replication of the virus being complete and
a large influx of virus being liberated. The virus immediately drops
on the day after reaching its highest level and the febrile response
occurs. This immediate drop in virus level would suggest rapid
neutralization of virus by the antibody.

The MCT, horse showed no initial infection with EIA; how-
ever, within eleven to fourteen days after inoculation with blood
from horse MC antigen was detectable in the peripheral blood.
The virus initially showed an increase with a concurrent tempera-
ture. The febrile response probably was due to the destruction of
cells in the host by the virus. The virus rose to highest levels after
the first febrile response. The virus then established a peculiar
stable state of equilibrium with its host. This stable state, however,
was disrupted by the liberation of large quantities of virus and
resulted in clinical manifestations as shown by a febrile episode.

The combination and comparison of the serologic and clinical
results obtained in these animals is merely suggested as a possible
in vivo course of the disease. The evidence obtained pertained to
only two studies of daily correlation of serologic results with clin-
ical signs and more information would be required before the ac-
tual in vivo course of the disease is adequately characterized.

The IPHA proved itself as sensitive, or even more sensitive, in
detecting virus than the IA test. The reactivity of the antibody
with sera from horses infected only with EIA indicated the detec-
tion of a highly specific antigen. The tests also were accurate in
the determination of those horses not infected with EIA. The re-
sults of this study verifies the reliability of the IPHA on both posi-
tive and negative sera and its sensitivity in daily testing of infected
horses.

The more distinct advantage of the IPHA over the IA is that the
IA only measures soluble antigens representing infection and repli-
cation of the virus within the host. The IPHA, on the other hand,
measures presumably both soluble viral antigen as well as the virus
particle. Thus, it is not surprising that in some instances one may
find one of these serologic tests (the IA or IPHA) more sensitive
than the other. It is interesting to note that levels of antigen vary

314 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

2-4 fold in practically all instances as depicted in Figure 4. In most
instances, the IPHA show the highest levels; however, in some in-
stances the IA shows the highest antigen levels. This might indi-
cate that the IPHA does not have the capability, as does the IA, of
detecting soluble viral antigen as well. Further investigation on
the comparison of these two tests is warranted.

ACKNOWLEDGMENTS

This study was supported by the Michael G. Phipps Foundation
of Palm Beach, Florida, and four equine race tracks of Dade and
Broward counties who form the major contributors to the Florida
Atlantic Equine Research Institute.

LITERATURE CITED

ALLEN, P. Z., S. SmrisNrHaA, AND J. H. VAUGHAN. 1965. Immunological studies

of equine antibodies to human y,-globulin. Jour. Immun., vol. 95, pp.
918-928.

BoyDEN, S. V. 1951. Absorption of proteins on erythrocytes with tannic acid
and subsequent hemagglutination by anti-protein sera. Jour. Exp. Med.,
vol. 93, p. 107.

BrapsHAw, L. 1966. Introduction to molecular biological techniques. Prentice-
Hall, Englewood Cliffs, New Jersey, pp. 114-119.

Cook, R. J. 1964. Reverse passive hemagglutination systems for the estimation
of tetanus toxins and antitoxins. Immun., vol. 8, p. 74.

Drecuss, M. N., anp L. S. Lomparp. 1954. Experimental studies in equine
infectious anemia. Univ. Pennsylvania Press, Philadelphia.

FAUCONNIER, B. 1966. Efficacité du lait sec écremé dans la prévention des
phénoménes d’autoagglutination non spécifique au cours de réactions
dhémagglutination passive utilisant les hématics formolées tannées.
Ann. Inst. Pasteur, vol. 110, pp. 453-457.

LEeFxowl71z, S. S., J. A. WitiiaMs, B. E. Howarp, AND M. M. SicEL. 1966.
Adenovirus antibody measured by passive hemagglutination test. Jour.
Bact., vol. 91, pp. 201-212.

Netson. R. A. 1953. The immune adherence phenomenon, an immunologically
specific reaction between microorganisms and erythrocytes leading to
enhanced phagocytosis. Science, vol. 118, pp. 733-735.

SAURINO, V. R., G. H. WaAppDELL, J. H. FLYNN, AND M. B. TEIGLAND. 1966.
Immunodiagnostic relations of three clinical types of equine infectious
anemia. Jour. Amer. Vet. Med. Assoc., vol. 149, pp. 1416-1422.

FLYNN ET AL.: Viremia in Horses oD

Sravirsky, A. B. 1954. Micromethods for study of protein and antibodies. I.
Procedure and general application of hemagglutination and hemagglu-
tination-inhibition reactions with tannic acid and protein tested red blood
cells. Jour. Immun., vol. 72, pp. 360-368.

Woop, W. A. 1964. Use of flow cells for scanning sucrose density gradients.
Gilford Instrument Laboratories, Report.

Florida Atlantic University, Boca Raton, Florida.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968 )

Temperature and Humidity Effects on Survival of Chickens

R. W. Dorminey, H. R. Wison, I. J. Ross, anp J. E. JONES

Witson and Armas (1963) found that with a relative hu-
midity of 75 per cent, the survival time of four-week old White
Leghorn chicks decreased sharply as the temperature was in-
creased from 40.5C to 48.8C. Lee et al. (1945) observed that
under natural conditions relative humidity was an important fac-
tor in the tolerance of laying hens to temperatures of 40.5C or
greater. However, very little is known concerning the exact rela-
tionships between temperature and relative humidity (R.H.) when
both were at high levels.

The following experiment was designed to determine the effects
of temperature, humidity, and air flow rate on the survival time
of five-week old White Leghorn chicks.

PROCEDURE

Two experiments were conducted with five-week old White
Leghorn chicks obtained from nine hatches. The chicks were
pedigree hatched from the F, generations of two genetic lines
which had been previously selected for a high and a low heat
tolerance (Wilson et al., 1966). Chicks were brooded in floor
pens with infra-red lamps as a heat source and were given water
and commercial type starter feed ad libitum.

The heat cabinet used was the same as that described by
Ross and Myers (1963) with the modifications listed by Wilson
et al. (1966). The temperature inside the chamber was controlled
within +0.2C, the R.H. within +2.5 per cent and the air flow
rate within +0.1416 cubic meters (m*) per minute. The chamber
temperature dropped approximately 1.94C while the chicks were
being placed in the chamber, but it recovered to approximately
0.4C of the desired temperature within three minutes after the
start of the test. The unit fully recovered within ten minutes
after the start of the test.

Temperatures of 40.5, 42.2, and 43.9C were used with 60,
75, and 90 per cent R.H. and air flow rates of 1.1327, 1.6990,
and 2.2653 m* per minute, resulting in 27 different environmental
treatments. Since only one heat cabinet was available it was im-

DoRMINEY ET AL.: Survival of Chickens Ot

possible to run 27 treatments per day. Therefore, nine different
hatches were made on different days to facilitate the use of chicks
when they were exactly five weeks of age. The same procedure
was used for the second experiment. A specially designed wire
cage with 30 individual compartments was used to keep the birds
separated and facilitate the maintenance of individual records.
Thirty chicks were equalized according to genetic line and sex in
each environmental treatment.

The survival time of individual chicks was measured in seconds
from time of placement in the chamber (Wilson et al., 1966)
and converted to minutes and tenths of minutes for analysis. The
data of both experiments were combined and subjected to analysis
of variance according to the procedure of Snedecor (1956).

RESULTS AND DISCUSSION

A significant difference (P<.01) was found in survival times
between the two genetic lines with the lo-line chicks surviving
an average of 72.4 minutes and the hi-lines an average of 99.9
minutes. The survival time of the males was 81.8 minutes and
the females 86.0 minutes. None of the interactions involving
genetic lines or sex were significant. Therefore, lines and sexes
were combined for this presentation.

Both temperature and humidity significantly (P<.01) affected
survival times (Table 1). Each time the temperature was in-
creased, survival time was decreased. Survival time was decreased
significantly when the R.H. was increased from 60 to 75 per cent,
and the decrease approached significance when the R.H. was in-
creased from 75 to 90 per cent. The effect on survival time of
increasing the R.H. from 60 to 75 per cent was similar to that
observed by Yeates et al. (1941) on body temperature when
the R.H. was increased from 65 to 75 per cent. Apparently,
temperature had a greater effect on survival time than humidity
within the ranges studied.

The temperature X humidity interaction was found to be signi-
ficant (P<.01) indicating the extent to which temperature affects
survival time is dependent upon the humidity and vice versa.
Survival time was shortened more when R.H. was increased from
60 to 75 per cent than from 75 to 90 per cent in both absolute
terms and on a percentage change basis (31.4 vs. 15.5 per cent

318 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SWAB TC Ere
Survival times of chicks as influenced by temperature,
relative humidity and air flow rates

Survival time (min.)+

REEL. Air Flow Temperature
(%) (m3 ) 40.5C AX2E 43.9C Average
60 IPS 27 204.92 102.4 AT.5 ITNSES
1.6990 199.1 100.4 49.3 116.3
2.2653 162.8 90.5 48.2 100.5
Av. 188.9 97.8 48.3 111.7
75 UB 27 LT) DED 43.7 (est
1.6990 129.9 64.1 39.0 Tile
22693 141.4 58.1 41.6 80.4
Av. 129.5 59.9 41.4 76.9»
90 S27; O72 44.4 32D 61.4
1.6990 109.9 44.4 33.9 62.7
PIGS TD) 52.0 31.9 65.0
Av. 109.4 46.9 32.8 63.0»
Av. IL AUS PA7/ 143.1 68.1 41.2 84.1
1.6990 146.3 69.6 40.7 85.5
2.2653 138.4 66.8 40.6 81.9

Av. 142.6* 68.2 40.87 83.9

1 Means with different superscripts differ significantly (P<.01).
2 Each value represents the average of 60 birds.

and 38.8 vs. 21.7 per cent, respectively) at both 40.5 and 42.2C.
However, at 43.9C this relationship was reversed (14.3 vs. 20.5
per cent). This demonstrates that survival time was affected more
by increasing the humidity at the lower temperatures than at the
higher temperatures and that increasing the R.H. from 60 to 75
per cent had a greater effect than increasing it from 75 to 90 per
cent, except at the highest temperature. This might have been
expected since Wilson and Edwards (1953) reported that the
effect of 72 per cent R.H. on respiratory rate and body tempera-
ture was more pronounced at 37.8C than at lower temperatures.

The survival time decreased more when the temperature was
increased from 40.5 to 42.2C than when it was increased from 42.2
to 43.9C in absolute terms at all humidities. However, the per-
centage change was approximately equal at 60 per cent R.H. (48.2

DorMINEY ET AL.: Survival of Chickens 319

vs. 50.6 per cent) and was much higher in the 40.5 to 42.2C change
than in the 42.2 to 43.9C change at 75 per cent R.H. (53.7 vs. 30.9
per cent) and at 90 per cent R.H. (57.1 vs. 30.1 per cent). There-
fore, at the higher temperature and lower humidity, the effect of
increasing the humidity was masked to some extent by the magni-
tude of the temperature effect. This may indicate that as tempera-
ture increased the critical effect of R.H. was reached at a lower
point.

Air flow rates did not significantly affect the survival time of
the chicks. This might have been expected since the major source
of heat loss in the chick is by evaporative cooling from the lungs
and in this study it was possible to maintain a constant relative
humidity at all air flow rates. However, with decreased air flow
rates this factor might become more important.

ACKNOWLEDGMENT

This work was supported in part by the Center of Tropical
Agriculture, Institute of Food and Agricultural Sciences, University
of Florida.

SUMMARY

Five-week old chicks from high and low heat-tolerant lines
were subjected to temperatures of 40.5, 42.2, and 43.9C with 60,
75, and 90 per cent relative humidity and air flow rates of 1.1327,
1.6990, and 2.2653 cubic meters per minute. Survival times were
142.6, 68.2, and 40.8 minutes, respectively, for temperature and
111.7, 76.9, and 63.0 minutes, respectively, for relative humidity.
Genetic line, temperature, and humidity were found to have signi-
ficant effects on survival times of the chicks while sex and air re-
placement rate did not significantly affect survival times. The
temperature < humidity interaction was significant.

LITERATURE CITED

LEE, D. H. K., K. W. Rosinson, N. T. M. YEATES, AND M. I. R. Scott.
1945. Poultry husbandry in hot climates—Experimental enquires.
Poultry Sci., vol. 24, pp. 195-207.

Ross, I. J., anp J. M. Myers. 1963. Humidity, temperature and air flow
control cabinets for experimentation in processing agricultural products.
Paper presented at the 1963 International Symposium on Humidity
and Moisture, Washington, D. C., May 20-23, 1963.

320 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SNEDECOR, G. W. 1965. Statistical Methods. 5th Ed. The Iowa College
Press, Ames, Iowa.

Witson, H. R., A. E. Armas, I. J. Ross, R. W. Dormingey, anp C. J.
Witcox. 1966. Familial differences of S$. C. White Leghorn Chickens
in tolerance to high ambient temperature. Poultry Sci., vol. 45, pp.
784-788.

Witson, H. R., anp A. E. Armas. 1963. The effect of temperature and
age on susceptibility of chicks to heat. Unpublished data. Fla.
Aor. Eixpreota.

Witson, W. O., anp W. H. Epwarps. 1953. Interaction of humidity and
temperatures as affecting “comfort” of White Leghorn hens. Poultry
Scit, vol. 382, p. 929)

Yeates, N. T. M., D. H. K. LEE, anp H. J. G. Hines. 1941. Reactions of
domestic fowl to hot atmospheres. Proc. Roy. Soc. Queenslands,
vol. 53, pp. 105-128.

Department of Poultry Science, University of Florida, Gaines-
ville, Florida 32601. Florida Agr. Exp. Sta. J. Series No. 2458.

Quart. Jour. Florida Acad. Sci. 30(4) 1967 (1968)

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1967

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Institute of Technology
Florida Presbyterian College
Florida Southern College

Florida State University
Jacksonville University
Marymount College

Miami-Dade Junior College
Mound Park Hospital Foundation
Nova University of Advanced Technology
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida

University of Tampa

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1967

President: JACKSON P. SICKELS
Department of Physical Sciences, University of Miami
Coral Gables, Florida

President Elect: CLARENCE C CLARK
Department of Physical Sciences, University of South Florida
Tampa, Florida

Secretary: Joun D. McCrone
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: JAMES B, FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: Pierce BRODKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges
should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

x

Quarterly Journal
of the

Florida Academy of Sciences

Vol. 30 March 1967 No. l
CONTENTS

Interferometry of Jupiter at 18 Mc/s
Jorge May and Thomas D. Carr

Effect of Hurricane Betsy on the southeastern Everglades
Taylor R. Alexander

Fishes of the St. Johns River, Florida
Marlin E. Tagatz

Composition and feed value of shrimp meal W. G. Kirk,
R. L. Shirley, J. F. Easley, and F. M. Peacock

Animal remains from a midden at Fort Walton Beach Elizabeth S. Wing
Fossil vertebrates from Navassa Island, W.I. Thomas H. Patton

Notes on the trematode genera Cleidodiscus and Urocleidus

C. E. Price
Aerial respiration in the Florida Spotted gar Brian McCormack
Invertebrates found in water hyacinth mats James O’Hara

Mailed April 16, 1968

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Mareins should be 114 inches all around.

Tires should not exceed 55 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and
ADDRESS in paragraph form following Literature Cited.

LITERATURE CiTED follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLEs are charged to authors at $19.95 per page or fraction. Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

Lecenps for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($17.30 per page; $15.80 per half
page). Drawincs should be in India ink, on good board or drafting paper, and
lettered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. Do not submit illustrations needing reduction by more than one-half.
PHotocraPus should be of good contrast, on glossy paper. Do not write
heavily on the backs of photographs. 3

Proor must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

Vol. 30 March, 1967 No. 1

Interferometry of Jupiter at 18 Mc/s
Jorce May Aanp THomas D. Carr

Ir is believed that the radiation emitted at decameter wave-
lengths from the planet Jupiter originates in areas which are
considerably smaller than the visible disc of the planet. Radio
interferometers must be employed in the measurement of the
angular sizes of such small sources. The Michaelson interferome-
ter, which is widely used in both radio and optical astronomy,
presents severe technical problems in the case of the Jovian radio
source. It is extremely difficult to maintain phase coherence be-
tween the signals arriving at the mixing point from the ends of the
baseline of a Michaelson interferometer when the baseline is very
long, and baselines of great length may be required to resolve
the radio sources on Jupiter. For this reason a type of interferom-
eter which does not require phase coherence in signal transmission
is being used at the University of Florida in an attempt to deter-
mine the sizes of the Jovian sources. This work describes the
system employed and presents some preliminary results which
have been obtained with it.

THE MICHAELSON INTERFEROMETER

The angular separation between the fringes of the Michaelson
interferometer, whether it be of the optical or of the radio variety,
is approximately equal in radians to the ratio of the wavelength
(X) to the baseline length (d). For sources of small angular
diameter compared with the fringe separation, the intensity at the
fringe minima is essentially zero. However, for progressively
larger sources or more closely spaced fringes, the fringes tend to
become more and more smoothed out. It is possible to deduce
the equivalent one-dimensional angular distribution of brightness

2, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

across a small source from the manner in which the relative fringe
depth decreases due to this smoothing process as the baseline is
made longer. The limit of resolution which can be obtained in
this way is of the order of \/d radians. Thus by using inter-
ferometers having baselines capable of being extended to great
lengths, it is possible to resolve much smaller sources than could
be resolved with a single lens, mirror, or antenna.

Interferometers have been widely used at the shorter radio
wavelengths to measure the angular sizes, and in some cases the
apparent shapes, of cosmic radio emitters subtending extremely
small angles. Such interferometers generally consist of a fixed
antenna and a movable one, the signals received by the two
antennas being conveyed to the receiver through transmission lines
or radio links of essentially equal electrical lengths. Intensity
maxima and minima (i.e., radio “fringes”) are produced as the
source moves relative to the lobe pattern of the interferometer.
In some cases the rotation of the earth is sufficient to produce the
desired relative motion; in others the lobe pattern is rotated by the
artifice of introducing a uniformly changing time delay into one of
the transmission lines.

Interferometric resolution of small-angle sources at the longer
radio wavelengths is much more difficult because of the extremely
long baselines which are required. The principal difficulty lies
in the transmission of the radio frequency signals from the antennas
at the ends of the baseline to the receiver at an intermediate
position. Transmission lines of lengths greater than a few miles,
and the associated repeater amplifiers which would be required
to overcome attenuation effects, are prohibitively expensive. Radio
relay links are practicable only up to line-of-sight distances, unless
expensive repeater stations are employed. Sky-wave propagation
cannot be used because the instability of the ionosphere prevents
the maintenance of phase coherence.

THe BrRrown-l[Iwiss INTERFEROMETER

To overcome these difficulties, Brown and Twiss (1954) de-
veloped an interferometer of a different type. In the postdetection
correlation interferometer, as it is called, there is a receiver at each
antenna, and the audio signals from the receiver outputs are

May AND Carr: Interferometry of Jupiter 3
transmitted to a common location where they are correlated. No

fringes are produced. The angular dimensions of the source in
this case are deduced from measurements of the degree of corela-

VY

RECEIVER RECEIVER

LOW FREQUENCY
BAND PASS
FILTER

LOW FREQUENCY
BAND PASS
FILTER

DELAY
NETWORK

TRANSMISSION
LINK

CORRELATOR
LINEAR LINEAR
DETECTOR DETECTOR
RECORDER RECORDER RECORDER

Fig. 1. Block diagram of the post-detector correlation interferometer
developed by Brown and Twiss.

4 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

tion between pairs of audio signals obtained with several baseline

lengths.

A simplified diagram of such an interferometer is shown in
Fig. 1. The audio output of the receiver at each end of the base-
line is rectified and is recorded on a pen recorder. The two audio
signals are also fed into the correlator, located at one of the sta-
tions, where they are multiplied and their averaged product is
recorded on a third pen recorder. A radio transmission link is
used. The great advantage of this system is that since information
is conveyed over the transmission link in the form of an audio
modulation rather than as a phase-coherent radio frequency signal,
transmission requirements are much less stringent. Since sky-wave
propagation could be employed for transmission, baselines of hun-
dreds or thousands of miles become possible. The purpose of the
delay network is to compensate for signal delay incurred in trans-
mission from the remote station, and also for the difference in
arrival time of the incident radiation at the two antennas when the
direction of the source is not normal to the baseline. The deflec-
tions of the three recorders are used to determine the normalized
correlation coefficient of the two audio signals. This quantity is
measured for each of a series of baseline lengths. It can be shown
that the normalized correlation coefficient for the rectified audio
outputs of the receivers is proportional to the square of the
modulus of the Fourier transform of the brightness distribution
for a given value of d/d, provided the source is assumed to be
radially symmetrical. Thus the angular distribution of brightness
across the source can be deduced from a series of measurements
of the normalized correlation coefficient for different baseline
lengths. The principal disadvantage of the post-detector corre-
lation interferometer is that its accuracy is greatly reduced by
background noise unless the source is relatively strong.

DECAMETER-WAVELENGTH RADIATION FROM JUPITER

The planet Jupiter sporadically emits powerful bursts of radio
noise at frequencies below 40 Mc/s. This decametric radiation,
as it is called, has been studied with simple broad-beam radio
telescopes at the University of Florida for a number of years
(Smith and Carr, 1964). Such instruments provide information
on the times of occurrence of the radiation and its intensity, but

May AND Carr: Interferometry of Jupiter 3)

cannot be used to measure the size of the radiating source. Early
experiments by Gardner and Shain (1958) indicated that rapid
fluctuations in the envelope of the Jovian noise received at two
locations a few kilometers apart were often largely uncorrelated.
These scintillations were attributed to the effect of drifting iono-
spheric irregularities, The relatively extreme degree of the scin-
tillation was interpreted as proof that the source is of very small
angular diameter in comparison with other known radio sources.
It was suspected by later workers that despite the presence of
strong scintillations which were often uncorrelated over distances
of a few kilometers, the ionospheric irregularities might produce
very little degradation in the correlation of radio frequency com-
ponents. Proceeding on the assumption that this is true, programs
were initiated independently at the C.S.I.R.O. Radiophysics Labor-
atory in Sydney, Australia and at the University of Florida Radio
Observatory to measure the size of the Jovian radio source inter-
ferometrically. Results were obtained first by the Australian group
(Slee and Higgins, 1966). The mountainous character of the
terrain near Sydney made possible the use of the Michaelson
(phase coherent) radio interferometer with very long baselines,
since the signal from the remote antenna could be relayed to the
mixing point by a line-of-sight radio link. Slee and Higgins, oper-
ating at 19.7 Mc/s, found that with a baseline of 85.5 km a stable
fringe pattern was observed despite the fact that the burst enve-
lope amplitudes obtained at the two stations were scintillating
more or less independently of one another. It was concluded,
as had been suspected, that the ionospheric irregularities affect
the correlation of the radio frequency components from the two
antennas relatively little. Slee and Higgins extended the baseline
to a maximum of 200 km. From the observed decrease in relative
fringe height with increase in baseline, they concluded from a
limited number of observations that the apparent source width
was approximately 15 sec of arc.

ReEsuLtTs WITH THE MopiFIeED BRown-Twiss INTERFEROMETER

The interferometer employed by the University of Florida is a
modification of the Brown-Twiss interferometer. A frequency of
18 Mc/s was selected, since it represents a compromise between
decreased Jovian activity at higher frequencies and increased

6 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

difficulty with ionospheric effects at lower frequencies. The re-
ceiving stations were at Gainesville and Ocala, Florida, defining a
baseline which is 55.3 km in length and is oriented in an approxi-
mately north-south direction.

A block diagram of the modified post-detector correlation inter-
ferometer is shown in Fig. 2. An antenna, a receiver, and a

RECEIVER

RECEIVER

LOW PASS BAND
TIMING SYSTEM FILTER

AMPLIFIER
TELEPHONE
LINE
MAGNETIC TAPE
RECORDER
CONTROL

SYSTEM

LOW PASS BAND
FILTER

AMPLIFIER

MAGNETIC TAPE
RECORDER

CONTROL
SYSTEM

Fig. 2. Block diagram of the modified post-detector correlation interfero-
meter used at the University of Florida.

dual-track tape recorder are located at each station. The filtered
audio output of the receiver is recorded on one track of the tape
recorder, and a timing signal on the other track. The same timing
signal is used at both stations; it is produced at the base station
and is transmitted to the other via telephone line. The Jovian
signal itself could not be transmitted by telephone line because of
excessive phase and amplitude distortion. After recordings have
been made, the tape from the remote station is brought to the
base station, where the signals on the two tapes are compared.
Tape recordings were made only during periods of intense emis-
sion from Jupiter. A single observer controlled the operation of
both stations.

On play-back, the recorded Jupiter noise from one station was
fed to a cathode ray oscilloscope, and the associated timing signal
was made to flash a small neon lamp. The cathode ray spot and
the flashing lamp were photographed simultaneously with a cam-
era in which the film travelled with continuous motion at 60 inches

May AND Carr: Interferometry of Jupiter 7

per second. The marks produced on the film by the lamp flashes
permitted the time of any part of the Jupiter noise waveform to
be read with an accuracy of about 0.2 millisecond. Tapes which
had been recorded at both stations were played back individually
through the same transcription system, eliminating in this way
any possible difference introduced by the equipment. Correspond-
ing sections of the films from the two stations were identified
with the aid of the timing signals, and pairs of sections containing
simultaneously-recorded signals were aligned side-by-side for mea-
surement of the degree of correlation and for other studies. A
sample of a pair of these films is shown in Fig. 3.

\

AUATAT CR yf po,

AW

Fig. 3. Audio-frequency noise from Jupiter obtained with receivers 55.3
km apart during the 1964-65 apparition. Time marks are milliseconds.

The degree of correlation between a pair of rf signals entering
separate receivers is closely related to that of the audio signals at
their outputs. The necessary condition for good rf correlation, and
hence for good audio correlation, is that the source diameter in
radians be small compared with X/d. From Fig. 3 it is evident
that there was good pulse-for-pulse correlation when the receivers

8 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

were spaced at 55.3 km (3324). We can therefore set an upper
limit of about one minute of arc (the apparent angular size of
the visible disc of the planet is close to this figure) for the size
of the source of decametric radiation from Jupiter. This is con-
sistent with the results of Slee and Higgins.

Longer baselines will be required to resolve the source. The
correlation coefficient obtained with the 55.3 km baseline was
obviously close to unity. As the baselines are increased, the corre-
lation coefficient will of course become less and will become more
difficult to estimate. A calibration system is being developed to
aid in making quantitative determinations of the correlation co-
efficient. Pairs of noise signals having various known values of
correlation coefficient will be produced by the calibrator and will
be recorded. A curve can thus be obtained of the average number
of pulse coincidences per unit time interval for the two noise
signals as a function of their correlation coefficient, Then the
pulse coincidence rate for the pair of Jupiter signals can be mea-
sured from the photograph, and with the aid of the calibration
curve the correlation coefficient can finally be determined.

Two new stations providing longer baselines are planned. One
is at St. Petersburg, Florida, at a distance of 193 km from the
base station in Gainesville, and the other is at Maipu, Chile, which
is 7040 km distant. With the latter baseline it is theoretically
possible to achieve a resolution of 0.5 second of arc. Timing sig-
nals from radio station WWYV, corrected for propagation effects,
will be used in place of those sent by telephone line. It is expected
that these increased baselines will permit the source to be re-
solved and angular estimates of its size to be made on a routine
basis.

ACKNOWLEDGMENTS

We wish to thank Mr. C. N. Olsson, Mr. G. F. Walls, and
Miss Dolores Smoleny for valuable assistance in the development
and operation of the equipment and in the reduction of data.
We gratefully acknowledge the financial support of the National
Science Foundation, the Office of Naval Research, and the Army
Research Office (Durham).

May ANpD Carr: Interferometry of Jupiter 9
LITERATURE CITED

Brown, R. H., Anp R. Q. Twiss. 1954. A new type of interferometer for
use in radio astronomy. Philosophical Mag., Serial VII, vol. 45, pp.
663-682.

GaRpDINER, F. F., AND C. A. SHatn. 1958. Further observations of radio
emission from the planet Jupiter. Australian Jour. Physics, vol. 11,

pp. 55-69.

SLEE, O. B., AND C. S. Hiccrns. 1966. The apparent sizes of the Jovian

decametric radio sources. Australian Jour. Physics, vol. 19, pp.
167-180.

SmirH, A. G., AND T. D. Carr. 1964. Radio exploration of the planetary
system. D. Van Nostrand Co., Princeton, New Jersey, 148 pp.

Department of Physics and Astronomy, University of Florida,
Gainesville, Florida 32601; Observatorio Radioastronomico de
Maipu, Universidad de Chile, Maipu, Chile.

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Effect of Hurricane Betsy on the Southeastern Everglades

TAYLOR R. ALEXANDER

On September 7-8, 1965 Hurricane Betsy affected the lower
southeast coast of Florida with winds of 120 miles per hour and
tides 6 feet above normal. Extensive flooding by salt water blown
in from the east occurred. Damage in the form of total kill to
much of the native flora was spectacular in parts of the area
indicated in Fig. 1. The affected area is a part of the region
described and designated by Egler (1952) as the Southeastern
Saline Everglades. This extensive mainland region lies east, south-
east, and south of the Miami Oolite rock rim and extends to salt
water. Severe damage was confined to parts of the areas desig-
nated by Egler as Belts 3 and 4. They are generally classified
as coastal glades and mangrove physiographic divisions and extend
inland as much as 10 to 12 miles from salt water. A part of the
Everglades National Park lies within the affected area.

The entire region is characterized as a very flat sawgrass
prairie, dotted at frequent intervals by tree islands (low hammocks )
and a mangrove fringe belt at salt water. The soils are Perrine
or Flamingo marl under most of the sawgrass, and peat under
the tree islands. Mangroves occur on both marl and peat. The
region encompasses fluctuating boundaries of fresh to brackish to
salt water. Hence, from the inland edge of the Southeast Saline
Everglades there is a gradual but steady change in vegetation
toward completely salt tolerant species. Natural drainage is to-
ward the southeast.

Anthropic activity in the area has had important influences
on the direction and amount of water flow and associated with
these have been inevitable vegetational changes. Roads on ele-
vated and inadequately culverted roadbeds and canals with spoil
banks act as channels, dams, and barriers for both fresh and salt
water movement. Some of these have been in place since 1905.
The damage patterns from Betsy and other recent storms, in part,
are definitely related to man-made structures. In this instance,
the roadbed of U. S. highway 1 south from Florida City and a new
canal known as C 111 and its spoil banks are especially significant
(Fig. 1). This canal was begun in 1964 and completed in 1966
(Klein, 1965).

ALEXANDER: Effect of Hurricane Betsy Ta

lon

| Everglades National Park Uy

Heed

Miles

= @ = Canals
e eee Severe Damage Line |

Fig. 1. Map of southeastern Florida, showing location of severe damage
area east of dotted line.

Canal 111 is located just north of the northeast boundary of the
Everglades National Park and was constructed without a salinity
control structure near its mouth that would protect against high

12 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

and storm tides, and salt water intrusion. For these reasons it
could be assumed that ecological changes may occur in the near-
by Park areas. Since a pattern of damage did develop involving
C 111, the data reported herein is especially significant to this
type of ecological problem and can serve as a reference for
further studies in this location.

The vicinity of the intersection of U. S. 1 and C 111 has been
the site of continuing field studies by the author since 1947. This
long-term low hammock study is still in progress and was initially
reported by Alexander (1955). Some of the long-term plot sites
were bisected and some destroyed when C 111 was dug. This
allowed a series of reference and continuing measurements to
determine the differences, if any, that might develop north and
south of the canal. Early in 1965 a study was started on the re-
lationship of blue green algae to marl formation in the area. A
number of soil, water, and algae collecting and measuring sites
had been visited monthly before the hurricane. Data from these
afforded an opportunity to compare the “before and after” situa-
tion that developed.

This report is an account of the observations on plant kill,
survival, natural post-storm seeding, and of certain edaphic condi-
tions east of the damage line shown in Fig. 1. Most herbaceous
plants were badly broken, defoliated, and in many instances rotted
so quickly that their remains were impossible to identify. Hence,
much of the emphasis on plant damage relates to woody plants.
Measurements of chloride content, electrical conductivity as a
measurement of total salts concentration, salinity, and pH of sur-
face water and soil are included. The period reported includes
the sampling date, August 13, before the storm and the following
six months from September 1965 to March 1966.

MrtTHOpDS

Plant lists were available from previous plot and transect
studies (Alexander, 1955; Egler, 1952) and were used as a basis
for determining the plant population. The prairie was almost a
pure stand of sawgrass in most areas. Several hammocks in
different locations were checked every month to see if defoliated
and broken plants were sprouting. Recovery in the form of root
sprouts and seed germination during the first six months after the

ALEXANDER: Effect of Hurricane Betsy 13

storm was so limited that plot and transect methods were not
practical. Instead, lists were made as hammocks were searched
each month for signs of survival or recovery. Plant names used
are from the checklist of Lakela and Craighead (1965).

The sawgrass prairie about one quarter mile west of U.S. 1
and both north and south of C 111 was analyzed in February
after the storm using 120 randomly located meter square plots to
determine frequency and density of sawgrass. Dead and living
culms were counted so the data could be used to determine sur-
vival.

Surface water samples were collected in 500 ml bottles by
carefully moving an open bottle from top to bottom so as to
equally sample the entire water layer. Three sites were used for
soil collections: (1) prairie marl under sawgrass; (2) near-by
peat at the margins of low hammocks; and (3) peat near the cent-
ers of the same hammocks. Soils were collected from three posi-
tions at each site and a composite sample of 500 cc made for
analysis. Analyses for chlorides, electrical conductivity, and pH
were made following procedures used by the Dade County ( Flor-
ida) soils laboratory (Llewellyn, 1963). Electrical conductivity,
reported as millimhos per centimeter at 25 C, was measured with
a “Solubridge” RD 15 instrument (Industrial Instruments). A
Beckman model 76 pH meter was used in the laboratory and a
battery operated pocket pH meter (Analytical Measurements Inc. )
in the field. Salinity was measured with a set of three hydro-
meters (Sea Water Test Set-G. M. Mfg. & Instrument Corporation ).
Chloride determination was by silver nitrate titration.

VEGETATIONAL DAMAGE

The area of severe plant kill is shown in Fig. 1. Two weeks
after the storm the area looked scorched as far as one could see,
except that a few very salt tolerant species were still green or
leafing out (scored minor or no injury in Tables 1, 2, and 3).

It was obvious that many of the plants in the low hammocks as
well as the sawgrass were severely injured or dead. In the area
to the west of U. S. 1 the plants in at least 350 low hammocks
looked dead. Plants in the triangular area northwest of the inter-
section of U. S. 1 and C 111 were hurt the worst especially for a
distance about three miles from the point of intersection. The kill

14 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

was severe just west of U. S. 1 and generally decreased in severity
toward the west.

Another damaged area, not shown in Fig. 1, was within the
Park and located just north of West, Cuthbert, and Seven Palm
Lakes. This was observed and reported by Robertson (personal
communication from William B. Robertson, Jr., Research Biologist,
Everglades National Park, Homestead, Florida, 1965). Observa-
tion was done by low level aircraft reconnaissance. This area
centered about 20 miles west southwest of the intersection of
U.S. f and C 111) Plant kill occurred in“anareayabouteasix
miles long, east to west, and about one-half mile wide at its
widest. Within this area the degree of damage was spotty.
Effects were similar to those seen to the northeast. Dense and
tall sawgrass was killed and Paurotis wrightii growing in this area
was severely damaged. Robertson presumed that the natural
levee north of the lakes (Craighead, 1964) trapped saltwater as
the Geri bankacire:

In the area east of U. S. 1 the damage was not as extensive and
uniform as it was to the west of this highway. One reason is the
fact that proportionally a greater part of the vegetation is salt-
tolerant mangroves. However, species in low hammocks through-
out the area were affected severely, especially near the coast.
Another reason is that considerable inland acreage has been or is
being farmed and the original vegetation is either missing or
greatly disturbed. Furthermore, the exotic Casuarina equisetifolia
has become well established and dominates sizeable areas. This
species tolerated the change in salt concentration and survived
with little damage.

The hammock trees were wind-damaged considerably but not
to the extent to cause death. When leafing-out failed to occur and
cambial zones turned brown, it became obvious that most of the
defoliated plants were dead. Tables 1-3 show the effect on each
species. Included species are from the entire area east of the sev-
ere damage line in Fig. 1. Where a species is checked in the first
two columns, it usually indicates a high percentage of kill but
that survivors were found. Those that finally root sprouted were
included under the column, severe injury.

Large and high windrows (up to two feet high) of trash and
plant remains were deposited on the eastern edge of each ham-

ALEXANDER: Effect of Hurricane Betsy

TABLE 1.

Effect on woody plants in area of greatest damage

Complete Severe Minor or New
Species kill Injury No Injury Seedlings

Annona glabra — x — x
Baccharis spp. x ax — Xi
Berchemea scandens 5X

Borrichia spp. — — Ke —
Bumelia celastrina — — x

Calyptranthes pallens —
Casuarina equisetifolia —
Cephalanthus occidentalis —

x
x
me

Chiococca alba — x x —
Chrysobalanus icaco x x
x

Coccoloba uvifera —

x
x
Conocarpus erecta — — 25. Og
Eugenia spp. — X
Ficus aurea — x
Hippocratea volubilis —
Ilex cassine x
Magnolia virginiana x — — —
Metopium toxiferum Xe x
Morinda roic — x
Myrica cerifera x x ~— —
Persea borbonia x

Pithecolobium
gaudelupense —

x
Randia aculeata — x — —
Rapanea guianensis x x x
Rhizophora mangle — — x x

Salix caroliniana x — -—
Schinus terebinthifolius — — x

wm

Swietenia Mahagoni x
Taxodium distichum x

wm mM OM
ws
|

Toxicodendron radicans —
Trema micrantha

va

|

|
wm A

Vitis munsoniana X — —

16 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 2.

Effect on herbaceous dicotyledonous plants in area of greatest damage

Species

Aster exilis
Centella erecta
Chamaesyce scoparia

Eupatorium leptophyllum
Heliotropium parviflorum

Ipomoea cathartica
Ipomoea sagittata
Mikania_ batatifolia

Complete

kill

Parthenocissus quinquefolia x

Pass‘flora pallida
Pluchea spp.

Samolus ebracteatus
Solanum verbascifolium

Severe

Injury

TABLE 3.

Minor or
No Injury

New
Seedlings

Paes See eee ie

Effect on monocotyledonous plants in area of greatest damage

Severe

Minor or

New

Complete

Species

Kill

Injury

Seedlings

No Injury

Crinum americanum
Distchol's spicata
Eleocharis cellulosa
Epidendrum tampense
Juncus roemerianus
Mariscus jamaicensis
Muhlenbergia capillaris
Paspalum vaginatum
Pe'tandra_ virginica
Rhynchospora tracyi
Sabal palmetto
Serenoa repens

Smilax spp.
Sporobolus virginicus
Tillandsia spp.
Typha domingensis
Vanilla _ dilloniana

ayeu lueee ene | eee oe

ALEXANDER: Effect of Hurricane Betsy Wf

mock. The usual loose and rather thick mat of organic debris
and duff on the soil was floated away, becoming a part of the
windrow deposit for the next hammock to the west. Very little
new sediment from the Bay was deposited anywhere. Hence, the
soil was cleaned of its organic litter and exposed directly to the
sun since the canopy was also gone. In some hammocks numerous
windthrows occurred. In others most trees were left broken, but
upright and leafless. Roots and the organic soil layer were severely
disturbed and loosened in every hammock.

The sawgrass areas looked dead above the water but leaves
were green below the surface. However, most of these turned
brown as the water level dropped. Only on the elevated edges of
the hammocks were there areas of relatively undamaged sawgrass.

Epiphytes, orchids, and bromeliads were damaged according
to position above ground level. Those that were high enough
to escape being covered by salt water survived with little damage.
In the case of vanilla orchid vines, the parts of vines up in the
tree were unhurt, while the parts of the same vines that were
near the ground and were salt affected died. Hence, such species
were checked in the first three columns of Table 3.

The algae mat that normally covers the surface of the marl
prairie had disintegrated within two weeks, particularly north of
C 111 and near the coast. A fine flocculent sediment apparently
derived from the disintegrating mats covered the marl to a depth
of an inch and a half. As this settled an unusual reddish tan color
developed on top. Where the mat remained partially intact a
black color developed through most of its profile. When stirred
the bottom materials smelled strongly of hydrogen sulfide.

VEGETATIONAL SURVIVAL

The more southernly located hammocks looked comparatively
green after the storm owing to the abundance of the red man-
grove, Rhizophora mangle, that was hardly damaged and Cono-
carpus erecta that locally suffered some defoliation but leafed out
quickly. The latter tends to replace the Rhizophora in hammocks
away from the coast. These two trees afforded most of the green
seen in the hammocks for the first six months. Nearer the coast,
species typical of salt water areas were not damaged and were
not tabulated.

18 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The frequency of sawgrass on marl from the meter square plot
study was 100 per cent with a culm density of 32. In the area of
U. S. 1 and C 111, the survival north of the canal was only 4 per
cent while south of the canal it was 23 per cent. Survival of
sawgrass increased away from the canal and coast. Eleocharis
cellulosa survived the storm and spread rapidly in the areas of
sawgrass kill. Especially to the north of C 111 this sedge was
the only green plant above the water between the hammocks.

Other plants that survived without extensive damage were
Borrichia arborescens, B. frutescens, Bumelia celastrina, Casuarina
equistifolia, Sabal palmetto, Serenoa repens, and Solanum verbas-
cifolium.

The first seedlings of any great number, other than red man-
grove, were seen on December 12 three months after the storm.
Among these were sawgrass and poison ivy. Six months after the
storm seedlings of Aster exilis, Baccharis halimifolia, Mariscus
jamaicensis, Mikania batatifolia, Parthenocissus quinquefolia, Per-
sea borbonia, Pluchea foetida, P. petiolata, Smilax, and Vitis mun-
soniana were rather common, but not abundant. Gametophytes of
a moss and two ferns, Acrostichum spp. and Blechnum serrulatum,
were noted in several hammocks. The fern sporophyte plants
had been injured but had recovered quickly. All of these young
plants were in hammocks on soil that was above water. During
the first six months after the storm the marl prairie was continu-
ously under water and no seedlings of any species were seen.
However, an alga, Chara sp., developed in water over the marl
areas and became abundant especially north of C 111.

ENVIRONMENTAL CONDITIONS

Heavy rainfall in southern Florida for several months preceed-
ing Betsy had raised the water table above the soil in the marl
prairie. Only the elevated interior of the low hammocks was free
of standing water when the storm struck. The August 13 hygro-
meter datum on the water north of C 111 (Fig. 2) indicates fresh
water. This instrument was not accurate at very low salinities.
Conductivity measurements of the surface water on August 13
were 5.9 north of C 111 and 8.3 to the south. These data reflect
the fact that the area was under salt influence. (For comparison,
surface water from these same localities at the end of a prolonged

ALEXANDER: Effect of Hurricane Betsy 19

Surface Water Salinities

30 AN AUG-MAR.

onion en N of Canal 111
20

uum) Of Canal ILI

8-13 9-26 | I=6 [2-12 I-8 212 3-3

Fig. 2. Graph showing differences in water salinity in parts per thousand
over marl north and south of Canal 111.

rainy season has been found to be as low as 0.85 and 1.4, re-
spectively. )

Salt water pushed inland during the storm overflowed the tops
of roadbeds and in some cases became impounded since it could
not follow the natural drainage back to the bay. Salt water also
flowed up the canals and flooded out on the land. This gave rise
to a variety of soil conditions. This impoundment was most
severe west of U. S. 1 and north of C 111. Other lesser areas of
impoundment occurred along canal banks and roads elsewhere.
The level of salt water at the peak of the storm tide over the
marl soil near the coast was about four feet. Two weeks after
the storm, water eighteen inches deep covered the marl north of
C 111 and west of U. S. 1 and only the very highest parts of the
hammocks were above water.

The extent of surface water contamination by salt water ex-
pressed as salinity (sea water at 35 parts per 1000) is presented
in Fig. 2. Also the rapid rate of recovery toward fresh water
condition is apparent. Similar data were obtained throughout the
area of extensive kill.

The salt content of the soils of the area expressed as ppm
chlorides is shown in Table 4. Electrical conductivity measure-
ments correlated very well with all the other soil measurements
taken. For example, on the August 13 collection date, mar] soil
of the high-damage area was 2.3 millimhos per centimeter. On
September 26 the marl south of C 111 was 6.8 and north of C 111

20 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 4.

Soil chlorides (ppm) in area of greatest kill

Gollection Hammock peat Prairie marl Rainfall
Date Margin Center ING Ge bia Sri at Site*
ANH MB, (GS) 8,650 5,850 2,900 3,700 —
Sept. 7 & 8 Hurricane Betsy 8.00**
Sept. 26 10,450 — 19,300 ~ 10,850 —
Nov. 6 1222.0 1,100 8,000 10,050 9.50”
Dec. 12 9,250 3,000 4,250 2,600 Wh 8)5)
janwoa 66 5,850 850 3,100 1,850 1.20”
Feb. 12 5,100 600 2,850 DMO) 2.40”
Mar. 3 4,500 500 2,600 2,100 1.00”

* Since last date
** Fi stimated

it was 13.0. The soil of hammock margins also regularly tested
much higher in conductivity than samples taken from hammock in-
teriors. Except for the September 26 measurement, hammock mar-
gin soils were higher in conductivity than nearby marl.

Soil pH remained fairly constant and was usually above 7.0
for all samples. Marl samples were 8.0 or slightly higher; ham-
mock interiors were slightly less alkaline and hammock margins
were occasionally as low as 7.0.

DISCUSSION

The data from several measurements of different soil factors
supports the conclusion that the death of plants in this storm was
primarily due to the sudden overland influx of salty water (salinty
of 30) that penetrated the root zone raising soil chlorides to as
high as 19,000 ppm. Apparently, there was not sufficient time for
root cells to adjust osmotically, even in many species considered
salt tolerant. Furthermore, the entire flora had been exposed dur-
ing previous months to a freshening water cycle. For comparison,
a level of 1000 ppm chloride in soil is critical for the most tolerant
crops and is considered a problem. Soils that have an electrical
conductivity of over 4 millimhos per cm are unsuitable for many
crop plants. One mmho/cm is equivalent, on the average, to 640
ppm of salt (Bernstein, 1964). After the storm, electrical con-

ALEXANDER: Effect of Hurricane Betsy 21

ductivity measurements ranged above 6 mmhos/cm. These data
also support the salt kill concept.

Samples of surface water were not collected between Septem-
ber 26 and November 6 and it is known that water levels dropped
drastically before heavy rains occurred at the end of October.
These restored water levels to the levels that prevailed just after
the storm. Evaporation rates measured in comparable areas are
known to be about 0.3 pan inches per day (Tabb, Dubrow, and
Manning, 1962). It is very likely the salt content of the water
in the impounded areas increased somewhat above the peak
shown in Fig, 2 rather than decreased steadily as indicated. The
November 6 sample was diluted with fresh rain water and the
real salt peak was missed. The surface water freshened consider-
ably within two months after the storm. This agrees with pub-
lished data on Hurricane Donna of 1960 when salinities in Florida
Bay and adjacent waters returned to normal in six weeks (Tabb
and Jones, 1962).

This interpretation of salt water being the principal contribut-
ing factor in the plant kill is at variance with reasons given for
the Hurricane Donna kill in 1960. Craighead and Gilbert (1962)
reported that “. . . no abnormal increase (chlorides) followed the
receding tidal water. Probably the copious rainfall during and
just after the storm diluted the sea water.” It should also be noted
that Betsy did not leave the extensive silt and marl deposit that
characterized the area of the Donna kill and neither was impound-
ment of water behind man-made structures a problem with the
return of the Donna storm tide to the Bay.

One of the unexpected effects was the kill of sawgrass over
such an extensive area. Sawgrass is known to be euryhaline and
is frequently found growing robustly in saline and brackish areas.
Kills of this sort may be the reason for the relative thin stand of
sawgrass in these areas as compared with thick stands in more
inland locations where such salt fluctuation is not a factor.

Equally spectacular was the rapidity of spread of Eleocharis
cellulosa north of C 111 forming thick stands in what was pre-
viously an almost pure stand of sawgrass. This may be one ex-
planation of the rather mystifying pattern of distribution of these
and similar plants in the Saline Everglades. It is interesting to note

22 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

that Egler (1952) studied 50, randomly chosen, 10 sq. m. plots
in 1948 and found Eleocharis (listed as sp.) with a frequency of 83
per cent but with only rare or occasional occurrence. Observations
by the writer on the occurrence of E. cellulosa in this study and
prior to Hurricane Betsy support the type of distribution reported
by Egler. Hence, it seems possible that E. cellulosa was present
but not abundant in most of the affected area. Post storm condi-
tions seem to have favored rapid growth for Eleocharis but not
sawgrass. Measurements indicate a rate of spread as much as 10
inches a month.

Sawgrass seedlings were found three months after the storm.
Seeds germinated only in the peat soils. Seedlings were most
common in a very narrow band just above the water level at the
margins of the low hammocks. No seedlings were observed any-
where on the marl which was frequently as much as twelve inches
under water.

Algal mats are a consistant feature of the Saline Everglades.
It has been suggested that the algae are involved in the formation
of marl. There is usually a very prominent layering observable
in the mats and the upper part of the marl. Whether or not these
mats are the source of this marl is not known. Also the importance
of these mats to the ecosystem is not clearly understood. In any
event, the algal mats were affected by the sudden changes in the
water after the storm. Six months afterwards there were still
sizeable areas where the mats had not reformed or had abnormal
characteristics.

Study of the recovery of the tree islands known as low ham-
mocks during the next decade should yield information on their
ecology. It is possible that kills of this sort in the past decades have
been effective in controlling low hammock development. It is
already apparent that their flora will be different in the immediate
future than that existing prior to hurricane Betsy, September 8,
1965. Of equal interest will be the future of the sawgrass areas
that were replaced by Eleocharis cellulosa.

This report documents an important example of the effect
man-made structures can have on native vegetation. It is the
writers opinion that the vegetational kill northwest of the inter-
section of U. S. Highway 1 and Canal 111 should serve as a

ALEXANDER: Effect of Hurricane Betsy 23

warning. Wherever extensive low areas exist near seacoasts, and
where native vegetation is of primary concern extreme care should
be exercised in the building of structures that may impound storm
driven salt water.

ACKNOWLEDGMENTS

This project was supported by a National Science Foundation
Institutional Grant, University of Miami, GU 0728.

The writer wishes to express his appreciation for the coopera-
tion given by personnel of the Everglades National Park.

LITERATURE CITED

ALEXANDER, T. R. 1955. Observations on the ecology of the low hammocks
of southern Florida. Quart. Jour. Florida Acad. Sci., vol 18, no. 1,
pp. 21-27.

BERNSTEIN, LEON. 1964. Salt tolerance of plants. Agric. Inform. Bull.,
WAN Dept Agric, no. 283; pp. 1-23.

CRAIGHEAD, FRANK C, AND V. C. Giutpert. 1962. The effects of Hurricane
Donna on the vegetation of southern Florida. Quart. Jour. Florida
Aeadescin vol, 25, no. 1, pp. 1-28.

CRAIGHEAD, FRANK C. 1964. Land, mangroves, and hurricanes. Fairchild
Tropical Garden Bull., vol. 19, no. 4, pp. 5-32.

Ecuier, FRANK E. 1952. Southeast saline everglades vegetation, Florida and
its management. Vegetatio, vol. 3, fasc. 4-5, pp. 213-265.

KiEIN, Howarp. 1965. Probable effect of Canal 111 on saltwater en-
croachment, southern Dade County, Florida. U.S. Dept. of Interior,
Geological Survey, Water Resources Division, Open-File Report, Miami,
Florida, 26 pp.

LAKELA, OLGA AND F. C. CraicHEAp. 1965. Annotated checklist of the
vascular plants of Collier, Dade, and Monroe, Counties, Florida. Fair-
child Tropical Garden and University of Miami Press, Coral Gables,
Florida, 95 pp.

LLEWELLYN, W. R. 1963. Soil testing in Southern Dade County. Florida
Agriculture Extension Service, Dade County Agricultural Agents Office,
Homestead, Florida, 26 pp.

Tass, D. C., D. L. Dusrow, AND R. B. MAnninc. 1962. The ecology of the
northern Florida Bay and adjacent estuaries. Florida Board of Con-
servation, Tech. Ser., no. 39, pp. 1-81.

24 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Tass, D. C. anp A. C. Jones. 1962. Effect of Hurricane Donna on the
aquatic fauna of north Florida bay. Trans. Amer. Fisheries Soc., vol.
91; no. 4, pp. 375-378.

Department of Biology, University of Miami, Coral Gables,
Florida.

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Fishes of the St. Johns River, Florida

MARLIN E.. TAGATZ

Tue St. Johns River watershed has an area of approximately
§,350 square miles. The headwaters are about 50 miles north of
Lake Okeechobee and 15 miles inland from Florida’s east coast.
From there, the river flows northward 260 miles to the vicinity of
Jacksonville, and then swings eastward for about 25 miles before
emptying into the Atlantic Ocean. Tidal influence extends as far
upstream as Lake George, 115 miles above the mouth. Salinity
varies greatly in the lower reaches, but south of Jacksonville it is
normally less than 1 0/oo.

This system is of particular interest because of the extent to
which marine forms penetrate the strictly fresh waters in the river’s
upper reaches. Such fishes as Archosargus probatocephalus, Sciae-
nops ocellata, Micropogon undulatus, Pogonias cromis, Lutjanus
griseus, and Dasyatis sp. have been found far upstream. As a re-
sult, the so-called primary-division freshwater fishes, i.e., those
fishes that are intolerant of salt water (Myers, 1938, 1951), often
occur together with various marine species.

The St. Johns estuary is a nursery area for the young of many
marine fishes that spawn in the ocean, and also is a feeding and
spawning area for various adult marine fish. Spawning popula-
tions in the river consist of species that are anadromous, those that
extend their breeding range into the river, and some that have
established isolated resident populations in the fresh water.

The abundance of marine fishes beyond the zone of brackish
water (salinity of 50/oo or less) probably is due to the configura-
tion of the river and to the presence of moderate traces of salt in
its upper reaches. Many tributary creeks and large coves along
the lower 25 miles of river provide areas of intermediate salinity
where fish can undergo gradual transition from salt to fresh water.
Odum (1953) stated that the main reasons for the extensive move-
ment of marine forms into the St. Johns system appear to be the
abundance of calcium chloride in the water and the presence of
the salt springs that drain into the river. Salinity actually increases
with distance upstream between Palatka and Lake George from
the discharge of a number of these springs ( Beck, 1965).

The boundary between brackish and fresh water, which moves

26 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

back and forth from its usual location in southern Jacksonville, de-
termines the upstream or downstream penetration of many species.
The following selected salinity readings indicate the effect of tidal
stage and runoff on the vertical and horizontal location of brackish
waters. In October, at low tide, salinity in the channel, 29 miles
upstream, was ()0/oo at the surface and 4.6 0/oo at the bottom; at
high tide, it was 10.2 0/oo at the surface and 10.9 0/oo at the bot-
tom. Salinity on the channel bottom normally is greater than that
at the surface. Freshwater fishes rarely occur downstream of the
boundary between brackish and fresh water, and, conversely, marine
forms that cannot tolerate fresh water seldom occur upstream of
the boundary. For example, Ictalurus catus and I. punctatus were
the only freshwater forms collected at salinities above 1 o0/oo (up
to 90/oo).

The abundance and size of many fishes changes seasonally with
temperature. Gunter (1945) stated that the temperature cycle was
more definite than general salinity changes and was chiefly respon-
sible for the seasonal movements and other recurrent activities of
marine fishes in Texas. The occurrence in the St. Johns of marine
forms that reach the northern or southern limit of their ranges off
northeast Florida is particularly dependent on favorable seasonal
temperatures.

Various scientists have contributed to knowledge of the fish
fauna of the St. Johns River, particularly since the turn of the cen-
tury. The first comprehensive checklist of Florida fishes included
112 species from the St. Johns, of which 28 were found only at the
mouth (Evermann and Kendall, 1900). Fowler (1945) included
a number of records from the system in his study of fishes of the
southern Piedmont and Coastal Plain. The most extensive study
was a 10-year investigation by McLane (1955), who provided an
annotated list of the species known to occur in the St. Johns drain-
age, but excluded those forms found only near the mouth. Of the
118 forms he reported, 52 are strictly freshwater species. Some of
McLane’s data (collected in 1953) were from a long-term study of
the exploited fish populations of the river (Moody, 1961). Carr
and Goin (1959) reported the occurrence of certain marine species
in the St. Johns in their guide to the freshwater fishes of Florida.
Briggs (1958) summarized the ranges of St. Johns River fishes in
his checklist of Florida species. 3

Tacatz: Fishes of St. Johns River 27

From April 1961 to November 1963, the seasonal occurrence of
fishes in trawl and seine collections was noted during a study of
the blue crab (Callinectes sapidus) in the St. Johns River by the
Bureau of Commercial Fisheries. Capture of fish was incidental
to the “random” or regular sampling of juvenile crabs. The num-
ber and size range of many of the fish collected from 10 to 135
miles upstream were summarized and a complete checklist was
prepared.

A 70-foot seine having a funnel-shaped bag 15 feet long (net,
%e-inch, and bag, 5/32-inch stretched mesh) and an 8-foot trawl
(net, %-inch, and bag, 44-inch stretched mesh) were used to obtain
nearly all of the fish. Specimens of Sphyrna lewini were obtained
from commercial fishermen.

At the time of each collection we determined surface salinity
in parts per thousand (hydrometer readings corrected for tempera-
ture) and surface water temperature in degrees centigrade. All
salinity readings beyond 30 miles upstream were less than 1 0/00
and were recorded as 0 0/oo.

Specimens in each sample were identified, counted, and meas-
ured in the field. Representative individuals from almost all col-
lections were preserved in 10 per cent formalin and sent to the
Florida State Museum, Gainesville, Fla., for identification and in-
clusion in its collection of fishes. Lengths of fishes with forked
tails were measured from the tip of the snout to the caudal fork;
those with nonforked tails were measured to the tip of the longest
caudal ray. Fork lengths were taken in preference to other meas-
urements because of the ease and speed with which they could be
obtained.

COLLECTING LOCALITIES

Collections were obtained from 12 localities or stations that were
assigned letters from A to L according to distance upstream ( Fig.
1). Fish from tributary streams (stations A, C, E, and F) were
captured 1 to 3 miles from the mouth of the tributary. Specimens
from the main river and from Lake George (stations B, D, and G
to L) were taken in the channel and adjacent waters.

The various stations represented different types of habitat. The
bottom material at stations C, E, and F was muck; at all others it
was firmer mud and sand. Bottom vegetation was most dense at
stations H and I where it consisted of coontail-moss (Ceratophyllum

28 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

DUNN CLAPBOARD
CREEK CREEK

TROUT RIVER JETTIES

oo \S : - MAYPORT

ty JACKSONVILLE BEACH °

ORTEGA RIVER

DOCTORS
LAKE

© SWITZERLAND

(H) ST. AUGUSTINE

rv,
=
—
Oo
x=
z
“
ba
<
m
z

FLORIDA

OKLAWAHA
RIVER

LAKE
GEORGE

Fig. 1. Location of the 12 collecting stations, A to L, in the St. Johns
River.

Tacatz: Fishes of St. Johns River 29
sp.) and eelgrass (Vallisneria sp.). The maximum depths were 6
feet at station C, 11 feet at F and K, and more than 20 feet at all
other stations.

The following is a complete list of collections, numbered con-
secutively by station and date. Data for each station are given in
the order: year, collection number (in parentheses), month and
day, surface salinity (o/oo) and surface temperature (°C). Salin-
ity is omitted for stations G to L, where all readings were 0 0/oo.

Station A (10 miles upstream ).
IGGIE Gl
(3) June 7—21.5, 28.8

HOG2 eens) Mian. 13—28.2> 17.4:
(6) Apr. 9—25.8, 19.3;
1963: (8) Feb. 14—7.7, 11.0;

10) Mar. 5—9.5, 19.0;

12) Apr. 9—22.1, 21.3;

14) Apr. 23—15.4, 23.5;
6) May 2223.1, 27.0;

~~

(
(
(
(1
(1
(20) July 23—14.5, 29.2;
(2

(24
(26

Station B (13 miles upstream )

1962; (1) Mar. 13—28.0, 17.0;
(3) June! 623.7, 28.5:
(5) Sept. 6B—16.0, 28.3;
1963: (7) July 23—19.6, 30.0

Station C (20 miles upstream )
HOGI lo) Oct 2413.4, 22.9.
Dec. 19—9.0, 19.8

1962: Jan. 15—24.5, 12.8;

July 12—17.5, 31.4;

Dec. 18—10.3, 13.0
Jan. 17—6.9, 14.0;
Mar. 4—0, 17.5;
Mar. 18—5.2, 21.0;
Apr. 10—10.5, 21.5;
May 8—21.3, 24.0;
June 4—22.7, 26.4;

)
)
)
)
)
)
)
) Nov. 19—8.8, 18.0;
)
1963: )
)
)
)
)
)

) April 18—14.6, 20.0;

(
(
(11) Mar. 21—11.4, 18.5;

GIS) Aor 22=—119) 552.7, 0:

8) June 18—22.5, 28.7;
0)
2) Aug. 26—20.1, 29.5;
)

Sept. 30—10.3, 24.0:
Nove 1S"=13:1) 17.4;

Mar. 29—18.0, 21.5;
May 25—25.0, 29.5;

Aug. 16—14.0, 31.0;
Sept. 17—10.2, 29.1;

(2) May 17—27.3, 25.2;

(5) Mar. 29—20.8, 21.5
7) Sept. 19—15.8, 30.0

9) Feb. 20—10.8; 13.0;
(15)) May 7—26°8, 23.5:
(17) June 5—24.7, 26.6;
(19) July 3—14.5, 27.4;
(21) Aug. 13—15.8, 29.8;
(23) Aug. 27—25.2, 28.7;
(25) Oct, 4—7.9, 26.2;

(2) May 15—30.2, 26.5;
(4) Aug. 6—14.1, 30.1;
(6) Sept. 19—15.8, 30.0

(2) Nov. 20—13.7, 20.3;

(5) Feb. 19—22.2, 18.5:
Co pelo 9010) 23.5.
(9) June 14—19.9, 29.2;
(11) Aug. 10—0, 30.6;
(13) Aus. 30—13.7, 30.8;
Cis) Octk<24——11 14 2120;
Chie) Decwlv— 120" 1250:

C20) Keb 4 earl 035:
(22) Mars —0. 16.8:

(24) Mar. 19—3.9, 25.7;
(26) Apr. 23—6.7, 23.0;

(28) May 21—16.6, 26.2:
(30) June 17—10.3, 28.5;

30 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

(31) July 2—0, 27.5; (32) July 23—5.8, 29.1;
(33) Aug. I3—I33, 31.0:) (84) Aue, 2676s oile>-
(35) Sept. 9—4.0, 28.5; (36) Sept. 30—O0, 25.4;
(37), Oct 8—0) 2451 (38) Oct 1810823105
(39) Nov.
Station D (22 miles upstream )

1961: (1) Aug. 16—24.5, 29.0

1962: (2) May 29—13.7, 29.7

1963:9 (3) Jans 29-1873; 1-7

Station EF, (25 miles upstream )
I961; (1) Apr. 18—5.7, 21.0; (2) May 9—Ill3n2a8:

(3) June 7—14.3, 30.8; (4) Aus. 16—13:5, 28:2;
(5) Oct. 24—9.5, 22.0; (6) Nov. 20—8.8, 20.5;
(7) Dec. 19—10.0, 20.5
1962: (8) Jan. 15—17.8, 14.0; (9) Feb. 19—19.0, 21.2:
(10) Mar. 29—16.9, 18.5; (11) May 25—22.1; 33.8;
(12) June 14—14.9, 29.9; (13) July 12—15.3, 34.1;
(14) July 27—11.0, 30.0; (15) Aug. 10—O, 32.6;
(16) Aug. 16—9.0, 31.8; (17) Sept. 17—8.4, 30.8;
(18) Oct. 24—8.0, 21.0; (19) Nov. 19—5.5, 19:4:
(20) Dec. 17—10.8, 11.0
1963: (21) Jan. 17—5.2, 13.0; (22) Mar. 6—2:0; 19:5;
(23) Mar. 20—3.0, 23.5; (24) Apr. 8—13.9, 20.3;
(25) Apr. 24—6.6, 24.3; (26) May 6—17.9, 22.8:
(27) May 20—14.9, 27.0; (28) June 3—22.1, 26.7;
(29) June 19—11.9, 29.1; (30) July 1—0, 29.8;
(31) July 16—10.5, 30.2:: (32) July 24—=siGR2 9:3;
(33) Aug. 14—12.2, 29.8; (34) Aug. 27—9.0, 31.0;
(35) Oct: 62 248= (386) nOck9— 0 ears.
(37) Nov. 20—2.2, 16.5

Station F (30 miles upstream )
1962: (1) May 29-119, 3274
Station G (40 miles upstream )
1961: (1) Apr. 17—20.0; . (2) May 8—25.5; (3) Jume 19—25:0
Station H (60 miles upstream )
1961: (1) Apr. 17—21.0; (2) Apr. 19—21.0; (3) May 8—25.5:
) June 16—29.0; (5) Aug. 25—29.5; (6) Oct. 23—24'5:
) Nov. 21—20.0; (8) Dec. 13—20.2
) Jan. 19—12.0; -(10) Feb. 21—19.0; (11) Mar. 28—20.5;
2) Apr. 20—21.2;. (13) May 14—26.8; (14) June 13—28.0;
) July 17—29.0; (16) July 30—29.3; (17) Aug. 7—30.5;
) Aug. 13—31.6; (19) Aug. 15—34.0; (20) Sept. 14—31.0:
) Oct. 19—241: (22) Nov. 20/7; 4(23)) 2 Decwia==se5
1963: (24) Jan. 23—14.0;. (25) Feb. 13—13.0; (26) Mar. 12—20.0;
(27) Mar. 25—20.0; (28) Apr. 11—22.2; (29) Apr. 26—23.8;
(30) May 9—25.0; (31) May 23—28.0; (32) June 14—28.9;
(33) June 24—28.8;. (34) July 11—28.4; (35) July 29—29.5;

Tacatz: Fishes of St. Johns River 31

(36) Aug. 1230.0; (37) Aug. 2829.8: (38) Sept. 1626.0:
om@ct 2245. (40) Oct. 10-235. (41) Nov. 21166
Station I (84 miles upstream )

1961: (1) Apr. 20—21.5; (2) May 11—19.0; (3) June 8—29.0;
(4) Aug. 29—28.8; (5) Oct. 12—25.2; (6) Nov. 28—20.8;
(7) Dec. 12—21.1

1962: (8) Jan. 16—12.9; (9) Feb. 12—16.0; (10) Mar. 28—19.7;
(11) Apr. 25—25.8; (12) May 28—30.2; (13) June 13—32.8;
(14) July 30—33.5; (15) Aug. 17—30.0; (16) Sept. 18—29.7;
(17) Sept. 26—30.0; (18) Oct. 12—27.4; (19) Nov. 21—19.0;
(20) Dec. 14—8.8

1963: (21) Jan. 23—15.5

Station J (105 miles upstream )
1961: > (1) May 11—22.0; (2) June 22—26.0; (3) Oct. 17—23.0;
(4) Nov. 23—19.9: (5) Dec. 21—17.8
IQo2 Reo) mane 2——1 2.0. (7) Keb. 15—20.5= (8) Dec. 19——12.2

Station K (120 miles upstream )
NOG Gle) Apr. 21=—20'5: (2) May 12—23.8: (3): June 20—26.5.

(4) July 10—29.0

1962: (5) May 2329.0

Station L (135 miles upstream )
LOGIE Gl meAors 2121.0 (2) May 12—28.5; (3) June 20—26.8

FISHES

The fish fauna of the St. Johns River consists of 55 species that
are typical freshwater forms and 115 that are euryhaline (occur
both in sea water and water of moderate to zero salinity). One
species, Cyprinodon hubbsi, is endemic to the river system. Fam-
ilies particularly well represented are Clupeidae, Cyprinodontidae,
Centrarchidae, Sciaenidae, and Gobiidae.

The list of marine fish for the St. Johns would be longer if fish
collected in the river within a few miles of the mouth had been
included. In this category, I collected Anchoa lyolepis (dusky
anchovy), Hippocampus erectus (spotted seahorse), Pomatomus
saltatrix (bluefish), and Sphaeroides spengleri (bandtail puffer).

The following list is a taxonomic grouping of all records of fish
for the St. Johns known to me, 170 species representing 55 families.
This listing is primarily based on 116 species reported by McLane
(1955) and 51 records from the present study. The changes that
were made in the records given by McLane are: Notropis sp. to N.
welaka, Ogcocephalus sp. to O. cubifrons, Ictalurus platycephalus
to I. brunneus (validated by Dr. Ralph Yerger, Florida State Uni-

32, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

versity) and deletion of Alosa pseudoharengus (a 19th-century
record considered in error). The checklist also contains Paralich-
thys albigutta (reported in Carr and Goin, 1959), and, from speci-
mens in the Florida State Museum collection, Elassoma okefenokee
and Umbra pygmaea. Nomenclature follows that of Bailey, Lach-
ner, Lindsey, Robins, Roedel, Scott, and Woods (1960).

For each of the 101 euryhaline species collected in the present
study, I have included a summary of the stations (collection num-
ber in parentheses) at which it was taken, the ranges in surface
salinity and temperature when collected, and number and length
range of fish, by month. Salinity is in parts per thousand, water
temperature in degrees C, and length (FL) in mm. Some speci-
mens were not identified to species and are listed as undetermined,
by genus or family. In addition to Anchoa hepsetus and A. mitch-
illi, one other engraulid (of uncertain identity) was collected and
is referred to as Anchoa sp. Fourteen euryhaline species, those
reported for the river but not obtained in this study, are listed with
the notation “None captured.”

Entered without annotation are the 55 species that are con-
fined to fresh water (indicated by an F). Except for an adult
specimen of Eleotris picta captured at station H on August 13, 1962
(first record of occurrence in the river) my samples of strictly
freshwater forms provided little new information.

Fifty-one species not previously recorded are designated by an
asterisk.

Family PETROMYZONTIDAE

Petromyzon marinus Linnaeus. Sea lamprey. None captured.

Family SpHyYRNIDAE

Sphyrna lewini (Griffith and Smith). Scalloped hammerhead*.
Station: B(3). Salinity 23.7, temperature 28.5. Number and
length: June (2, 324-381).

Family DAsyATIDAE

Dasyatis americana Hildebrand and Schroeder. Southern sting-
ray. None captured.
Dasyatis sabina (Lesueur). Atlantic stingray. Stations: A(6),

Tacatz: Fishes of St. Johns River 33

Paar 20) ee ( 7), D(1L), E(37), H(12, 16, 22, 38), 1(8, 9, 13, 16,
18-21), J(3-6, 8). Salinity 0-25.8, temperature 8.8-32.8. Number and
length: Jan. (5, 495-635); Feb. (3, 478-533); Apr. (3, 533-711);
June (2, 508-521); July (2, 394-660); Aug. (2, 330-700); Sept. (2,
310-432); Oct. (2, 500-508); Nov. (5, 381-711); Dec. (8, 432-648).

Family ACIPENSERIDAE

Acipenser brevirostrum Lesueur. Shortnose sturgeon. None
captured.

Acipenser oxyrhynchus Mitchill. Atlantic sturgeon. None cap-
tured.

Family LEPIsOSTEIDAE

Lepisosteus osseus (Linnaeus). Longnose gar. F
Lepisosteus platyrhincus DeKay. Florida gar. F

Family AMUDAE

Amia calva Linnaeus. Bowfin. F

Family ELOPMAE

Elops saurus Linnaeus. Ladyfish. Stations: A(25), C(33),
F(20, 37), H(17), 1(12). Salinity 0-13.3, temperature 11.0-31.0.
Number and length: May (1, 51); Aug. (3, 198-250); Oct. (1, 305);
Nova 195): Dec. (1, 239).

Megalops atlantica Valenciennes. Tarpon. None captured.

Family CLUPEDAE

Alosa aestivalis (Mitchill). Blueback herring. Stations: C(7),
L(2). Salinity 0-20.0, temperature 23.5-28.5. Number and length:
April (1, 28); May (1, 39).

Alosa mediocris (Mitchill). Hickory shad. None captured.

Alosa sapidissima (Wilson). American shad. None captured.

Brevoortia smithi Hildebrand. Yellowfin menhaden. Stations:
A(24); D(3). Salinity 10.3-18.3, temperature 11.7-24.0. Number
and length: Jan. (2, 201-266); Sept. (15, 228-254).

Brevoortia tyrannus (Latrobe). Atlantic menhaden*. Stations:

34 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

A(25), C(13). Salinity 7.9-13.7, temperature 26.2-30.8. Number
and length: Aug. (1, 123); Oct. (10, 69-74).

Dorosoma cepedianum (Lesueur). Gizzard shad. Stations:
C(22), G(3), HO, 37), 1(5, 6, 8, 9, 12. lay 20) Gee eee
Salinity 0, temperature 8.8-30.5. Number and length: Jan (4, 99-
121); Feb. (2, 55-64); Mar. (1, 111); May (1, 287); June 2, 75-80);
July (9, 25-157); Aug. (12, 90-272): Oct. (lO\S1E234) ee Nowe
115-220); Dec. (16, 68-237).

Dorosoma petenense (Gunther). Threadfin shad. None cap-
tured.

Harengula pensacolae Goode and Bean. Scaled sardine*. Sta-
tions: H(4, 33), I1(3, 12), J(2). Salinity 0, temperature 26.0-30.2.
Number and length: May (2, 23-27); June (103, 24-46).

Opisthonema oglinum (Lesueur). Atlantic thread herring*.
Stations: A(15, 16, 22, 23), B(6), 1(20). Salinity 0-26.8, tempera-
ture 8.8-30.0. Number and length: May (160, 21-53); Aug. (48,
73-121); Sept. (1, 59); Dec. (2, 73-81).

Undetermined Clupeidae. Stations: A( 10, 140 13,20) Gis
25-28, 32-34), E(22-24, 27), I1(4). Salinity 0-22.5, temperature
17.5-31.5._ Number and length: Mar. (19, 25-125); Apr. (50, 22-
97); May (58, 22-59); June (1, 61); July (45, 43-70); Aug. (46,
26-69 ).

Family ENGRAULIDAE

Anchoa hepsetus (Linnaeus). Striped anchovy. Stations: A(3,
12,24, 25), B(6); C(3), D(2), EC3, 12). ‘Salinity, 719222 eieatemd
perature 19.8-30.8. Number and length: Apr. (1, 94); May (2, 37-
45); June (23, 17-62); Sept. (2, 70-74); Oct. (2, 52-71) sWecm Gr
89).

Anchoa mitchilli (Valenciennes). Bay anchovy. Stations: A(2,
3, 7-9, 12, 14-22, 24, 25), B(6), C(1-9, 11-17, 19-22) 24-34536=39)),
E(3, 5-9, 11, 13-17, 21-29, 35, 37), FC), ©(1-3) (45 Gos ihiea
19, 23, 27; 28; 33) 35, 37, 40), 1¢ 7-11, 14, 15, 20). iG 27a ease kimnig,
0-27.3, temperature 8.3-34.1. Number and length: Jan. (281, 21-60);
Feb. (26, 31-75); Mar. (92, 29-71); Apr. (321, 40-78); May (658,
23-77); June (738, 19-73); July (897, 21-71); Aug. (1,501, 21-83);
Sept. (153, 19-62); Oct. (3,189, 10-68); Nov. (119, 22-61); Dec. (63,
27-71).

Tacatz: Fishes of St. Johns River 35

Anchoa sp.* Station: E(11). Salinity 22.1, temperature 33.8.
Number and length: May (1, 37).

Family UMBRIDAE

Umbra pygmaea (DeKay). Eastern mudminnow. F

Family EsociwarE

Esox americanus Gmelin. Redfin pickerel. F
Esox niger Lesueur. Chain pickerel. F

Family SYNODONTIDAE

Synodus foetens (Linnaeus). Inshore lizardfish*. Stations:
Pieleelia-21 24°25), B(6), C(8-10, 29, 36), E(12-14, 28, 29, 35).
Salinity 0-26.8, temperature 13.0-34.1. Number and length: Feb.
(1599): May (13, 39-104); June (73, 36-118); July (26, 37-136);
Aug. (1, 49); Sept. (8, 22-63); Oct. (9, 45-83).

Family CypriInIDAE

Hybopsis harperi (Fowler). Redeye chub. F

Notemigonus crysoleucas (Mitchill). Golden shiner. F
Notropis chalybaeus (Cope). Ironcolor shiner. F

Notropis cummingsae Myers. Dusky shiner. F

Notropis hypselopterus (Gunther). Sailfin shiner. F
Notropis maculatus (Hay). Taillight shiner. F

Notropis petersoni Fowler. Coastal shiner. F

Notropis welaka Evermann and Kendall. Bluenose shiner. F
Opsopoeodus emilae Hay. Pugnose minnow. F

Family CaTosTOMIDAE

Erimyzon sucetta (Lacépede ). Lake chubsucker. F

Family ARIDAE

Bagre marinus (Mitchill). Gafftopsail catfish. Station: D(3).
salinity 18.3, temperature 11.7. Number and length: Jan (1, 116).
Galeichthys felis (Linnaeus). Sea catfish. Stations: A(1, 6, 7),
C(12-14, 17, 25, 27), D(1), E(20). Salinity 10.2-25.8, temperature

36 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

11.0-31.0._ Number and length: Apr. (7, 86-186); May (1, 112);
Aug. (12, 74-187); Sept. (8, 59-103); Dec. (3, 81-91).

Family IcrALURDAE

Ictalurus brunneus (Jordon). Snail bullhead. F
Ictalurus catus (Linnaeus). White catfish. F
Ictalurus natalis (Lesueur). Yellow bullhead. F
Ictalurus nebulosus (Lesueur). Brown bullhead. F
Ictalurus punctatus (Rafinesque). Channel catfish. F
Noturus gyrinus (Mitchill). Tadpole madtom. F
Noturus leptacanthus Jordan. Speckled madtom. F

Family ANGUILLIDAE

Anguilla rostrata (Lesueur). American eel. Stations: C(30),
E(27, 28, 32, 35), G(1), H(1, 611, 14, 16, 18, 19, 22, 23, 25-28, 30-
33, 30, 36, 38-40, 42), I(1, 7, 8, 21), J( 5-6), K(2,4). Salimity 0222m¢
temperature 8.3-34.0. Number and length: Jan. (31, 52-533); Feb.
(6, 155-635); Mar. (16, 136-508); Apr. (7, 115-495); May (10, 165-
546); June (6, 118-508); July (11, 152-610); Aug. (12, 74-576);
Sept. (2, 274-379); Oct. (9, 251-611); Nov. (10, 120-512); Dee. (10,
279-533 ).

Family BELONIDAE

Strongylura marina (Walbaum). Atlantic needlefish. Stations:
AQT, 25), C(31), E(28, 30, 32), Hi( 15; 17, 29) eG es alana

74); June (3, 136-198); July (5, 76-483); Aug. (6, 252-545); Sept.
(2, 114-324); Oct. (1, 356).

Family CyPRINODONTIDAE

Cyprinodon hubbsi Carr. Lake Eustis minnow. F

Cyprinodon variegatus Lacépede. Sheapshead minnow. Sta-
tion: E(30). Salinity 0, temperature 29.8. Number and length:
July (19):

Fundulus chrysotus (Ginther). Golden topminnow. F

Fundulus cingulatus Valenciennes. Banded topminnow. F

Fundulus confluentus Goode and Bean. Marsh killifish. Sta-
tions: A(9), C(16). Salinity 8.8-10.8, temperature 13.0-18.0. Num-
ber and length: Feb. (5, 50-56); Nov. (10, 33-53).

Tacatz: Fishes of St. Johns River Sih

Fundulus heteroclitus (Linnaeus). Mummichog. _ Stations:
A(9, 21, 22, 25), C(16, 18, 21, 27-29, 31, 35, 37), E(22-25, 27-31,
33). Salinity 0-22.7, temperature 13.0-30.2. Number and length:
Feb. (7, 50-57); Mar. (42, 6-64); Apr. (11, 54-66); May (52, 34-68):
June (81, 24-93); July (48, 17-78); Aug. (6, 34-53); Sept. (2, 25-
30); Oct. (6, 28-55); Nov. (15, 31-63); Dec. (25, 35-65).

Fundulus majalis (Walbaum). Striped killifish. Station: B(6).
Salinity 15.8, temperature 30.0. Number and length: Sept. (2, 75-
io)

Fundulus notti (Agassiz). Starhead topminnow. F

Fundulus seminolis Girard. Seminole killifish. F

Fundulus similis (Baird and Girard). Longnose killifish*. Sta-
tions: A(9), B(5, 6). Salinity 10.8-16.0, temperature 13.0-30.0.
Number and length: Feb. (4, 60-62); Sept. (6, 22-56).

Jordanella floridae Goode and Bean. Flagfish. F

Leptolucania ommata (Jordan). Pygmy killifish. F

Lucania goodei Jordan. Bluefin killifish. F

Lucania parva (Baird and Girard). Rainwater killifish. Sta-
tion: H(18, 20). Salinity 0, temperature 31.0-31.6. Number and
length: Aug. (5, 22-30); Sept. (9, 29-43).

Family PorcILiDAE

Gambusia affinis (Baird and Girard). Mosquitofish. Stations:
C(28, 29), H(17). Salinity 0-22.7, temperature 26.2-30.5. Num-
ber and length: May (19, 34-40); June (10, 31-43); Aug. (2, 42-43).

Heterandria formosa Agassiz. Least killifish. F

Poecilia latipinna (Lesueur). Sailfin molly. Stations: C(17),
E(24). Salinity 12.0-13.9, temperature 12.0-20.3. Number and
length: Apr. (1, 38); Dec. (1, 44).

Family GADIDAE

Urophycis floridanus (Bean and Dresel). Southern hake*. Sta-
tion: A(8, 9). Salinity 7.7-10.8, temperature 11.0-13.0. Number
and length: Feb. (9, 34-120).

Urophycis regius (Walbaum). Spotted hake*. Station: A(8,
9). Salinity 7.7-10.8, temperature 11.0-13.0. Number and length:
Feb. (8, 36-121).

38 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Family SyYNGNATHIDAE

Oostethus lineatus (Kaup). Opossum pipefish. None captured.

Syngnathus fuscus Storer. Northern pipefish*. Station: E(7).
Salinity 10.0, temperature 20.5. Number and length: Dec. (1, 108).

Syngnathus louisianae Gunther. Chain pipefish. Stations: A(1,
15-17), C(15, 29, 36), E(11, 12, 35, 36). Salinity 0-26.8, tempera-
ture 20.0-33.8. Number and length: Apr. (1, 195); May (3, 115-
298); June (5, 56-146); Sept. (6, 78-166); Oct. (6, 76-213).

Syngnathus scovelli (Evermann and Kendall). Gulf pipefish.
Stations: A( 1), C(6), H(9, 11, 22, 25, 37), Hi 1oeise2 On 2) aaanee
J(8), K(5). Salinity 0-22.1, temperature 12.2-33.8. Number and
length: Feb. (2, 56-135); Mar. (13, 88-126); Apr. (4, 64-130); May
(2, 70-141); July (2, 77-136); Aug. (5, 77-107); Sept. (3, 113-126);
Nov. (5, 81-121); Dec. (6, 71-99).

Undetermined Syngnathidae. Stations: A(3, 14, 19, 25), C(30,
31, 34, 37, 38), E(2, 4, 27-30). Salinity 0-22.1, temperature 23.0-
31.5. Number and length: Apr. (1, 68); May (2, 68-126); June (6,
69-156); July (9, 32-157); Aug. (8, 36-179); Oct. (14, 78-156).

Family APHREDODERIDAE

Aphredoderus sayanus (Gilliams). Pirate perch. F

Family CENTROPOMIDAE

Centropomus undecimalis (Bloch). Snook. None captured.

Family SERRANIDAE

Centropristes philadelphicus (Linnaeus). Rock sea bass*. Sta-
tion: A(8). Salinity 7.7, temperature 11.0. Number and length:
Feb. (2, 129-131).

Centropristes striatus (Linnaeus). Black sea bass*. Station:
A(8, 11, 17, 19, 21, 26). Salinity 7.7-24.7, temperature 11.0-29.8.
Number and length: Feb. (1, 91); Mar. (2, 101-122); June (1, 28);
July 38S—7 Ip) Acree (alee 6))/- Noyes elenk7G))

Mycteroperca microlepis (Goode and Bean). Gag*. Station:
B(6). Salinity 15.8, temperature 30.0. Number and length: Sept.
(CL MS

Roccus saxatilis (Walbaum). Striped bass. Station: J(1).
Salinity 0, temperature 22.0. Number and length: May (1, 46).

Tacatz: Fishes of St. Johns River 39

Family LuryANIDAE

Lutjanus analis (Cuvier). Mutton snapper*. Station: E(12).

Salinity 14.9, temperature 29.9. Number and length: June (1, 65).

Lutjanus griseus (Linnaeus). Gray snapper. Stations: A(25),

C(35-37), E(8, 20, 35-37), H(42). Salinity 0-17.8, temperature
11.0-28.5. Number and length: Jan. (1, 67); Sept. (11, 19-54);
Oct. (27, 24-58); Nov. (4, 40-58); Dec. (1, 105).

Family CENTRARCHIDAE

Acantharchus pomotis (Baird). Mud sunfish. F

Centrarchus macropterus (Lacépéede). Flier. F
Chaenobryttus gulosus (Cuvier). Warmouth. F

Elassoma evergladei Jordan. Everglades pygmy sunfish. F
Elassoma okefenokee Bohlke. Okefenokee pygmy sunfish. F
Elassoma zonatum Jordan. Banded pygmy sunfish. F
Enneacanthus chaetodon (Baird). Blackbanded sunfish. F
Enneacanthus gloriosus (Holbrook). Bluespotted sunfish. F
Enneacanthus obesus (Girard). Banded sunfish. F
Lepomis auritus (Linnaeus). Redbreast sunfish. F

Lepomis macrochirus Rafinesque. Bluegill. F

Lepomis marginatus (Holbrook). Dollar sunfish. F
Lepomis microlophus (Ginther). Redear sunfish. F
Lepomis punctatus (Valenciennes). Spotted sunfish. F
Micropterus salmoides (Lacépéde). Largemouth bass. F
Pomoxis nigromaculatus (Lesueur). Black crappie. F

Family PERCIDAE

Etheostoma fusiforme barratti (Holbrook). Scalyhead darter. F
Etheostoma edwini (Hubbs and Cannon). Brown darter. F
Etheostoma nigrum Rafinesque. Johnny darter. F

Percina nigrofasciata (Agassiz). Blackbanded darter. F

Family CARANGIDAE

Caranx hippos (Linnaeus). Crevalle jack. Stations: A(25),

B(7), C(28, 32), E(30). Salinity 0-19.6, temperature 26.2-30.0.
Number and length: May (5, 33-54); July (5, 38-106); Oct. (1, 47).

Chloroscombrus chrysurus (Linnaeus). Bumper. Stations:

A(20, 22), B(6), C(2), E(7). Salinity 10.0-20,1, temperature 20.3-

40 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

30.0. Number and length: July (3, 22-52); Aug. (2, 61-67); Sept.
(3, 44-48); Nov. (12, 48-65); Dec. (1, 105).

Oligoplites saurus (Bloch and Schneider). Leatherjacket*.
Station: B(6). Salinity 15.8, temperature 30.0. Number and
length: Sept. (1, 55).

Selene vomer (Linnaeus). Lookdown*. Stations: A(19), E(29,
32, 35). Salinity 6.2-14.5, temperature 24.8-29.3. Number and
length: June (1,59) July (6, 27-76): Oct) (IN s3a)r

Trachinotus falcatus (Linnaeus). Permit*. Station: E(31).
Salinity 10.5, temperature 30.2. Number and length: July (1, 50).

Vomer setapinnis (Mitchill). Atlantic moonfish*. Station:
A(20). Salinity 14.5, temperature 29.2. Number and length: July
(dh, Sieh)

Family GERRIDAE

Diapterus olisthostomus (Goode and Bean). Irish pompano.
Station: C(32, 34-36). Salinity 0-7.6, temperature 25.4-31.5. Num-
ber and length: July (2, 26-40); Aug. (2, 40-68); Sept. (16, 24-78).

Diapterus plumieri (Cuvier). Striped mojarra*. Station: C(13,
15). Salinity 11.1-13.7, temperature 21.0-30.8. Number and length:
Aug. (2, 71-73); Oct. (3, 63-76).

Eucinostomus argenteus Baird and Girard. Spotfin mojarra.
Stations: A(20-22, 24, 25), B(6), C1, 13) 15, l6NS2ee4e3Gmeo)e
E(11, 12; 14, 17, 18, 31, 32, 36,37), H( 17-19) iC Gaae es alimony,
0-22.1, temperature 16.5-34.0. Number and length: May (1, 26);
June (2, 30-47); July (36, 36-85); Aug. (13, 28-98); Sept. (15, 22-
104); Oct. (61, 16-115); Nov. (134, 15-111); Dec. (2, 45-52).

Eucinostomus gula (Quoy and Gaimard). Silver Jenny*. Sta-
tions: A( 20-22), B(6), C(10), E(13, 31). Salinity 10.5-20.1, tem-
perature 29.2-34.1. Number and length: July (16, 46-83); Aug. (6,
78-100); Sept. (2, 91-103).

Undetermined Gerridae. Stations: A(18, 19), C(2, 3, 37, 38),
E:(6, 29, 30, 33-35). Salinity 0-22.5, temperature 19.8-31.0. Num-
ber and length: June (41, 36-58); July (66, 46-69); Aug. (19, 79-
95); Oct. (37, 22-102); Nov. (12, 46-73); Dec. (8, 44-73).

Family POMADASYIDAE

Orthopristis chrysopterus (Linnaeus). Pigfish. Stations: A(15-

Tacatz: Fishes of St. Johns River Al

17, 24), C(8), E(11-14, 16, 27, 32, 34, 35), H(15). Salinity 0-26.8,
temperature 23.5-34.1. Number and length: May (74, 17-54); June
(4739-69); July (19, 67-97); Aug. (3, 82-113); Sept. (1, 142); Oct.
(2, 115-118).

Family SCIAENIDAE

Bairdiella chrysura (Lacépéede). Silver perch. Stations: A(2,
Peostinie 19), C(2, 10, 11, 13, 14, 16, 29, 32, 38), D(3), E(3,
PeOwmlomliaw 22023, 28-30, 34), H(15, 17-20, 37). Salinity 0-27.3,
temperature 11.7-34.1. Number and length: Jan. (1, 142); Feb. (1,
81); Mar. (13, 104-137); May (10, 46-100); June (117, 22-97); July
(1315 45-124); Aug. (25, 71-122); Sept. (6, 83-91); Oct. (2, 135-
141); Nov. (3, 74-91).

Cynoscion nebulosus (Cuvier). Spotted seatrout. Stations:
C(15, 32, 34-38), E(18, 22, 35, 36), H(17). Salinity 0-11.1, tem-
perature 19.5-31.5. Number and length: Mar. (3, 97-118); July
(4, 64-84); Aug. (13, 13-136); Sept. (56, 15-128); Oct. (87, 19-173).

Cynoscion regalis (Bloch and Schneider). Weakfish*. Sta-
MONSEEP A Owlo)). (4), C(9 13), E(3). Salinity 13.7-26.8,. tem-
perature 23.5-30.8. Number and length: May (2, 31-32); June (13,
23-65); Aug. (5, 143-206).

Leiostomus xanthurus Lacépéde. Spot. Stations: A(2, 3, 8, 10-
eles 9s 21), C(1-3, 6-11, 13-15, 21-29, 31, 33-38), D(3), EC1,
6, 10, 12-16, 19, 22-31, 33-37), G(1, 2), H(2-4, 6, 10-12, 14-16, 21,
30), I(1-3, 10-12), K(2). Salinity 0-27.3, temperature 11.0-34.1.
Number and length: Jan. (9, 124-158); Feb. (508, 20-188); Mar.
(2, 186, 17-164); Apr. (1,250, 16-178); May (823, 38-160); June
(504, 40-138); July (255, 57-115); Aug. (60, 68-157); Sept. (15,
96-137); Oct. (26, 97-185); Nov. (15, 105-136); Dec. (2, 129-130).

Menticirrhus americanus (Linnaeus). Southern kingfish*. Sta-
tion: E(35). Salinity 6.2, temperature 24.8. Number and length:
Oct. (4, 28-44).

Micropogon undulatus (Linnaeus). Atlantic croaker. Stations:
A(1-3, 8-12, 14-16, 18), B(2), C(2-14, 16-22, 24-30, 32-35, 39),
DCS) eeki(lers, 6, 9-14) 16, 17, 19-25, 27-29, 31, 33, 37), F(1), G(1-
ay Hi 2-14, 16, 19, 20, 22, 23, 25-38, 40-42), 1( 1-16, 19-21), J(2, 6,
7), K(3-5), L(1). Salinity 0-30.2, temperature 8.3-34.1. Number
and length: Jan. (368, 16-105); Feb. (388, 5-147); Mar. (1,858, 16-
162); Apr. (1,711, 17-155); May (1, 110, 16-182); June (1,499, 24-

42 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

145); July (395, 59-162); Aug. (268, 78-171); Sept. (13, 123-154);
Oct. (92; 27-175): Nov, (220) 11-162): Dee~( 392) 17-1 aiae

Pogonias cromis (Linnaeus). Black drum. Stations: A(16),
C(4, 17, 22, 26, 27). Salinity 0-24.5, temperature 12.0-27.0. Num-
ber and length: Jan. (1, 194); Mar. (2, 181-184); Apr. (1, 194);
May (4, 178-203); Dec. (3, 174-195).

Sciaenops ocellata (Linnaeus). Red drum. Stations: A(20),
C(29, 39), E(25, 27, 28, 37), H(31). Salinity 0-22.7, temperature
16.5-29.2. Number and length: Apr. (6, 106-129); May (3, 116-
164); June (11, 110-160); July (1, 249); Nov. (6, 31-70).

Stellifer lanceolatus (Holbrook). Star drum*. Stations: A(7),
C(36), E(3, 20). Salinity 0-15.8, temperature 11.0-30.8. Number
and length: June (3, 94-95); Sept. (24, 12-150); Dee. (1, 77).

Family SPARIDAE

Archosargus probatocephalus (Walbaum). Sheepshead. Sta-
tions A(8, 12, 15-18, 20-22, 24, 25), C(14), E(6, 11) 13) 165246 2she
H(12, 15, 31). Salinity 0-26.8, temperature 11.0-34.1. Number
and length: Feb. (1, 95); Apr. (8, 75-258); May (7, 26-181); June
(3, 136-190); July (4, 59-166); Aug. (3, 166-194); Sept. (2, 106-
195); Oct. (4, 98-216); Nov. (1, 190).

Lagodon rhomboides (Linnaeus). Pinfish. Stations: A(8, 10,
12-14, 16, 19, 20, 25), B(6), C(8-11, 13, 15, 22, 30-32, 34, 38), D(3),
E(11-18, 22-33, 35, 37), H(15, 17, 20). Salinity 0-25.0, tempera-
ture 11.0-34.1. Number and length: Jan. (1, 94); Feb. (1, 127);
Mar. (14, 102-141); Apr. (14, 21-121); May (27, 22-157); June (32,
44-153); July (117, 43-159); Aug. (22, 73-170); Sept. (11, 104-146);
Oct. (23, 72-203); Nov. (2, 100-109).

Family EPHIPPIDAE

Chaetodipterus faber (Broussonet). Atlantic spadefish*. Sta-
tions: A(3, 21),/E(11, 13, 32, 36). Salinity 0-221) temperature
25.3-34.1. Number and length: May (2, 15-28); June (1, 98); July
(2943-58) Annee: (Ol, 32) Oct ile 24)p

Family TRICHIURIDAE

Trichiurus lepturus Linnaeus. Atlantic cutlassfish*. Stations:
B(3), C(1, 2). Salinity 13.4-23.7, temperature 20.3-28.5. Number
and length: June (3, 330-406); Oct. (1, 134); Nov. (1, 325).

Tacatz: Fishes of St. Johns River 43

Family SCOMBRIDAE

Scomberomorus maculatus (Mitchill). Spanish mackeral*.
Stations: A(18), B(6). Salinity 15.8-22.5, temperature 28.7-30.0.
Number and length: June (1, 108); Sept. (1, 58).

Family ELEOTRIDAE

Dormitator maculatus (Bloch). Fat sleeper. Station: H(18).
Salinity 0, temperature 31.6. Number and length: Aug. (5, 97-126).

Eleotris picta Kner and Steindachner. Spotted sleeper*. F

Family GoBIIDAE

Awaous tajasica (Lichtenstein). River goby. None captured.

Bathygobius soporator (Valenciennes). Frillfin goby. None
captured.

Gobioides broussonneti Lacépéede. Violet goby*. Stations:
B(3, 5), C(22), F(1), G(2). Salinity 0-23.7, temperature 16.8-
32.4. Number and length: Mar. (1, 597); May (2, 185-200); June
fel o02) sept. (3, 381-572).

Gobionellus boleosoma (Jordan and Gilbert). Darter goby.
Stations: A(8, 11, 14, 20-22, 25), B(6), C(15-17, 36, 39), E(17, 20,
21, 26, 27, 29, 34-37), H(20). Salinity 0-20.1, temperature 11.0-
31.0. Number and length: Jan. (1, 32); Feb. (1, 37); Mar. (5, 20-
39); Apr. (5, 32-44); May (3, 42-49); June (1, 43); July (11, 26-
37); Aug. (5, 14-39); Sept. (9, 15-36); Oct. (10, 19-38); Nov. (17,
18-40); Dec. (3, 24-28).

Gobionellus gracillimus Ginsburg. Slim goby*. Station: C(29).
Salinity 22.7, temperature 26.4. Number and length: June (2, 129-
159).

Gobionellus hastatus Girard. Sharptail goby*. Stations: C(17,
30), E(14, 20). Salinity 4.0-12.0, temperature 11.0-30.0. Number
and length: July (1, 218); Sept. (1, 84); Dec. (2, 91-178).

Gobionellus shufeldti (Jordan and Evermann). Freshwater
goby. Stations: A(10), C(10, 13, 17, 21-27, 31, 34, 36, 37), E(1, 2,
1059 22-26, 30, 37), G(1), H(18, 25, 27, 29), 1(7-9, 19). Salinity
0-21.3, temperature 12.0-31.6. Number and length: Jan. (2, 39-91);
Feb. (1, 67); Mar. (151, 28-74); Apr. (60, 28-77); May (11, 31-61);

44 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

July (3, 44-73); Aug. (8, 24-41); Sept. (5, 15-33); Oct. (1, 59);
Nov. (21, 35-92); Dec. (11, 40-66).

Gobionellus smaragdus (Valenciennes). Emerald goby*. Sta-
tion: E(13). Salinity 15.3, temperature 34.1. Number and length:
JulyaGi om):

Gobiosoma bosc (Lacépéde). Naked goby. Stations: A(8, 11,
14, 15), C(3, 16, 24, 28, 31, 35-38), E(12, 22-24 28 30°32) 3437)k
H(7, 9, 10, 20, 29, 33, 36, 40, 42), 1(15). Salinity 0-26.8, tempera-
ture 11.0-31.0._ Number and length: Jan. (91, 15-40); Feb. (7, 20-
37); Mar. (13, 33-50); Apr. (15, 24-42); May (2, 32-38); June (3,
29-34); July (7, 13-41); Aug. (10, 14-26); Sept. (38, 12-32); Oct.
(542, 14-47); Nov. (118, 16-33); Dec. (1, 20).

Gobiosoma robustum Ginsberg. Code goby*. Stations: E(14),
H(25). Salinity 0-11.0, temperature 14.0-30.0. Number and length:
Jan. (4, 20-23); July (1, 26).

Microgobius gulosus (Girard). Clown goby. Stations: C(9,
34, 36-38), E(12, 13, 35-37), F(1), H(4-9, 12, 19) 20, 222 bees
41, 42), I(1, -4, 6, 7, 9-11, 13, 15; 16, 18) 9S 21; eG omar
K(1, 4, 5). Salinity 0-19.9, temperature 12.0-34.1. Number and
length: Jan. (26, 28-52); Feb. (3, 43-49); Mar. (11, 47-62); Apr.
(14, 25-65); May (4, 17-61); June (5, 23-62); July (2, 35-60); Aug.
(18, 17-45); Sept. (13, 22-47); Oct. (85, 11-54); Nov. (92, 14-61):
Dec. (80, 13-56).

Microgobius thalassinus (Jordan and Gilbert). Green goby”*.
Stations: A(15, 20, 25), C(13, 15, 24, 36, 37), E(12, 14 30032) 35-
37). Salinity 0-26.8, temperature 16.5-30.8. Number and length:
Mar. (1, 43); May (2, 46-47); June (4, 23-35); July (12, 24-42);
Aug. (1, 32); Sept. (2, 19-21); Oct. (12, 27-42); Nov. (16, 30-36).

Family TRIGLIDAE

Prionotus scitulus Jordan and Gilbert. Leopard searobin*. Sta-
tion: A(17, 18, 24, 25). Salinity 7.9-24.7, temperature 24.0-28.7.
Number and length: June (9, 43-63); Sept. (7, 22-43); Oct. (9,
26-58 ).

Prionotus tribulus Cuvier. Bighead searobin*. Stations: A(3,
8-11, 12, 15, 19, 20), C( G6, 8, 15, 26, 35, 36, 39), HiGlOme IS za
Salinity 0-26.8, temperature 11.0-29.5. Number and length: Jan. (1,
68); Feb. (5, 61-88); Mar. (8, 19-93); Apr. (2, 28-71); May (2, 38-

Tacatz: Fishes of St. Johns River 45

67); June (1, 32); July (3, 32-36); Sept. (5, 37-72); Oct. (1, 39);
Nov. (2, 46-47).

Family URANOSCOPIDAE

Astroscopus y-graecum (Cuvier). Southern stargazer*. Sta-
tions: A(11), D(3). Salinity 11.4-18.3, temperature 11.7-18.5.
Number and length: Jan. (1, 65); Mar. (2, 45-55).

Family BLENNUDAE

Chasmodes bosquianus (Lacépede). Striped blenny*. Stations:
A(9), E(13). Salinity 10.8-15.3, temperature 13.0-34.1. Number
and length: Feb. (2, 51-63); July (1, 39).

Hypsoblennius hentzi (Lesueur). Feather blenny*. Station:
A(11, 17). Salinity 11.4-24.7, temperature 18.5-26.6. Number and
length: Mar. (3, 56-66); June (1, 65).

Hypsoblennius ionthas (Jordan and Gilbert). Freckled blenny*.
Station: A(17). Salinity 24.7, temperature 26.6. Number and
length: June (1, 36).

Undetermined Hypsoblennius. Station: A(16, 19, 21, 24-26).
Salinity 7.9-23.1, temperature 17.4-29.8. Number and length: May
eeeG2) aialy (ls 53); Aug. (1, 40); Sept. (3, 62-72); Oct. (2, 22-
52); Nov. (2, 61-62).

Family OPHIDIDAE

Ophidion welshi (Nichols and Breder). Crested cusk-eel*.
Station: A(2, 5). Salinity 20.8-27.3, temperature 21.5-25.2. Num-
ber and length: Mar. (9, 69-179); May (3, 128-145).

Family STROMATEIDAE

Peprilus alepidotus (Linnaeus ). Southern harvestfish*. Stations:
B(5), D(3). Salinity 16.0-18.3, temperature 11.7-28.3. Number
and length: Jan. (2, 81-90); Sept. (1, 96).

Family SPHYRAENIDAE

Sphyraena barracuda (Walbaum). Great barracuda*. Station:
A(20). Salinity'14.5, temperature 29.2. Number and length: July
(1, 41).

46 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Family MuciLarE

Agnonostomus monticola (Bancroft). Mountain mullet. None
captured.

Mugil cephalus Linnaeus. Striped mullet. Stations: A(10, 12,
17,18, 23; 25, 26), B(6), G(13, 18, 26) 29) 32°34) B22 a Ane enooe
36), H(17, 20, 29). Salinity 0-25.2, temperature 13.0-31.5. Num-
ber and length: Mar. (23, 24-166); Apr. (31, 22-330). June (69;
32-254); July (14, 76-330); Aug. (8, 122-381); Sept. (2, 131-159);
Oct. (3, 117-164); Nov. (6, 145-155); Dec. (13, 24-229).

Mugil curema Valenciennes. White mullet. Stations: A(19,
21), B(6), C(15, 16, 21, 31, 33°35), EG9) 22.2823 se spore
H(17, 20), 1(17). Salinity 0-22.1, temperature 16.5-31.5. Number
and length: Mar. (22, 22-139); June (56, 45-167); July (83, 48-92):
Aug. (24, 75-169); Sept. (14, 93-130); Oct. (9, 113-125); Nov. (13,
108-170).

Family ATHERINIDAE

Labidesthes sicculus (Cope). Brook silverside. F

Membras martinica (Valenciennes )._ Rough silverside. Station:
A(25). Salinity 7.9, temperature 26.2. Number and length: Oct.
(25; S2eKoy.

Menidia beryllina (Cope). Tidewater silverside. Stations:
A(16-19), C(15, 18, 28, 31, 32, 35-39), E(27, 28 30,3) same
H(3, 4, 8, 18, 20), 1(2, 3, 6, 17), J(2-6, 8), K(3) Ge) easalnmniay
0-24.7, temperature 12.0-31.6. Number and length: Jan. (1, 45);
May (51, 25-42); June (363, 18-53); July (71, 38-57); Aug. (6, 38-
56); Sept. (38, 19-52); Oct. (28, 23-42); Nov. (8, 27-49); Dec:
(13, 21-54).

Menidia menidia (Linnaeus). Atlantic silverside. Stations:
A(10-12, 15, 20-22, 24-26), C(26, 34, 36), E(35, 36))) Ssalinuty,
0-26.8, temperature 17.4-31.5. Number and length: Mar. (100, 65-
96); Apr. (54, 57-96); May (3, 69-79); July (50, 55-65); Aug. (520,
38-59); Sept. (21, 54-63); Oct. (40, 32-71); Nov. (17, 37-72).

Family BoTHIDAE

Ancylopsetta quadrocellata Gill. Ocellated flounder*. Station:
A(6, 8, 11). Salinity 7.7-25.8, temperature 11.0-19.3. Number and
length: Feb. (4, 55-67); Mar. (1, 85); Apr. (3, 34-59).

Tacatz: Fishes of St. Johns River AT

Citharichthys spilopterus Giinther. Bay whiff. Stations: A(15-
pt 20)), BiG), C(2, 7-9, 11-15, 25, 27-29, 33-35, 39), E(3, 11-18,
28, 30, 32, 34-37), H(5, 19, 40), I1(2). Salinity 0-26.8, temperature
16.5-34.1. Number and length: Apr. (3, 38-86); May (45, 26-93);
June, (90, 31-97); July (45, 38-121); Aug. (35, 41-125); Sept. (14,
97-126); Oct. (8, 26-124); Nov. (5, 51-62).

Etropus crossotus Jordan and Gilbert. Fringed flounder*. Sta-
MONG AS. 9, I, 24-26), B(1, 6), E( 14). Salinity 7.7-28.0, tem-
perature 11.0-30.0. Number and length: Feb. (2, 69-72); Mar. (1,
$9); June (3, 35-42); July (1, 78); Sept. (4, 33-88); Oct. (6, 28-
89); Nov. (1, 56).

Paralichthys albigutta Jordan and Gilbert. Gulf flounder. Sta-
mouse ll6,.17), C(6, 16), E(17). Salinity 7.7-24.7, tempera-
ture 11.0-30.8. Number and length: Feb. (5, 127-168); Mar. (2,
plea Nay al 71): june (3, 66-68); Sept. (1, 139); Nov. (1, 159).

Paralichthys dentatus (Linnaeus). Summer flounder*. Sta-
tions: A(15), C(7). Salinity 20.0-26.8, temperature 23.5. Number
and length: Apr. (1, 73); May (2, 65-86).

Paralichthys lethostigma Jordan and Gilbert. Southern flound-
er. Stations: A(9, 21, 22), B(2), C(6, 8, 14, 16, 17, 21, 29, 30, 35,
So pee Oe 12213), GC2), H(3, 20; 29, 38), 1(2). Salinity 0-30.2,
temperature 12.0-31.0. Number and length: Feb. (1, 311); Mar.
(19, 22-56); Apr. (6, 40-78); May (6, 67-155); June (43, 61-145);
Aug. (3, 257-503); Sept. (6, 106-201); Oct. (1, 139); Nov. (3, 112
202); Dec. (6, 129-144).

Undetermined Paralichthys. Stations: A(1, 3, 10-12, 15, 18-20),
C(10-13, 21-28, 30-32, 36-38), E( 13-15, 22-25, 27, 32, 34), H(4, 19,
230, 32. 33, 37, 41), 1(4, 12, 13). Salinity 0-26.8, temperature
14.0-34.1. Number and length: Jan. (1, 210); Mar. (126, 15-254);
Apr. (104, 32-178); May (43, 25-362); June (9, 71-240); July (27,
54-231); Aug. (13, 89-356); Sept. (3, 106-191); Oct. (4, 110-164).

Scophthalmus aquosus (Mitchill). Windowpane*. Station:
E(19). Salinity 5.5, temperature 19.4. Number and length: Nov.
(le 122.)

Family SOLEIDAE

Achirus lineatus (Linnaeus). Lined sole*. Stations: A(15),
C(15, 21), E(20). Salinity 0-26.8, temperature 11.0-23.5. Number
piemencth: Mar. (1, 96); May (1, 31); Oct. (1, 77); Dec. (1, 92).

A8 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Trinectes maculatus (Bloch and Schneider). Hogchoker. Sta-
tions: A(le5, 8) ll), €(9; 10, 25, 34 36, 37), Dis), Ga aiigaseE
22, 35-37), F(1), G(2, 3), H(1, 2, 414, 16, 18-22, 25-42), I( 1-16,
18-21), J(1-8), K(1-5), L(1-3). Salinity 0-28.2, temperature 8.8-
34.1. Number and length: Jan. (116, 26-108); Feb. (90, 27-128);
Mar. (785, 16-172); Apr. (230, 25-112); May (170, 36-114); June
(220, 15-122); July (140, 16-118); Aug. (189, 17-123); Sept. (70,
15-103); Oct. (366, 17-140); Nov. (240, 28-135); Dec. (231, 25-121).

Family CyNOGLOSSIDAE

Symphurus plagiusa (Linnaeus). Blackcheek tonguefish. Sta-
tions: A(1, 8-12, 15-22, 24-26), B(6), C(2, 9-1), 14, 16 ie 225:
27, 29-39), D(3), E(10, 13-15, 22, 24, 25, 28, 30-32, 34-375) saline
ity 0-26.8, temperature 11.0-34.1. Number and length: Jan. (8, 32-
81); Feb. (21, 44-106); Mar. (50, 31-88); Apr. (7, 32-115); May
(4, 25-77); June (20, 19-81); July (47, 19-68); Aug. (35, 11-76);
Sept. (55, 18-92); Oct. (91, 18-84); Nov. (28, 22-152); Dec. (3, 33-
37).

Family GoBrEsOCcIDAE

Gobiesox strumosus Cope. Skilletfish*. Stations: A(8), C(21),
E(19). Salinity 0-7.7, temperature 11.0-19.4. Number and length:
Feb. (1, 40); Mar. (1, 40); Nov. (1, 38).

Family BALISTIDAE

Stephanolepis hispidus (Linnaeus). Planehead filefish*. Sta-
tions: A(15-18), C(27, 29), E(3, 11-14, 26-29). Salinity 11.0-26.8,
temperature 22.8-34.1. Number and length: May (12, 17-60); June
(14, 21-59); July (3, 65-70).

Family TETRAODONTIDAE

Sphaeroides maculatus (Bloch and Schneider). Northern puf-
fer*. Stations: A(16-19), C(10, 26, 29), E(12, 13, 28, 32, 33).
Salinity 6.7-24.7, temperature 23.0-34.1. Number and length: Apr.
(1, 16); May (1, 19); June (25, 18-56); July (4) 42-64) -eAueer Ge
40).

Tacatz: Fishes of St. Johns River 49

Family D1IopONTIDAE

Chilomycterus schoepfi (Walbaum). Striped burrfish*. Sta-
tions: C(15), E(3, 4). Salinity 11.1-14.3, temperature 21.0-30.8.
Number and length: June (1, 23); Aug. (1, 101); Oct. (1, 39).

Family BATRACHOIDIDAE

Opsanus tau (Linnaeus). Oyster toadfish. Stations: A(1, 11,
Eielo- 2h 24-26) B(6), C19), E(6, 18, 24, 30, 32, 33). Salin-
ity 0-24.7, temperature 14.0-30.0._ Number and length: Jan. (1, 51);
Mar (3, 133-233); Apr. (4, 130-161); June (2, 118-172); July (22,
18-289); Aug. (3, 26-163); Sept. (2, 135-229); Oct. (4, 89-205);
Nov. (7, 88-205).

Family OGCOCEPHALIDAE

Ogcocephalus cubifrons (Richardson). Shortnose batfish. None
captured.

ACKNOWLEDGMENTS

Dr. Carter R. Gilbert, Assistant Curator in charge of fishes, the
Florida State Museum, Gainesville, identified representative speci-
mens from almost all of the collections, read the manuscript, and
offered helpful suggestions on treatment of the data. Dr. William
M. McLane, Florida Aquatic Nurseries, Ltd., Ft. Lauderdale, gen-
erously allowed me to include the records from his unpublished
dissertation on the fishes of the St. Johns River. Martin A. Roessler,
University of Miami, identified the fine-scaled species of Gobionel-
lus. |

LITERATURE CITED

BaiLEy, REEVE M., ERNEsT A. LACHNER, C. C. LinpsEy, C. RicHARD ROBINS,
Pum. M. Roepe, W. B. Scorr, Loren P. Woops. 1960. A list of
common and scientific names of fishes from the United States and Can-
ada. 2nd Ed., Amer. Fish. Soc., Spec. Pub. 2, 102 pp.

Breck, WILLIAM M., Jr. 1965. The streams of Florida. Bull. Florida State
Mus., vol. 10, no. 3, pp. 91-120.

Briccs, Joun C. 1958. A list of Florida fishes and their distribution. Bull.
Florida State Mus., Biol. Sci., vol. 2, no. 8, pp. 223-318.

50 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Carr, ARCHIE, AND COLEMAN J. Gorn. 1959. Guide to the reptiles, amphibi-
ans, and fresh-water fishes of Florida. Univ. Florida Press, Gainesville,
341 pp.

EVERMANN, BARTON W., AND WILLIAM C., KENDALL. 1900. Check-list of the
fishes of Florida. U.S. Comm. Fish and Fish., pt. 25, Rep. Comm.
1899, pp. 5-103.

Fow.er, Henry W. 1945. A study of the fishes of the southern Piedmont
and Coastal Plain. Monogr., Acad. Nat. Sci. Philadelphia, vol. 7, pp.
1-408.

GuNTER, Gorpon. 1945. Studies on marine fishes of Texas. Pub. Inst. Mar.
Sci., Univ. Texas, vol. 1, no. 1, pp. 1-190.

McLane, WitLiAM M. 1955. The fishes of the St. Johns River System. Un-
published Ph.D. Dissertation, Univ. Florida, 362 pp.

Moopy, Harotp L. 1961. Exploited fish populations of the St. Johns River,
Florida. Quart. Jour. Florida Acad. Sci., vol. 24, no. 1, pp. 1-18.

Myers, Georce S. 1938. Fresh-water fishes and West Indian zodgeography.
Rep. Smithsonian Inst. 1937, pp. 339-364.

Myers, Georce S. 1951. Fresh-water fishes and East Indian zodgeography
Stanford Ichthyol. Bull., vol. 4, no. 1, pp. 11-21.

Opum, H. T. 1953. Factors controlling marine invasion into Florida fresh
waters. Bull. Mar. Sci. Gulf Caribbean, vol. 3, no. 1, pp. 134-156.

Bureau of Commercial Fisheries, Biological Laboratory, Beau-
fort, N. C. 28516

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Composition and Feed Value of Shrimp Meal
W. G. Kirk, R. L. Suimuey, J. F. EASLEY, AND F’. M. PEAcock

SHRIMP meal, a by-product of the shrimp industry, is assuming
more importance as the yearly catch of these crustaceans increases
and as ways are found to utilize the inedible portions. On ship
processing is begun with the removal and discarding of the head,
followed by freezing. When delivered to the processing plant, the
shrimp are thawed, and the shell, legs, and vein removed. This
material is then steamed, dried, bagged, and sold under the name
of Shrimp Meal or Shrimp Gel. It has been found to have com-
mercial value as animal feed and fertilizer.

This report gives data on the chemical composition of shrimp
meal and its use as a protein supplement in cattle fattening rations.

MATERIALS AND METHODS

Shrimp meal consists mainly of small irregular flakes with many
finer particles. Some flakes are pale in color and reflect light while
others are of various shades of grey and brown. The meal has a
distinct shrimp or fish odor. Samples collected for analysis con-
sisted of small aliquots of meal from several bags fed in each of the
four trials. Dry matter, crude protein, ether extract, crude fiber,
ash, phosphorus, urea, uric acid, and alkaline distillable ammonia
were determined by A.O.A.C. methods (1960); calcium by pro-
cedure outlined by Welcher (1957); nitrate and nitrite by methods
of Nelson et al. (1954); and acetylglucosamine (chitin) determined
according to Boas (1953).

CHEMICAL COMPOSITION

The main constituents of shrimp meal as shown by the proximate
analysis (Table 1) are crude protein, crude fiber, and ash with a
limited amount of ether extract and a negative NFE (nitrogen-free-
extract) value. The A.O.A.C. method of analysis using copper sul-
fate as a catalyst gave an average of 45.3 per cent crude protein.
The negative NFE value suggests that an “unknown” factor had
entered into the calculation of the composition of shrimp meal.
When the samples were analyzed for acetylglucosamine an average

52 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

value of 27.2 per cent was obtained. When the nitrogen of this
compound was subtracted from the total nitrogen and the difference
multiplied by 6.25, an average value of 34.4 per cent crude protein,
not acetylglucosamine, was found. Essentially all the crude fiber
residue by the A.O.A.C. method was accounted for as glucosamine.
This indicated that the acetate group of the acetylglucosamine was
removed by the acid and alkali extractions of the crude fiber deter-
mination. The polymeric acetylglucosamine (chitin) is similar in
structure to the polyglucose structure of cellulose and thereby
might be expected to be in a proximate analysis crude fiber fraction.

TABLE 1

Composition of shrimp meal by the usual proximate analysis procedure

Nitrogen
Dry Crude Crude Ether free
Sample Matter Protein! Fiber? Extract Ash Extract
1 89.0 42.3 Zon 0.42 27.8 — 6.623
2 91.5 48.1 20.0 1.87 2m =) '57
3 92.9 46.0 230 0.92 ADA) On
4 O2N7 44.7 PASS} OM. 25.7 = se
Average 91.5 45.3 O88) 0.98 252, — oo

1 Per cent total nitrogen in samples times 6.25.

2 Analysis indicated the crude fiber was glucosamine, which is chitin without
the acetyl group.

3 Nitrogen-free extract values are obtained by subtracting the sum of percentage
values for moisture, crude protein, crude fiber, ether extract, and ash from 100.
The negative values obtained this way for NFE suggests that a spurious value
for crude protein had been calculated when the percentage N in the samples
was multiplied by 6.25 to obtain the per cent crude protein.

The results of further analyses of shrimp meal show the high
fiber content (Table 1) to be due to the nitrogenous substance glu-
cosamine which results in spurious values for crude protein and
crude fiber. More appropriate composition values are given in
Table 2. When the shrimp meal was analyzed for nitrates, pyri-
dine-ring type nitrogen, urea, uric acid, and ammonia salts, only a
trace was found. These data indicate that the crude protein fraction
is essentially all amino-nitrogen or true protein except that found
in the acetylglucosamine (chitin). The average percent of ash was
25.2 per cent with calcium 10.7 per cent and phosphorus 2.7 per
cent making 53.2 per cent of the total.

Kirk ET AL: Value of Shrimp Meal 53

According to the Merck Index (1960) chitin is a white amor-
phorus material forming the harder part of the horny substance of
crabs, beetles, and shrimp and is found in fungi and bacteria. It
is insoluble in the ordinary solvents, dilute acid and alkali used in
crude fiber determinations. It contains 7.2 per cent nitrogen.

TABLE 2

Corrected composition of shrimp meal accounting for chitin composition

Crude Acetylglu- Nitrogen

Dry Protein cosamine! Ether free

Sample Matter (not chitin) (chitin) Extract Ash Extract
i 89.0 S216 24.3 0.4 27.8 3.9
2 91.5 38.1 24.9 1.9 2 ali 4.5
3 92.9 34.0 29.9 0.9 WB 2.4
4 92.7 32.8 29.8 0.7 D5 3.7
Average 91.5 34.4 Dl 1.0 25:3 3.6

1Contains 1.744% nitrogen equivalent to 10.9% crude protein.

CATTLE FEEDING TRIALS

In a preliminary feeding trial of 102 days grade steers fed a
balanced ration of cottonseed hulls, a mixture of shrimp meal and
cottonseed meal, citrus pulp and alfalfa meal had an average daily
gain of 1.93 pounds. The steers on the average ate 1.96 pounds
shrimp meal daily.

In the second trial of 129 days cattle fed a balanced ration con-
taining cottonseed meal had an average daily gain of 1.90 pounds,
consuming 984 pounds feed per 100 pounds gain. Animals fed
shrimp meal had an average daily gain of 2.19 pounds, eating 921
pounds feed for each 100 pounds gain. Improvement in slaughter
grade was from U. S. Utility to U. S. Good. Steers fed shrimp meal
made the fastest and most efficient use of their feed for gains but
this advantage was offset by a 2.4 per cent lower carcass yield com-
pared to those fed cottonseed meal.

Eighteen grade steers in both Trials 3 and 4 were divided into
three lots of six steers each and fed in drylot for 124 and 125 days,
respectively. The three balanced rations of hay and cottonseed
hulls, citrus pulp, alfalfa meal and corn meal contained the same
weight of protein supplement: Lot 1, cottonseed meal; Lot 2, %4 as

54 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 3

Average gain, feed consumption, carcass grade, and yield for steers fed in

Lot No.
Protein supplement

No. steers
Age, months
Weight, lb.:
Initial
Gain
Daily gain
Daily protein supplement:
Cottonseed meal, lb.
Shrimp meal, lb.
Daily ration, lb.
TDN daily, lb.
TDN/100 pounds gain, lb.
Carcass grade
Carcass yield

Trial 3

1
Cotton-
seed
meal

6
17

612
301
2.43

3.02
24,32
16.04

660
High Good
60.91

TABLE 4

2
%4 cotton-
seed meal,
Y% shrimp
meal

6
7

596
283
2.28

2.14
.70
PAS)
14.73
646
High Good
60.22

3
Shrimp
meal

Average gain, feed consumption, carcass grade, and yield for steers fed in

Lot No.
Protein supplement

No. steers
Age, months
Weights, lb.:
Initial
Gain
Daily gain
Daily protein supplement:
Cottonseed meal, lb.
Shrimp meal, lb.
Daily ration, lb.
TDN daily, Ib.
TDN/100 pounds gain, lb.
Carcass grade

Trial 4

1
Cotton-
seed
meal

627
High Good
61.15

2
34 cotton-
seed meal,
Yq shrimp
meal

6
22

805
323
2.58

2.28
505
27.20
16.49
639
High Good
60.48

3.03
PA sil
ape
680
High Good
61.30

Carcass yield

Kirk ET AL: Value of Shrimp Meal 55

much cottonseed meal as Lot 1, and “4 shrimp meal; Lot 3, shrimp
meal. The results of these two trials are summarized in Tables 3
and 4.

DISCUSSION

Rations with shrimp meal as a protein supplement were eaten
readily and cattle remained in a healthy state. According to Morri-
son (1956) cottonseed meal, 41 per cent protein, has a 69 per cent
TDN (total digestible nutrients) and shrimp meal 43.5 per cent
TDN. This is a difference of 0.76 pounds more TDN per animal
when the protein supplement fed is 3 pounds daily in favor of cattle
fed cottonseed meal compared to animals fed shrimp meal. Shrimp
meal has approximately 19 per cent more mineral ash than cotton-
seed meal which probably accounts for much of the difference in
TDN in the two protein feeds. This may be a reason for the higher
average daily gain by steers fed cottonseed meal of 0.12 in Trial 3
and 0.32 pound in Trial 4.

The roughage feeds, hay and cottonseed hulls, fed in Trial 3
made up 15-17 per cent of the feed eaten. In Trial 4, hay, cotton-
seed hulls, shuck, and cob in the ground snapped corn, supplied
30-31 per cent of the total ration. Lot 1 in Trial 3 had 0.20 pound
higher average daily gain than in Trial 4, while Lot 3 fed shrimp
meal had the same rate of gain in both trials.

The steers fed in Trials 3 and 4 were of similar breeding but
those in Trial 4 averaged 5 months older and 194 pounds heavier
than the steers in Trial 3. Trial 4 animals had increased carcass
yield. This might be attributed to their greater age and weight or
to the higher percentage of roughage in their diet.

ACKNOWLEDGMENTS

The authors are indebted to Treasure Isle Ocean Products, Inc.,
Tampa, Florida, who furnished the shrimp meal fed in the four
cattle fattening trials. The chemical study was supported in part
by a grant-in-aid from the Moorman Manufacturing Company,
Quincy, Illinois.

LITERATURE CITED

ASSOCIATION OFFICIAL AGRICULTURAL CHEMISTS. 1960. Official Methods of
Analysis. Washington, D. C., 9th edition, 832 pp.

56 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Boas, Norman F. 1953. Methods for the determination of hexosamines in
tissues. Journ. Biol. Chem., vol. 204, pp. 553-563.

Morrison, F. B. 1956. Feeds and feeding. Morrison Publ. Co., Ithaca, New
York, 22nd edition.

NELSON, J. L., L. T. Kurtz, ANp R. H. Bray. 1954. Rapid determination of
Nitrates and Nitrites. Anal. Chem., vol. 26, pp. 1081-1082.

Merck INDEX OF CHEMICALS AND Drucs. 1960. Merck & Co., Inc., Rahway,
Neale

WELCHER, FRANK J. 1957. The analytical uses of ethyenediamine-tetraacetic
acid. D. Van Nostrand Company, Inc., Princeton, N. J.

Range Cattle Experiment Station, Ona, Florida, and Animal
Science Department, University of Florida, Gainesville, Florida.
Florida Agricultural Experiment Stations Journal Series No. 2353.

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Animal Remains from a Midden at Fort Walton Beach
ELIZABETH S. WING

A SMALL sample of animal bones was excavated by the late
William Lazarus from a Deptford midden site which dates from
300 B.C. and is located beneath the Fort Walton Temple Mound
in Fort Walton Beach, Okaloosa County, Florida. This sample is
of interest in that knowledge of the Deptford period economy is
scant. It represents all the bone within a portion of a level
(pit 5, level 11) 3 feet square and 0.7 feet deep and is composed
primarily of the remains of small fish.

For the various species minimum numbers of individuals are
given below and are the same as the largest number of identical
elements of skeleton present. The number of bones or bone
fragments of each species is not given, because fish vertebrae, com-
prising the bulk of the material, are difficult to identify. Further-
more, some species are represented by only one preservable ele-
ment, as teeth of shark and ear bones of shad, which would Eve
a bias to the bone count.

With few exceptions the remains are of small individuals,
comprising 20 species of fishes and six other vertebrates. Jack
(Caranx), catfish (Galeichthys felis), and shad (Brevoortia ctf.
B. smithi) are by far the most abundant fishes at this site repre-
sented by at least 132, 25, and 21 individuals, respectively. The
species of jack that were identified are blue runner (Caranx cry-
sos) and crevalle jack (C. hippos). Sheephead (Archosargus sp.),
mullet (Mugil sp.), spotted sea trout (Cynoscion nebulosus ), and
flounder (Bothidae) are represented by at least 12, 6, 5, and 4
individuals, respectively. The following fishes are represented by
at least one individual: bull shark (Carcharhinus cf. C. leucas),
bowfin (Amia calva), lady fish (Elops saurus), gafftopsail catfish
(Bagre marinus), snook (Centropomus sp.), kingfish (Menticir-
rhus sp.), drum (Pogonias cromis), pinfish (Lagodon sp.), barra-
cuda (Sphyraena sp.), flounder (Paralichthys sp.), and toadfish
(Opsanus sp.). A few fragments of box turtle (Terrapene caro-
lina), terrapin (Pseudemys sp.), and gopher tortoise (Gopherus
polyphemus ) were found in other parts of the Deptford midden
and could be considered occasional food animals in the economy
of this site. Finally two mammals, rabbit (Sylvilagus sp.) and

58 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

white-tailed deer (Odocoileus virginianus), and one bird, a loon
(Gavia immer), are each represented by at least one individual.
Although the fish probably did not provide much more food than
the one deer represented, the Indians must have occupied much of
their time in fishing.

On the basis of the fishes that were caught, it is possible to
suggest some of the fishing techniques that may have been used.
All the fishes represented are found in shallow water along shore
and around rocks, so that boats would not have been essential
for catching the fish that are represented at the site. Most of the
fish that are represented in the midden are either carnivores or
omnivores and could have been caught with hook and line or nets.
The two vegetarian species, shad and mullet, are more easily
caught with nets. In fact shad, a plankton feeder, cannot be
caught with hook and line. This species requires saline conditions
and would be more abundant in the Gulf than in the Bay. Great
numbers are now caught in nets along the northern part of the
Gulf Coast, and nets may have been used in aboriginal times.
Mullet, one of the most important commercial fishes of the Gulf
Coast today, are more adapted to brackish situations than shad.
The Indians probably caught the mullet in nets or impoundments
in the Bay. Although undoubtedly abundant in the Gulf Coast
waters, this species is represented in Florida coastal Indian sites
only by vertebrae of a few individuals. Perhaps mullet were
caught incidentally to other fish and were prepared differently.

ACKNOWLEDGMENTS

I gratefully acknowledge the support of this work by grant
GS-284 and GS-1018 of the National Science Foundation.

Florida State Museum, University of Florida, Gainesville, Flor-
ida, 32601.

Quart. Jour. Florida Acad. Sci. 30(1) 1966 (1968)

Fossil Vertebrates from Navassa Island, W. I.

THOMAS H. PATTON

In October, 1965, members of the staff of the Florida State
Museum were given permission by the United States Coast Guard
to visit the island of Navassa, W. I., as guests aboard the USCG
Cutter Hollyhock. The major purpose of the trip was to prospect
for possible fossil vertebrate accumulations on the island. This
investigation is part of a continuing study of the systematics and
zoogeography of West Indian fossil vertebrates. Because the
island of Navassa was previously unknown insofar as fossil verte-
brates are concerned, a preliminary report on our findings may be
of interest to zoogeographers and paleontologists.

Navassa Island is a flat-topped limestone block situated 34
miles west of Cap de Irois on the southeast tip of Haiti at approxi-
mately 75° longitude and 18° 25’ N latitude. The island is about
2.2 miles long and 1.2 miles wide and narrows towards the north-
west. Its most conspicuous features are an ancient wave-cut
terrace and the high bluff into which the terrace has been cut.
The surface of the terrace lies about 50 feet above sea level and
is bordered seaward by a steep scarp formed by waves at present
sea level.

The virtually flat upper surface of the island, also a wave-cut
feature, reaches a height of approximately 250 feet towards the
southeast and is tilted gently towards the northwest, where it is
about 140 feet above sea level. The upper surface is relatively
featureless, except for the presence of innumerable limestone
solution pits and cavities. It is within these cavities that the
fossiliferous matrix occurs. The island is honeycombed with such
solution cavities, which are primarily of a vertical position. Within
most of them a cemented, phosphatic, sometimes fossiliferous,
rock is found adhering to the walls of the parent limestone. Ap-
parently the phosphatic rock completely filled the majority of the
cavities at one time, but has been mined to the extent that in most
places little of the original material is left. Where the matrix occurs
in too small amounts to have made mining practical, it is still
undisturbed, but such situations were seldom encountered by our
party.

In addition to the hard phosphatic deposits already mentioned,

60 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

soft sediments were observed in the bottoms of some of the deep
vertical-walled, solution pits that occur commonly in the southeast
part of the island. We were unable to climb down into these
pits on our first visit.

The phosphatic matrix has been treated with acetic acid and
so far has proved to be rather sparsely fossiliferous. Several
reptiles were recovered, however, from a deposit approximately
150 yards from a lighthouse at bearing 20° NE. Three forms are
previously unreported from Navassa (Schmidt, 1921; Barbour,
1937 ).

A preliminary list of fossil vertebrates of probable Pleistocene
age from Navassa is given below, with the additions to the fauna
indicated by asterisks.

Class REPTILIA
Order CHELONIA:
Family Testudinidae: Geochelone sp.*
Family Emydidae: Pseudemys sp.*
Order SQuaAMATA: Suborder Lacertilia
Family Gekkonidae: ?Aristelliger sp.

Family Iguanidae: Cyclura sp.
Anolis sp.

Order SQuAMATA: Suborder Serpentes

Family Colubridae: Alsophis sp.*
LITERATURE CITED

Barsour, THoMas. 1937. Third list of Antillean reptiles and amphibians.
Bull. Mus. Comp. Zool., vol. 82, no. 2, pp. 72-166.

Scumipt, K. P. 1921. The herpetology of Navassa Island. Bull. Amer.
Mus. Nat. Hist., vol. 44, art. 18, pp. 555-559.

Florida State Museum, University of Florida, Gainesville, Flor-
ida 32601.

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Notes on the Trematode Genera Cleidodiscus and Urocleidus

C. E. Price

THE monogenetic trematode genus Cleidodiscus Mueller, 1934,
is one of approximately 50 genera included in the subfamily
Ancyrocephalinae. Descriptive work within this subfamily began
for North America when Van Cleave and Mueller (1932) de-
scribed Cleidodiscus aculeatus from the gills of Stizostedion vitre-
um.

The Ancyrocephalinae have long been in a state of taxonomic
disorder. One of the major reasons for this unsettled condition
has been the tendency of some workers to base new genera and
species on anatomical features of insufficient magnitude.

Between 1934 and 1938, J. F. Mueller proposed nine genera of
Ancyrocephalinae (then called Tetraonchinae), namely Actinoclei-
dus, Aristocleidus, Cleidodiscus, Haplocleidus, Leptocleidus, On-
chocleidus, Pterocleidus, Tetracleidus, and Urocleidus. Mizelle
and Hughes (1938) reduced these nine genera to three, Actino-
cleidus, Cleidodiscus, and Urocleidus. Actinocleidus, with articu-
lated haptoral bars and cirri with basally articulated accessory
pieces, appears to be a valid genus. Cleidodiscus and Urocleidus,
however, are not so well differentiated, and require a close com-
parison.

Generic diagnoses are quoted below from Mizelle et al. (1956).

Cleidodiscus Mueller, 1934

Synonyms: Leptocleidus Mueller, 1936, in part; Tetracleidus Mueller,
NOSOseim apart

Diagnosis: Tetraonchinae (Ancyrocephalinae) somewhat flattened dorso-
ventrally and with trunk narrowly elliptical in outline (dorsal view). Eyes
four, one pair larger than, and posterior to, the other. Gut bifurcate,
without diverticula; rami confluent posteriorly. Gonads near middle of body.
Cirrus usually a simple cuticularized tube. Accessory piece always present
and generally articulated with the cirrus. Vagina usually present and
opening on left margin near mid-length of trunk (the vagina is dextral in
C. banghami; see Mizelle 1940). Vitellaria consisting or numerous, small,
discrete follicles arranged in a pair of lateral bands extending from the
pharynegeal region to, or into, the peduncle; the bands always posteriorly,
and sometimes anteriorly, confluent. Haptor generally distinct, discoidal or
subhexagonal; armed with two pairs of anchors and seven pairs of hooks.

62 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Anchors, one pair dorsal and the other pair ventral, with superficial roots
of each pair connected by a transverse bar; bars nonarticulate with each other.
Parasitic on gills of freshwater fishes.

Urocleidus Mueller, 1934. Emended by Mizelle and Hughes, 1938

Synonyms; Onchocleidus Mueller, 1936, in part; Tetracleidus Mueller,
1936, in part; Aristocleidus Mueller, 1936, in part; Haplocleidus Mueller,
1937, in part; and Pterocleidus Mueller, 1937, in part.

Diagnosis: Tetraonchinae (Ancyrocephalinae) with trunk, eyes, gut, go-
nads, vitellaria, and haptoral bars as described for Cleidodiscus. Cirrus a
cuticularized tube, straight or undulate, with or without a cirral fin or
thread, infrequently corkscrew-like. Accessory piece never basally articu-
lated with the cirrus. Vagina, when present, opening on the right margin
near mid-length of trunk. Haptor generally distinct, subhexagonal; armed
with two pairs of anchors and seven pairs of hooks. One pair of anchors
dorsal, the other pair ventral, in position. Parasitic on gills of freshwater

fishes.

As an additional character, Yamaguti (1963) states that the
haptoral bars are similar in Urocleidus and dissimilar in Cleidodis-
Cus.

Comparison of the characters of the two genera indicates that
the morphology of the trunk, eyespots, gut, gonads, vitellaria,
and haptoral bars is quite or nearly identical to that of Cleidodis-
Cus.

It is readily ascertained that the haptor armaments of the two
genera are very much alike in all respects. Each haptor contains
two pairs of anchors, the bases of each pair supported by a trans-
verse bar. The anchors are not articulated to each other in either
case, and the anchors are located in the same portions of the
haptors of both genera. All members of both genera possess 14
haptoral hooks, these hooks in each case being arranged in an
essentially common spatial organization. Yamagutis (1963) pro-
posed differentiating factor of dissimilar bars in Cleidodiscus
and similar ones in Urocleidus is not a valid one. Several species
of Cleidodiscus, including the new species described later in this
paper, possess bars very similar in shape. It thus appears that no
significant differences in the haptoral armaments of these genera
exist.

The presence or absence of a vagina can scarcely be considered
a trait of generic magnitude. Mizelle and Hughes (1938) state

Price: Notes on Trematodes 63

that the absence of a vagina in North American fresh-water forms
is not of generic significance since this condition sporadically oc-
curs in several of the old genera.

A survey of North American monopisthocotylean Monogenea
indicates without any serious question that position of the vagina
is a poor generic trait. The vagina of Cleidodiscus banghami,
for example, is dextral (Mizelle, 1940), whereas the vaginae are
sinistral in all other congeneric species. In the genus Dactylogy-
rus the vagina has been reported as on the right ventral margin
in D. amblops Mueller (1938), as being located near the mid-
ventral surface in D. columbiensis Monaco and Mizelle (1955),
and as being sinistral and lateroventral in D. vancleavei Monaco
and Mizelle (op. cit.).

In final analysis, only one character appears to differentiate
Cleidodiscus and Urocleidus. The accessory piece is said to be
basally articulated to the cirrus in Cleidodiscus, whereas the cir-
rus is not joined to the accessory piece in Urocleidus. The articu-
lated accessory piece is apparently derived from ancestral forms
possessing the non-articulated accessory piece. As it seems plaus-
ible that this character might furnish significant information of an
evolutionary nature, both genera should stand, but with revised
limits.

As concerns the species lacking an accessory piece entirely, it
has not been established beyond question that any North American
parasite related to either Cleidodiscus or Urocleidus actually lacks
the accessory piece. If any such species do in fact exist, Oncho-
cleidus Mueller, 1936, is available for their inclusion. Onchocleidus
was established by Mueller to include Urocleidus-like forms which
lacked an accessory piece, but Mizelle (1955) stated that an
accessory piece has been observed in practically every species of
Onchocleidus and probably exists in all of them.

As several species presently classified in Cleidodiscus possess
accessory pieces which are apparently not actually articulated to
the cirri, these forms should be transferred to Urocleidus. They
will be known as U. alatus (Mueller, 1938), U. brachus (Mueller,
1938), U. chautauquansis (Mueller, 1938), U. chavarriai (E. Price,
1938), U. megalonchus (Mueller, 1936), and U. travassoi (E. Price,
1938 ).

64 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

For any worker of the opinion that the trait of an articulated
or nonarticulated accessory piece does not exhibit enough impor-
tance to justify generic separation, the only alternative would be
to synonymize Cleidodiscus and Urocleidus.

A New Species oF Cleidodiscus

The author wishes to express his sincere thanks and apprecia-
tion to Mr. Emory Milner for a single host specimen of Lepomis
auritus (Linnaeus) from his branch near Hollanville, Georgia.
The branchial material and resultant parasites were treated as
prescribed by C. Price and Mizelle (1964). Measurements were
made as outlined by Mizelle and Klucka (1953) and are expressed
in microns.

Cleidodiscus georgiensis sp. n.

Host and locality. Lepomis auritus (Linnaeus), the red-bellied
sunfish; Milner’s Branch, 3 miles SE of Hollanville, Georgia.

Number of specimens studied. Two.
Region of host's body inhabited by parasites. Gill filaments.

Types. Holotype deposited in the helminthological collection
of the U. S. National Museum, accession no. 61201. Paratype in
author's collection.

Description. Dactylogyridae, Ancyrocephalinae. A form of
relatively small size provided with a thin, smooth cuticle devoid
of scales or spines. Body constricted somewhat near mid-length.
Body length 340 (322-358); greatest body width 63 (58-68),
just anterior to constriction. Two pairs of eyespots, members of
posterior pair much larger and somewhat closer together than other
members. Comprising eyespot granules exhibit a slight tendency
to dissociate; a few comprising granules scattered in the cephalic
region. Pharynx subspherical in dorsal view; transverse diameter
16 (14-18). Peduncle short and narrow, with result that haptor
is well set off from body proper. Haptor essentially subspherical
in outline, with a somewhat irregular posterior border; length 83
(75-91), width 80 (76-84).

Two pairs of anchors, all members similar in both size and

Price: Notes on Trematodes 65

shape (Figs. 1-2). Each anchor composed of: (1) a solid base
provided with a prominent superficial and a short deep root, (2)
a solid shaft, joined to (3) a solid point. Shaft and point connect
at a definite angle. Length of ventral anchor 37 (35-39), width of
base 13 (12-14), length of dorsal anchor 36 (34-38), width of
base 14 (13-15). Each of the haptoral bars simple and quite
similar (Figs. 3-4). Length of ventral bar 20 (18-21), length of
dorsal bar 23 (21-24).

: 3 5
Are

= if
i

8
| 0.05 mm

Figs. 1-8. Cleidodiscus georgiensis sp. n., illustrating the major scleroti-
zed structures. 1. Ventral anchor. 2. Dorsal anchor. 3. Ventral bar. 4.
Dorsal bar. 5, 6. Haptoral hooks. 7. Cirrus. 8. Accessory piece.

Haptoral hooks 14 (seven pairs), similar in shape, subequal in
size, and arranged in agreement with Mizelle and Crane (1964).
(Figs. 5-6). Each hook composed of a solid base, a solid shaft, and
a sickle-shaped termination provided with an opposable piece.
Hook lengths: no. 1— 28; nos. 2, 4— 22; no. 3— 23; nos. 6, 7— 24;
no. 5— 18.

Copulatory complex composed of a cirrus and an accessory
piece (Figs. 7-8). Cirrus arises from an inflated base as a tube
of narrow diameter, and opening ventrally; length 25 (23-28).
Accessory piece basally articulated to cirrus; as the accessory piece
shaft distally approaches the cirrus tube, it divides into two short
rami of approximately equal length; length of accessory piece 18
(16-20). Vagina not observed with certainty. Testis postovarian.
Vitellaria moderately well developed, forming two lateral bands.
Intestinal crura confluent posteriorly.

It is difficult to determine which North American Cleidodiscus

66 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

species most clearly resembles the present form. Several species
of this genus possess haptoral armament similar to this new para-
site, but none as yet described possesses a copulatory complex
closely resembling that of C. georgiensis.

SUMMARY

The monogenetic trematode genera Cleidodiscus and Uroclei-
dus are very nearly identical in morphology. The sole differ-
entiating feature is the possession by Cleidodiscus of an accessory
piece basally articulated to the cirrus; in Urocleidus the accessory
piece is not joined to the cirrus base. Several species of Clei-
dodiscus are transferred to Urocleidus on that basis.

A new species, Cleidodiscus georgiensis, recovered from the
gills of the red-bellied sunfish, Lepomis auritus (Linnaeus), is
described.

LITERATURE CITED

MizELLeE, J. D. 1940. Studies on monogenetic trematodes. III. Redescrip-
tions and variations in known species. Jour. Parasitol., vol. 26, no.

Opp ps 6S A/S:

——. 1955. Studies on monogentic trematodes. XIX. The status of
North American Dactylogyrinae and Tetraonchinae. Proc. Indiana

Acad. Sci., vol. 64, pp. 260-264.

MizELLE, J. D., AND J. W. Crane. 1964. Studies on monogentic trema-
todes. XXIII. Gill parasites of Micropterus salmoides (Lacépéde)
from a California pond. Trans. Amer. Micros. Soc., vol. 83, pp.
343-348.

MizELLE, J. D., ano R. C. Hucues. 1938. The North American fresh-
water Tetraonchinae. Amer. Midl. Nat., vol. 20, no. 2, pp. 341-353.

MizELLE, J. D., anp A.:R. Kuucxa. 1953. Studies on monogenetic trema-
todes. XIV. Dactylogyridae of Wisconsin fishes. Op. cit.,. vol. 49,
no. 3, pp. 720-733.

MizELLE, J. D., P. S. Sroxiry, B. J. Jaskosxi1, A. P. SEAMSTER, AND L. H.
Monaco. 1956. North American freshwater Tetraonchinae. Op.
Cre SHO, G5, ia, Jy jojyon IUGHEl re):

>

Monaco, L. H., ann J. D. Mizettr. 1955. Studies on monogenetic
trematodes. The genus Dactylogyrus. Amer. Midl. Nat., vol. 53, no.
2, pp. 455-477.

Price: Notes on Trematodes 67

MuELLER, J. F. 1936. Studies on North American Gyrodactyloidea. Trans.
Amer. Micros. Soc., vol. 55, no. 1, pp. 55-72.

1938. Additional studies on North American Gyrodactyloidea. Amer.
Midl. Nat., vol. 19, no. 1, pp. 220-235.

Price, C. EF. anp J. D. Mizevite. 1964. Studies on mongenetic trema-
todes. XXVI. Dactylogyrinae from California with the proposal of a
new genus, Pellucidhaptor. Jour. Parasitol., vol. 50, no. 4, pp. 572-
578.

Prick, E. W. 1938. The monogenetic trematodes of Latin America. In
Livro Jubliar Prof. Travassos, Rio de Janeiro, Brazil, vol. 3, pp. 407-
413.

Van CieEAve, H. J. anp J. F. Muevver. 1932. Parasites of Oneida Lake
fishes. Part 1. Descriptions of a new genera and new species.
Roosevelt Wild Life Ann., vol. 3, no. 1, pp. 5-71.

Yamacutr, S. 1963. Systema helminthum. Vol. IV: Monogenea and
Aspidocotylea. Interscience Publishers, New York, 699 pp.

Department of Biology, Augusta College, Augusta, Georgia
30904.

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Aerial Respiration in the Florida Spotted Gar

BrtiAN McCorRMACK

Many physotomous fishes use their gas bladder as a temporary
or supplementary organ of respiration. Among these are included
the Osteoglossidae, Mormyridae, Characidae, and Holostei (Ber-
tin, 1957). Among the holostean fishes, the gars (Lepisosteus )
occur naturally in fresh and brackish waters, generally in ponds,
lakes, canals, and estuaries. They are very common in the Mis-
sissippi Valley and Florida. L. platyrhincus De Kay is quite
common in the Everglades, where it feeds on Gambusia affinis,
Fundulus chrysotus, Jordanella floridae, and the young of many
other fishes. It is itself an important prey of the alligator (Alli-
gator mississippiensis). Its chief method of catching food is to
float motionless in mid-water and dart into a school of smaller
fish, seizing one of them in its long jaws armed with needlelike
teeth.

It has long been known that gars may survive in waters of low
oxygen concentration, such as often occur during dry periods in
the Everglades, when numerous fish crowd into the larger ponds.
The gars have often been seen to gulp air at the surface by
rising parallel to the surface and, when their backs touch it, they
thrust their snout out of the water and gulp some air. I have
noticed in the tank that the fish almost invariably, after this,
let out a few bubbles of air through their gill openings. This
“exhalation” can occur at other times as well, however, such as
just before they gulp air, or even while they are merely swimming
or resting on the bottom.

For a long time there was uncertainty as to whether or not the
gas bladder was really the organ whereby the garfish utilized
oxygen from the air. It was not until Potter (1927) analyzed
samples of gas drawn from the bladders of long-nosed gars (Lepi-
sosteus osseus) and showed that a gaseous exchange of oxygen
and carbon dioxide indeed takes place in the gas bladder. The
oxygen concentration was highest immediately after the fish gulped
air and then gradually decreased, while the concentration of car-
bon dioxide increased correspondingly. The gas bladder is thus
definitely the organ of aerial respiration in the garfish. Potter
further estimated that 50 per cent of the oxygen in the air was

McCormack: Aerial Respiration in the Gar 69

absorbed by the gas bladder. When a screen prevented the fish
from going to the surface to gulp air, the fish would die in five to
six hours in water of low oxygen concentration. It is this second
set of experiments that I have tried to repeat and expand, using
L. playtyrhincus as the subject. In this way one can compare how
a different species of gar reacts to the same conditions. Temper-
ature considerations were introduced here, which Potter had almost
entirely ignored, so that the correlation between the two species
can only be approximtae.

My thanks to Professor L. R. Rivas for allowing me the use
of the Ichthyological Laboratory and his own library. My thanks
also goes to Drs. Hunt and Rich for allowing me the use of the

YSI Oxygen Meter.

MATERIALS AND METHODS

The fish used in the experiments were collected at “40 Mile
Bend” on the Loop Road (U. S. 94) off the Tamiami Trail. They
varied from 11-14 inches in total length. Before the experiments
they were kept for several weeks in an unaerated 35 gallon tank
and fed live Gambusia. At the time of the experiments all the fish
were transferred to another tank and then were tested one by one
in their original tank. Some fish were tested in another tank in
which the water temperature was lower. Oxygen concentration
was controlled by means of an air pump. Without a heat control
system, however, the temperature could not be regulated.

The experiments consisted essentially in placing a fish in the
tank, waiting for an hour or so until it got adjusted to its new
environment, which could be inferred when it began to swim
normally and gulp air. A flexible plastic screen was placed on
top of the tank below the surface and wedged in with wooden
supports. In this manner a free exchange of oxygen from air to
water could be maintained, so that the oxygen content of the
water would remain constant. Measurements of temperature and
oxygen concentration in the water were made during the course
of the experiments by a YSI Model 51 Oxygen Meter.

The first three experiments show that Lepisosteus platyrhin-
cus cannot survive without breathing air at temperatures of 20-21
C and 2-3 parts per million of oxygen. When access to air was

70 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

permitted under the same conditions the fish survived for over a
month. L. osseus survived for 5-6 hours in what were probably
the same conditions (Potter, 1927), while L. platyrhincus died in
about an hour. We may thus have here a significant species
variation.

At 21-22 C, and much higher oxygen concentration, as shown
by fish no. 5, L. platyrhincus can survive indefinitely without
aerial respiration. Fish no. 6 shows that this is also true for
temperatures lower than 21 C. This is qualitatively true for many
other air-breathing fishes (Willmer, 1934; Horn and Riggs, MS).
What is interesting to note here, however, is that fishes nos. 5 and 6

TABLE I

Effect of oxygen concentration and temperature on survival of garfish not
given access to surface

Fish No. Time 0, ppm Temp® C Total time

1 1230-1345 2.0 20.0 hrs Semin
2eD 20.4

oe 1600-1650 225 21.0 50 min
25 21.0

3 1400-1500 De 20.4 Ie aie

| NS 20.4

A 1330-1830 6.5 24.0 5 hr
6.5 24.0

5 1800-2300 DD 21.5 5 hr
6.0 22.0

6 1330-0930 6.5 14.5 PAN joie
6.5 15.0

*Fish removed alive.

actually tried to gulp air, even though the experiments showed
that they did not need it. Other fish in conditions identical to
experiment 6, but without the screen, were observed to gulp air
once every 10-15 minutes. This implies that aerial breathing be-
havior may be partially independent of temperature and oxygen
concentration influences. Fishes nos. 1-3 were observed to gulp
air every 2-5 minutes. Thus the rate of aerial gulping does depend

McCormack: Aerial Respiration in the Gar val

somewhat on oxygen concentration, but the behavior is shown
independent of oxygen values as well.

Fishes nos. 4 and 6 show that, as expected, at higher termper-
atures the fish need greater amounts of oxygen. Much more work
needs to be done on this with adequate temperature controls.

The characid, Erythrinus unitaenatus, does not use its gills
at low concentrations of oxygen (Willmer, 1934). The same was
noted in the experiments on Lepisosteus; at oxygen concentra-
tions of 2.0-2.5 ppm the opercles are shut tight. At such low
concentrations of oxygen the fish’s blood may contain more oxygen
than the medium; to expose its gills to the water at such a time
would result only in the loss of more oxygen. Thus the fish con-
serves oxygen by not using its gills.

L. platyrhincus, however, shows opercular activity under these
conditions. Just before the fish gulped air at the surface its
opercules expanded and contracted several times, although the gill
membranes remained shut. This may be the. way the fish draws
out stale air from its bladder preparatory to gulping new air.
Another series of opercular puffings was seen after the fish had
gulped air and exhaled through its gill openings. Perhaps the
fish now was forcing the new air into its bladder.

At higher concentrations of oxygen L. platyrhincus clearly uses
its gills for respiration. Fish no. 4, for example, had a respiratory
rate of 20 opercular pumpings per minute when it was also allowed
to gulp air. Once the screen was put on this rate rapidly climbed
to 40 pumpings per minute.

SUMMARY AND CONCLUSIONS

L. platyrhincus has both aquatic and aerial respiration and
can exhibit both at the same time or either one alone, depending
on the conditions. At low temperatures and high oxygen con-
centrations the fish can survive indefinitely without aerial respira-
tion. The gills are adequate, but the fish still tries to gulp air,
perhaps to ease the work load of the gills. At very low oxygen
concentrations, the fish closes the gill covers and may live indefi-
nitely on aerial respiration alone. At intermediate levels of oxygen
and temperature the fish exhibits both aerial and aquatic respira-
tion.

2 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

LITERATURE CITED

Bertin, LEoN. 1957. Organes de la respiration aérienne. Traité de
Zoologie, vol. 13, fasc. 2, pp. 1363-1398, figs. 972-1002.

Carter, G. S. 1957. Air breathing. In M. E. Brown, Physiology of Fishes,
vol. 1, pp. 65-79.

Porrer, G. E. 1927. Respiratory function of the swim bladder in Lepidos-
teus. Jour. Exper. Zool., vol. 49, pp. 45-67.

Wititmer, E. N. 1934. Some observations on the respiration of certain
tropical fresh water fishes, Jour. Exper. Biol., vol. 11, pp. 283-306.

Ichthyological Laboratory and Museum, Deparrtment of Zo,-
ology University of Miami, Coral Gables, Florida. Contribution No.
60.

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

Invertebrates Found in Water Hyacinth Mats
James O'HARA

Tue water hyacinth, Eichornia crassipes (Mart.), is a perennial,
floating, mat-forming aquatic plant which is known to be estab-
lished in most of the southern states. In many areas where it
flourishes, it produces mats that cover thousands of acres of fresh
water and often develops so thickly as to impede navigation. The
plant has an extensive root system that often reaches a length of
three feet. This gives the hyacinth mat an interface area that is
greater than any other floating aquatic plant. Thus, the presence
of water hyacinths creates a vast new habitat for colonization by
aquatic invertebrates in areas of open water that would normally
be dominated by larger predaceous forms.

A number of studies have shown that direct relationships exist
between aquatic plants and many species of aquatic invertebrates.
Krecker (1939) and Rosine (1955) demonstrated a relationship
between invertebrate populations and the surface area of sub-
merged aquatic plants. Scotland (1934) investigated the species
of animals associated with a floating plant. There have been no
studies done on the relationship between aquatic invertebrates and
water hyacinths.

Goin (1943) studied the lower vertebrate fauna associated
with water hyacinths, and Dickinson (1949) had occasion to
investigate hyacinth infested ponds in a study of the biota of
ponds and ditches. However, he gave no information regarding
the animals associated specifically with hyacinths.

In view of the lack of information concerning this plant-animal
relationship, this study was designed to survey the invertebrate
animals that are associated with the hyacinth root mat and obtain
an indication of their relative abundance. I would like to thank
Dr. B. P. Hunt, Dr. D. R. Paulson, and Dr. E. R. Rich for their
continuing help and guidance throughout this project. I would
also like to thank Dr. F. G. Butcher, Dr. J. P. Moore, Dr. H. van
der Schalie, and Dr. J. E. Wilson for identifying several of the
specimens collected.

This study is based on 11 collections made between December,
1959, and May, 1960, south of Lake Okeechobee, Florida (see
Appendix). Collections were generally made in slow flowing

74 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

canals up to one and one-half meters deep. All collections were
made in water deep enough to allow the collecting net to pass
beneath the roots without disturbing them. The use of the net is
similar to that described by Goin (1942). The sampling differed
from Goin’s by having a smaller mesh screening (30 meshes-to-
the-inch) in the net. One portion of the net was isolated as a
center well with a known area and all quantitative measurements
were based on the organisms collected in this center well. The
animals and hyacinths were placed in a large plastic bag, pre-
served in 10 percent formalin, and taken to the laboratory. The
animals were then washed free of the root mass, identified and
counted. Details of these procedures are given by O'Hara (1961).

The invertebrate fauna associated with water hyacinths is gen-
erally typical of bottom fauna communities. The invertebrates
collected totaled over 44,000 specimens comprising over 55 species.
The total list of species is not complete since not every animal
was fully identified.

The most abundant invertebrate forms collected were the Gas-
tropoda with a total of 2,098 specimens in 11 collections. This
total consisted of about six species and comprised an average of
28 percent of the animals collected (Table I).

TABLE 1.

Invertebrate populations found in hyacinths

Hyallela Insect Other
Collection Animals/m2 Gastropoda azteca larvae animals

i 3,446 62% 3% 2% 33%
2 7,242 40 Pat 18 Pal
3 9,573 60 Hi 10 29
4 8,650 2 61 9 28
» 7,626 2 70 5 23
6 6,505 7 72 3 18
Tl SDT 33 oni 8 Daye
8 10,370 36 0 54 10
9 11,864 49 5 28 18
10 24513 18 2; 35 45
et 84,223 0 95 0 5

Average 16,484 28% 33% 15% 23%

O'Hara: Invertebrates in Hyacinth Mats 75

Hyallela azteca was the most common single species with a
total of 1,622 specimens obtained in collections 1 through 10.
Collection 11 contained a total of 81,430 H. azteca per square
meter. This exceedingly dense population occurred in a perma-
nent lake situation as opposed to the slow flowing canal environ-
ment of the other collections. An average of 33 percent of the
animals collected belonged to this species, although this figure is
heavily weighted by the large population in collection 11.

The total number of insect larvae collected was 1,242 for 11
collections. This total was composed of about 33 species and
comprised an average of 15 percent of the total animals collected.

Other animal groups comprised a very small percentage of the
total animal numbers. In collection 10 Oligochaeta comprised 41
percent of the total number of animals. This high percentage can
be correlated with strong evidence of agricultural pollution at this
location. The worms (Tubifex sp.) were almost totally absent
at this same location one week before collection 10 was made.

The following is a list of the invertebrates collected in water
hyacinths, with notes on their frequency of occurrence.

Class TURBELLARIA

Planarians were found in all collections in small numbers. These
forms apparently become abundant in localized situations with
the greatest concentration found being over 1,000 per square meter
in collection 1.

Class OLIGOCHAETA

Tubifex sp. were noted to be in abundance only in collection
10. These forms are often considered to be indicators of organic
pollution, which agrees with the water condition at the site of
collection 10. Three unidentified species of Oligochaeta were also
found to live in hyacinths.

Class HrupDINEA

Dina lateralis was the most common leech found to live in
association with water hyacinths. Macrobdella ditetra, Helobdella
triserialis, and Placobdella parasitica were also collected. Hiru-
doids were represented in all collections except 1 and 4 but
seldom in large numbers.

76 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Class GASTROPODA

Physa pomilia was the most frequently observed form in all
collections except collections 1, 3, and 11. Helisoma trivolvis
was the most common species found in collections 1 and 3. Collec-
tion 11 contained no Gastropoda. Pomacea miamiensis was repre-
sented occasionally as was Pseudosuccinea columella. Amnicola
sactijohannis and Gyraulus parvus was relatively rare.

Class PELECYPODA

Sphaerium partumeium and Sphaerium securis occurred only
in collections 3 and 5 and not in large numbers.

Class ARACHNIDA

Arrenurus birgei, Arrenurus major, Arrenurus spetiolatus, Ar-
renurus magnicaudatus, and Arrenurus falcicornis were found to
occur in water hyacinths but generally in very small numbers.
One form or another was found in all collections. <A _ birgei
appeared to be the most common form. One unidentified hydra-
carina was also collected.

Class CRUSTACEA

Hyallela azteca was represented in all collections except col-
lection 8. The relative abundance of this form is discussed earlier.
Palaemonetes paludosa occurred in collections 3, 4, 5, 6, 7, 9, and
11 but rarely in very large numbers. Procambarus alleni was
taken in collections 2, 3, 5, and 6, but no more than one specimen
was taken in each.

Class Insecta (Ephemeroptera )

Berner (1950) indicated that Caenis diminuta is rarely found
in association with water hyacinths because of lack of oxygen.
In collection 3, 4, 6, 7, and 8, this species and Callibaetis floridanus
were very common. Collection 8 had a population of 1,652
Ephemeroptera per square meter, of which the greatest number
were C. diminuta.

O'Hara: Invertebrates in Hyacinth Mats el

Class InsEcra (Odonata, Anisoptera )

Of the twenty-nine Odonata nymphs collected, eighteen were
Erythemis simplicicollis. Cannacria gravida, Miathyria marcella,
Pachydiplax longipennis, and Tetragoneuria sp. were represented
in the collections. Anisoptera nymphs were taken in all collections
exeepimcollections 6, 7, 10, and 11.

Class INsEctra (Odonata, Zygoptera )

Ischnura ramburii and Enallagma sp. were identified as occur-
ring in water hyacinths but were never present in large numbers.

Class INsecra (Diptera )

Because of difficulty in obtaining exact taxonomic determina-
tions, insect larvae and pupae other than Ephemeroptera and
Odonata were largely identified only to family groupings. Insect
larvae were found in all collections except collection 11. Although
the representation in each collection was frequently very different,
the total relative abundance is presented here as a composite view
of the insects found in association with water hyacinths.

Chironomidae were by far the most abundant insect larvae
found in the hyacinth community. This group appeared to contain
several species in both larval and pupal stages, although the exact
number of species was not determined.

Ceratopogonidae were fairly abundant and only two species
were distinguished. Both larvae and pupae were collected.

Stratiomyidae larvae were taken only in collection 10 but
they were fairly abundant in this location.

Several unidentified Diptera larvae were also collected.

Class Insecta (Coleoptera )

Hydrophilidae and Dytiscidae adults were rare and no larvae
were taken.

Omophronidae adults were very common, being found in over
half the collections. Haliplidae adults were quite abundant in
localized situations. Haliplus was the only genus noted.

Gyrinidae was represented only by the larval stage and never

78 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

in great abundance, although the adults were commonly observed
in nearby open water. Gyrinus was the only genus noted.

Class Insecta (TRICHOPTERA )

No identification was made on these forms. There are at least
three species present in the hyacinth environment, as determined
by the types of cases found, although all are relatively rare.

Class Insecta ( HEMIPTERA )

Nepidae were uncommon in water hyacinths. Ranatra sp. was
collected only twice.

Naucoridae were common in localized situations and all stages
of development of Pelocoris sp. were taken.

Belostomatidae were infrequent in most collections but oc-
casionally nymphs were observed in large numbers. This was
probably due to a hatching of an egg mass. All stages of de-
velopment of Belosoma sp. were taken.

CONCLUSION

The total number of organisms per square meter shows an
extreme variation from 3,446 to 84,223 per square meter with an
average of 16,484. Attempts to relate invertebrate population
size to-size of the root mass were unsuccessful. The animal
populations are considerably larger than are generally found in
bottom samples and in other plant associations. Ball (1948) re-
ported 2,641 bottom organisms per square meter and 175 organisms
per pound wet weight of Potamogeton in Third Sister Lake in
Michigan. This compares with a calculated minimum of 902
organisms per pound wet weight of hyacinth roots. Cooper
(1941) found the average number of organisms in 44 Maine
lakes to be about 1,600 organisms per square meter of lake bottom.
Unpublished data of the author show an average of 2,449 or-
ganisms per square meter of lake bottom in three Florida lakes.

It is quite evident that the animal populations in water hya-
cinths are usually far larger than the populations obtained from
other aquatic habitats. One major factor contributing to a larger
population is the three dimensionality of the hyacinth en-
vironment compared to the relatively thin bottom environment.

O'Hara: Invertebrates in Hyacinth Mats 79

APPENDIX

All collections except number 11 were obtained in the canals
located alongside the highways indicated.

Collection 1. December 23, 1959. Dade County. 0.5 miles south of 224th
Street on 87th Avenue near Perrine, Florida.

Collection 2. January 15, 1960. Broward County. 1.7 miles south of Florida
Highway 810 on Lyons Road near Margate, Florida.

Collection 3. February 27, 1960. Collier County. 10.4 miles west of Collier-
Dade County line, on U. S. Highway 41.

Collection 4. March 11, 1960. Glades County. 3.8 miles north of U. S. 27
on Florida Highway 78.

Collection 5. March 11, 1960. Glades County. 8.2 miles north of U. S. 27
on Florida Highway 78.

Collection 6. April 29, 1960. Broward County. 6.0 miles north of Andytown
on U. S. Highway 27.

Collection 7. April 29, 1960. Hendry County. 0.6 miles east of Florida
Highway 80 and U. S. Highway 27.

Collection 8. April 29, 1960. Collier County. 36.4 miles north of Carnes-
town on Florida Highway 29.

Collection 9. April 29, 1960. Collier County. 6.4 miles east of Monroe
Station on U. S. Highway 41.

Collection 10. May 6, 1960. Collier County. 36.4 miles north of Carnestown
on Florida Highway 29.

Collection 11. May 6, 1960. Collier County. Peppers Cove on Lake Trafford.

LITERATURE CITED

Batt, R. C. 1948. Relationship between available fish food, feeding
habits of fish, and the total fish production in a Michigan lake. Mich.
State Univ., Agr. Exp. Sta. Tech. Bull., vol. 206, pp. 1-59.

BerRNER, L. 1950. The mayflies of Florida. Univ. Florida Stud., Biol.
Sei, Ser., vol. 4, pp. 1-267.

Cooper, G. C. 1941. A _ biological survey of lakes and ponds of the
Androscoggin and Kennebec River drainage systems in Maine. Maine
Dept. Inland Fish and Game, Fish Sur. Rep., vol. 4, pp. 1-238.

Dickinson, J. C. 1949. An ecological reconnaissance of the biota of some
ponds and ditches in northern Florida. Quart. Jour. Florida Acad.
Sei vol, I pp. 1-28.

Goin, C. J. 1942. A method for collecting the vertebrates associated
with water hyacinths. Copeia, vol. 1942, pp. 183-184.

80 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

1943. The lower vertebrate fauna of the water hyacinth community.
Proc. Florida Acad. Sci., vol. 6, pp. 143-153.

Krecker, F. H. 1939. A comparative study of the animal population
of certain submerged aquatic plants. Ecol., vol. 20, pp. 553-562.

O'Hara, J. 1961. The invertebrate fauna associated with water hyacinths
in south Florida. Unpublished master’s thesis. University of Miami,
66 pp.

Rosine, W. N. 1955. The distribution of invertebrates on submerged plant
surfaces in Mahee Lake, Colorado. Ecol., vol. 36, pp. 380-314.

ScotLanp, M. B. 1934. The animals of the Lemna association. Ecol.,
vol. 15, pp. 290-294.

Department of Biology, Central State University, Wilberforce,
Ohio

Quart. Jour. Florida Acad. Sci. 30(1) 1967 (1968)

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1967

Archbold Expeditions

Barry College

Central Florida Junior @ollese
Florida Atlantic University —
Florida Presbyterian College
Florida Southern College
Florida State University
Jacksonville University
Marymount College
Miami-Dade Junior College
Moin Park Hospital Foundation
Polk Junior College

Rollins College

St. Leo College -

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida
University of Tampa |

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1967

President: JAcKsON P. SICKELS
Department of Chemistry, University of Miami
Coral Gables, Florida

President Elect: CLARENCE C CLARK
Department of Physical Sciences, University of South Florida
Tampa, Florida

Secretary: Joun D. McCrone
Department of Zoology, University of Florida
Gainesville, Florida

Z'reasurer: JAMES B. FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: PreERcE BRODKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges

should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

CY
NA
Fo}G63

~I Quarterly Journal
of the
Florida Academy of Sciences

Vol. 30 June, 1967 No. 2

CONTENTS

Rainfall distribution in the Miami area Louis C. Sass 81

Two ancient Florida dugout canoes
Ripley P. Bullen and Harold K. Brooks 97

Variation in plantar tubercles in Peromyscus polionotus
Michael H. Smith 108

A new gill trematode from Georgia Charles E. Price 111
A new centrolenid frog from Guyana Coleman J. Goin 115
Behavior of iguana during rain storms Coleman J. Goin 119
Additional Miocene anurans from Florida J. Alan Holman 121

Variation in bovine cholinesterase levels
C.J. Wilcox and H. H. Head 141

Officers and members of the Academy 145

SSERARI A Mailed July 24, 1968

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Mares should be 112 inches all around.

Tires should not exceed 55 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and
ADDRESS in paragraph form following Literature Cited.

LITERATURE CiTeED follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TABLES are charged to authors at $19.95 per page or fraction. Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

Lecenps for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($17.30 per page, $15.80 per half
page. Drawincs should be in India ink, or good board or drafting paper, and
lettered by letting guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. Do not submit illustrations needing reduction by more than one-half.
PHotocrapus should be of good contrast, on glossy paper. Do not write
heavily on the backs of photographs.

ProoF must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
FLORIDA ACADEMY OF SCIENCES

Vol. 30 June, 1967 No. 2

Rainfall Distribution in the Miami Area

Louts C. Sass

In the Miami area, one has only to drive down the streets during
a shower to become aware of the highly local and very spotty
nature of many of the area’s rain storms. It may rain hard one day
in one area, while only a few blocks away no rain at all falls. The
newspapers regularly report daily weather data for 4 or 5 of the
local official Weather Bureau observation stations, and these pub-
lished figures for rainfall frequently vary widely from station to
station for the given day. Questions follow as to whether these
evident station-to-station irregularities reported for individual days
will tend to cancel out with time. Or will some meaningful rainfall
distribution pattern appear, and, if so, how long a period of time
might it take for such a pattern to show up? This study answers
these speciiic questions, and then goes on to cover other interesting
aspects of rainfall distribution in the subject area.

SOURCE OF Basic DATA

The basic data behind this report are the raw rainfall statistics,
station-by-station, month-by-month, and year-by-year, for all Dade
County cooperating weather observation stations, that appear, along
with a great mass of other weather material all related to Florida,
in the U. S. Department of Commerce, Weather Bureau's “Clima-
tological Data, Florida’, vols. 52 (1948), through 70 (1966).

SYSTEM OF COMPILATION FOLLOWED

The objective was to develop rainfall data over a long time span
at as many widely separated geographic points as possible within
Dade County. In 1948 there were 11 cooperating weather observa-
tion stations serving the Weather Bureau in Dade County, but two
of these stations (Pennsuco and Tamiami Canal) proved short-

82 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

lived and of little use in this report. Over the next 19 years two
completely new observation stations were added (Royal Palm and
South Miami); three of the old stations changed name with only
insignificant changes in geographic location; and most of the other
old stations underwent slight changes in geographic location but
retained their original names. These complications were straight-
ened out for the author with help from the local Weather Bureau
personnel, and Fig. 1 was developed which graphically portrays
the plan that was followed in compiling the statistics presented
herein. Fig. 1 shows 11 arrows, each arrow representing a “sta-

1948 1951 1954 1957 1960 1963 1966
= es) = == Sa aes = oe] =A

Miami City Weather Bureau 5668
|

! Miami Bayfront Park 5653 fone
Homestead 4091
2 IO yrs.
Coconut Grove I7I16 Miami l2SSW 5678
a 2) 19 yrs.
oO
= Miami Airport 5663
= 4 19 yrs.
3 Hialeah 3909
8 5 19 yrs
oe Miami Beach 5658 9 yrs
e
© Kendall 4518
2 7 19 yrs.
- :
x 5 Peters 7033 Perrine 7020 ree
40 Mile Bend 8780
9 I9yrs.
| iat
10 Royal Palm 60 ISyits

South Miami 3W 8396
It eS [2 yrs,

Fig. 1. Key to plan followed in compiling the statistics presented in this
report.

tion” or locality for which it was possible to develop long term
data. Official Weather Bureau station names and numbers exactly
as given in the Department of Commerce's climatological reports
are shown just above each arrow, and the length of each arrow—
or portion of an arrow—depicts the years that the station was in
service. Finally, the numbers 1 through 11, in order, just to the
left of the arrows, are the simple numbers the author used to iden-
tify his 11 resulting data points, or “stations”, on the maps and
tables presented herein. Fig. 1 is, therefore, the “key” between the
published Weather Bureau records and the numbers presented in
this report.

“INAVIAD HLNOS ©
Alvd WwAOU -
OnN3G JIA OF ©
BNIYY3Id -SY313d |
TVWONIM 4

HIV38 IAVIA

HV31VIN ©
LV0dUIV IAVIA

“ASS 21 IMVIPE-3A089 LNNODOD
Qv31S3R0H
WYVd LNOUS AVE IAVIA-IAVIA

NOILVIIZILNIO! NOlLWLS

|
‘

=“—NmMeHwoer Ooang

8
3NINU3d

fe | LuOdyl

v
TN, LNI AVIA

®
3

S3Y¥OHS (22?

IAVIA

Vek

MISA 1 i 7

Fig. 2. Base map, locating Weather Bureau observation stations.

84 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The map, Fig. 2, geographically locates all 11 of the resulting
“stations” as small numbered black circles against a background
showing the familiar Miami coast line, along with a few well-known
locality names and a few of the principal streets and highways that
traverse the area. This map reveals the areal distribution of the
stations with respect to one another and with respect to the area as
a whole. Good coverage is certainly indicated for the several hun-
dred square miles in the immediate vicinity of the city of Miami,
and extending as far south as Perrine. Unfortunately, coverage
over the remainder of the map area is limited to only three stations
(nos. 2, 9, and 10). Just two or three more stations, judiciously
placed throughout this large open area, certainly would have
strengthened the pictures developed.

AREAL DISTRIBUTION OF ANNUAL RAINFALL

Table 1 presents rainfall statistics (stations versus years ) for the
1] stations in the Miami Area compiled in accordance with the plan
detailed in Fig. 1. At the bottom of the table, the totals of recorded
rain have been divided by the number of years of data involved,
to arrive at a simple arithmetical-average annual rainfall value, rep-
resentative for the locale of each station. These average values
were then transferred to the base map and 5-inch isohyets were
drawn depicting the areal distribution of this rain. Fig. 3 presents
the resulting picture. This figure establishes that over the 19 year
period, while many of the marked differences in rainfall that are so
commonly noted for individual storms in the area do tend to cancel
out with time, there is still a simple, definite, interesting, large scale
pattern that remains. This pattern consists of increasing rainfall
as one moves inland from the Miami Beach station (No. 6), to the
Miami station (No. 1), to the International Airport station (No. 4),
and then on to the Hialeah station (No. 5) or to the South Miami
station (No. 11). Furthermore, a reversal of this trend takes place
somewhere west of the airport—possibly near the junction of Krome
Avenue and the Tamiami Trail—and the rainfall decreases west-
ward from that point out to beyond the station at 40 Mile Bend
(No. 9). There can be no question regarding the validity of the
general distribution pattern as shown on Fig. 3. A glance at Table
1 proves that Station 6 (Miami Beach) and Station 1 (Miami) al-
most every year have had rainfall figures on the low side, whereas
Station 4 (International Airport) and Station 5 (Hialeah) are

V3uV IWVIN Y¥3LV3N9 qzed uy *‘pezemT.ee BEA oindTZ se7eoT PUL
SNOILVLS NOILVAY3SSGO I! 1V

S °29 91°95 e e ; e 5 e iz e e@ ; e
NIVY S3HONI TVANNY 8 ea°cs §—€RtSy 629 ER*BS = dnd 20°19 RES | ar/av |
I 37evWL g9°9€g = OT” Z90T S9°660T 2g°0dg 99°S6TT 2d°dTTT Gh°6STT 26°€20T | eteso,

9°95 42°062

0°9S £5°66S
T°9S €9°2e9
0°9S 9E°HgS
T°9S 45° 21S
8°95 n2°2Sh
0°9S 04°062
6°9S 00°906
S° 4S 68°T69
T°Es 62°€29
2°2es 49°204
T°HS é4°60S
2°ss £9°9T9

2°nS 88°cn9g
0°2s 25° Thh
2°€S 1S°0SE
0°9S GL°22h

S°€9 4Q°62S

2°99 €2°eT9

IvexR 10g But yszodoy
e3er9ay aZeraay guot3e4s
Butuuny | o¢zomyytay 3° °ON

‘IAVIAD HLNOS ©
Wivd WwAOU ©
Onw3@ 2IIN OF ©
BMIdeId-SUILId ©
TIVONIM 4

HIVIG AVIA
HV3ZIVIH ©
LBOd¥IVY INVIA

‘ASS 21 INVIN-3A0¥9 1NNODOD

QV3I1S3W0H ©
NUVd LNOMJ AVE IAVIF-IAVIA

WOILVIIZILNZO! WOILLVLS

: / a

gy} pNNVIN

e
v

VA

@
S

AA ff
by /
CS TISISIY.
Vb Wf,

/

/
Wife. ey
/ ys OSL,
/
/ , 5
/
A / ‘ /
// ‘
/// WWOILWN y

CY ne

4 ’
(f/f SIOW1OMZAa

Isohyets depicting the long-term annual distribution of rainfall in

Fig. 3.
the Miami Area.

Sass: Rainfall in Miami Area 87

almost always high. The data for a few individual years may singly
afford a general isohyet delineation which closely approaches the
long-term pattern, but pictures for most years differ in small de-
tails, one from another, and from the long-term picture. However,
review reveals that these differences in details smooth out with
time, and that something rather close to the general trend will usual-
ly have started to emerge after compiling 4 or 5 years of records,
and will certainly be evident after 10.

The general delineation of the isohyets shown on Fig. 3 bears
a reasonably close resemblance to those shown on the small scale
map of Florida rainfall which is presented on page 22 of the Wea-
ther Bureau’s “Climates of the States, Florida” (2), but Fig. 3 af-
fords far greater detail. In spite of the sparsity of control points
over the western and southern portions of Fig. 3, the author went
ahead and planimetered the land areas between isohyets in order
to arrive at a weighted average annual rainfall for the land area
as a whole. The resulting number (57 inches) came out very close
to the simple arithmetic average of the annual rainfalls for all 1]
stations, which suggested that this given combination of stations
may, where necessary, collectively be considered as reasonably rep-
resentative of the area as a whole.

Figure 4 presents miniatures of the same base map as that of
Figs. 2 and. 3, and on these miniatures 2.5-inch isohyets have been
sketched depicting the areal distribution of the 19 years of rainfall
(1948-1966) by quarters—ie. by three-month periods. It is of in-
terest that a suggestion of the same crude isohyet pattern can be
seen in all four pictures. This tends to deny the possibility that
the residual annual pattern indicated in Fig. 3 is due to anomalous
happenings in any one season of the year.

YEAR TO YEAR VARIATIONS IN RAINFALL

The suggestion previously mentioned, that the simple arithmetic
average of the annual rainfalls of all of my given stations affords a
single figure reasonably representative of the average yearly rainfall
spread over the area as a whole, has been pursued and, at the right
hand side of Table 1, the totals of the recorded rain for all the re-
porting stations for each year are divided by the number of stations
reporting to arrive at yearly averages. These averages are plotted,
year by year, on Fig. 5 which clearly depicts wide variations in
annual rainfall—another characteristic of Miami's precipitation. The

FEB. MAR. APR

(7

a

aNd
IN2z
aNo
oOo 4
a -
Ww qa
> z
w

EVERGLADES —

G

Scale
Miles

NOV. DEC. JAN.

NSS MANS
N MQNAAGGRS
RA MA GEA

RM AAV OO yg \\
\ ~

Fig. 4. Isohyets depicting quarterly distribution of rainfall in the Miami
Area.

AUG. SEP, OCT.

MAY JUNE JULY

Sass: Rainfall in Miami Area 89

range of the values plotted is from a low of 36.60 inches in the dry
year of 1956, up to a high of 82.37 inches in the wet year of 1959
—with the wettest year having more than twice the rainfall of the
driest. The suggestive rather orderly cyclic repetition of peaks and
troughs every 5 to 7 years, evident in Fig. 5, is certainly of interest
and may shed some light on Dade County’s recurrent water table
and drought problems. The horizontal line on Fig. 5 at 56.8 inches
represents the arithmetic average of the 19 individual yearly

90"
80"
i 70'
C) Running Cumulative Average
\4
Q I9 YEAR AVERAGE
XN me
‘ - fi 8 60
Q Ce ©
() s OS 0) 2
So-
1948 1951 1954 1957 1960

Fig. 5. Graph of rainfall in the Miami Area, by years.

90 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

values. The irregular dashed line, broken by circles, shows the
running cumulative average, year-by-year, from 1948 through 1966.
This latter line suggests that at least 10 years of records—and pos-
sibly even 15 years—must be considered in order to arrive at a
tentative “norm” for the area. It is of interest to note that, with
the wide variations in rainfall reported from year to year, even the
full 19-year average of 56.8 inches can itself rise or fall approxi-
mately one inch after the 20th year (1967), should that year happen
to repeat either the previously established maximum or the previ-
ously established minimum.

DIsTRIBUTION OF RAINFALL, BY MONTHS

The same arithmetic average approach that was applied to
yearly data in the preceeding paragraph was applied to monthly
data, and the results are presented in Table 2 and in Fig. 6. The
rainy season is quite clearly May through October. Just over 75
percent of the total rainfall for the year comes in these six months.
The shaded blocks in Fig. 6 depict the 19-year mean values as cal-
culated for each month. The upper dashed line plots the maximum
values that were recorded for each month during the 19 year period,
and the lower dashed line plots minimum values. Table 2 also
gives the years in which the maximum and minimum rains fell, for
each month. The maximum and minimum lines plotted on Fig. 6,

TABLE 2
Monthly rainfall (inches) in Greater Miami Area, 1948-1966

19 Year 19- Year 19 Year 19 Year
Month Maximum Minimum Mean Median
Jan. 5.88 (1958 ) 21 (1949) 1.81 81
Feb. 5:8 p (1957) Al (1962) 1.97 LS
Mar. 6.28 (1959) 3) E1956)) 2.01 1.50
April 10.07 (1960) 24 (1961) 3.26 2 OTe
May 1:82, C1958) 82 (1965) 5.65 4.99
June 18.20 (1966) 3.08 (1952) 8.08 7.74
July 8.73 (1959) 2.78 (1963) 6.25 6.32
Aug. HOS RGl9 57 ) 4.19 (1954) 6.57 6.04
Sept. 21.81 (1960) 2.95 (1961) 9.10 8.11
Oct. 12575) (Gl9G65:) 1.99 (1962) 8.02 7.92
Nov. 9.49 (1959) 43; G1952)) 2.34 2.08
Dec. 5.64 (1958 ) .26 (1962) 1.89 TL OAS)

Totals: 56.95 51.08

Sass: Rainfall in Miami Area 91

when compared against each other and against the means, clearly
demonstrate wide variations in monthly rainfall—yet another char-
acteristic of the rainfall of this area. In fact, the numbers that were
averaged to arrive at the monthly means were spread over such a
wide range of values, and sometimes appeared to present such a
poor distribution of values, that an experimental column was added
to Table 2 to present 19-year median values for each month. These,
with one exception, proved to be lower than the 19-year means,
most of them, by some 5 to 25 percent.

The two summer peaks of rainfall so evident on Fig. 6 are an-
other interesting characteristic of the rainfall of this area. One
peak occurs in June, and the second, in September-October. These

| INCHES OF RAINFALL
fo) ou ro) a

ela ||

+02
G2

‘av [a | aad

190 |udas| ‘onv | Aine] anne! av

Fig. 6. Graph of rainfall in the Miami Area, by months. Shaded area plots
the mean values; the upper dashed line plots maxima; and the lower dashed
line, minima.

92 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

peaks are correlated by Riehl (3) with the mean position of an up-
per air trough which Riehl says extends southward from the middle
latitudes and lies over southern Florida in June. He goes on to
state that this trough is displaced westward into the Gulf of Mex-
ico duing July and August, and then returns to again overlie south-
ern Florida in September-October. Riehl concludes that the heavi-
er and more frequent rainfall occurs when the trough is overhead,
and that the characteristic July-August drop-off in rainfall in south-
ern Florida is due to this trough’s seasonal displacement westward.

RAINFALL DISTRIBUTION By DAYS

The International Airport station (No. 4) and the Miami Beach
station (No. 6) were selected as representative of the area as a
whole for a 10-year study (1955-1965) of a different type—a study
of rainfall by days. This study was planned to afford a measure
of the average number of days per year in the area that have meas-
urable rainfall, and to afford a break-down of these rainy days into
categories based on the amount of rain that fell in the 24 hours
that comprised each of these days. Six categories were arbitrarily
chosen as follows:

Category Range (inches per day)

1 Otome tO
2 JO 25
3 26etoy a0
4 Will 1) we)
5 POQ0stoyeIg
6 2.00 and up

Such a study would reveal how many days on the average have
light showers, medium showers, heavy rains, etc., and just how
much of the total yearly precipitation falls in each category. Re-
sults are summarized in Table 3 and Fig. 7.

In Table 3 the data for the International Airport and for Miami
Beach are shown side by side for ready comparison. It is interest-
ing to note that at the Airport, for example, there were on the ay-
erage 125 days per year with measurable rain, and that, again on
the average, 58.81 inches of rain fell annually. Very light rains
(.01 to .25 inches) accounted for 56.7 percent of the rainy days,
but accounted for only 10.5 percent of the total inches of rainfall.
By contrast, heavy rains (1.00 inch or more) occurred on only 14

Sass: Rainfall in Miami Area 93

percent of the rainy days, but accounted for 58 percent of the
inches. Putting this another way, very light rains fell on 71 days
in the average year but accounted for only just over 6 inches of
rainfall, whereas heavy rains fell on only 18 days, but accounted for
34 inches. The study shows, therefore, that a large percentage of
the year’s rain falls on the relatively few days that the area is sub-

TABLE 3

Airport rainfall (inches) vs. Miami Beach rainfall (inches) on an intensity
basis, with days as the unit of time (1956-1965 )

Miami Miami Differ-

Airport Beach ence
1. Av. no. of days per year with measurable rain 125 118
2. Av. total inches of rain per year 58.81 46.30 = PAS
3. Av. no. of days per year with
between .01” and .10” of rainfall 47 53
3a. Total inches per year of such rain 2.03 QHOM sta OA
Percentage line 3 is of line 1 (days) 37.6 45.3
Percentage line 3a is of line 2 (inches) 3.4 4.5
4. Av. no of days per year with
between .11” and .25” of rainfall 24 23
4a. Total inches per year of such rain 4.20 4.03 ey
Percentage line 4 is of line 1 (days) 19.1 19.5
Percentage line 4a is of line 2 (inches) 7.1 8.7
5. Av. no. of days per year with
between .26” and .50” of rainfall iil ile
5a. Total inches per year of such rain 1.83 6.34 alta
Percentage line 5 is of line 1 (days) 16.9 14.4
Percentage line 5a is of line 2 (inches) 13.3 Sa
6. Av. no. of days per year with
between .51” and .99” of rainfall 15 £3
6a. Total inches per year of such rain 10.60 9.25 — 1.35
Percentage line 6 is of line 1 (days) 8) 10.7

Percentage line 6a is of line 2 (inches) 18.0 20.0
7. Av. no. of days per year with

between 1.00” and 1.99” of rainfall rS 8
7a. Total inches per year of such rain Gil 10.98 = 16,03
Percentage line 7 is of line 1 (days) 10.1 6.8

Percentage line 7a is of line 2 (inches) 29.9 Del
8. Av. no. of days per year with

more than 2.00” of rainfall Sy 4
8a. Total inches per year of such rain 16.54 13.63 — 2.91
Percentage line 8 is of line 1 (days) 3.9 3.4

Percentage line 8a is of line 2 (inches) 28.1 29.4

94 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

jected to larger storms. These larger storms are usually associated
with organized temporarily disturbed bands of “weather” and this,
in turn, explains why rainfall in individual months sometimes de-
parts so drastically from the average. If just one or two heavy
storms fail to materialize within a given 28 to 31 day period, or
just one or two extra storms slip in during that period, the inches
data for that particular month are sharply affected. Fig. 7 plots
the cumulative percentage of days with measurable rain versus
the cumulative percentage of the total annual rainfall, with the
shaded area depicting the whole range of the curves that developed
for the two representative Miami area stations, calculated for in-

CUMULATIVE % OF DAYS WITH RAIN

| zi ec

O 20 40 60 80 100
CUMULATIVE % OF TOTAL RAINFALL

Fig. 7. Percentage plot; days with measurable rain versus total rainfall.
After Riehl.

Sass: Rainfall in Miami Area 95

dividual years as well as for their ten-year composites. These
Miami curves proved to be just a bit more flexed than a similarly-
based curve presented by Riehl (3) which portrayed Argentine
data. The dashed diagonal line on Fig. 7 represents the situation
for a hypothetical region where the same quantity of rain falls on
every single rainy day, and the amount of departure—or flexure—
from this base line is a measure of the variations present in the
quantities of rain that fall on the individual rainy days in the area
under study. Fairly sharp flexures, such as depicted on Fig. 7 and
by Riehl (3), must be generally characteristic of much of the
world’s rainfall. In any event, Miami's flexure is certainly high.

SUMMARY

Rainfall characteristics of the Miami Area can be summarized

as follows:

1. The average annual rainfall based on 19 years of records is
56.8 inches, but wide variations occur from year to year. The
wettest year has more than twice the rainfall of the driest.
The 19-year picture shows interesting rather orderly cyclic
repetitions of peaks and troughs every 5 to 7 year.

2. Some rain occurs in every month of the year, but the rainy
season is from May through October. The rainy season ex-
hibits two maxima, the first in June, and the second in Sep-
tember-October. Wide extremes from year to year in the
amounts of rain that fall in individual months are common.

3. During the dry season it rains on the average on about one
day out of every four or five; during the rainy season it rains
on almost half of the days. Most of the storms are light
showers (less than .25 inches of rainfall). Heavy rains (in
excess of 1.00 inch) occur on only some 10 to 25 days during
the year. Most, but not all, of these heavier storms come in
the rainy months.

4. Many of the storms are highly localized, with good rain in
one locale, and no rain at all in others.

5. A regional pattern of areal distribution of rainfall develops
over the years, with the least rain on Miami Beach, increas-
ing inland to Miami, and continuing to increase on to the
International Airport, and possibly as far west as the junction
of Krome Avenue and the Tamiami Trail. Beyond this max-
imum point, rainfall tends to decrease westward.

96 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ACKNOWLEDGMENTS

Thanks for first sparking the idea to write this report are extend-
ed to my good friend and professor, Dr. Paul S. Salter of the De-
partment of Geography, University of Miami. Thanks are also due
Mr. Leonard G. Pardue of the local U. S. Weather Bureau office on
the University of Miami campus for graciously making the neces-
sary source records available to me. Dr. Luis R. Rivas, of the Uni-
versity of Miami faculty, provided real help and encouragement
toward getting the work published. Finally, the author must
acknowledge his appreciation for the seven excellent line drawings
which were made in the Drafting Room of the Coral Gables office
of Latin American Gulf Oil Co.

LITERATURE CITED

U. S. DEPARTMENT OF COMMERCE, Weather Bureau. 1948-1966. Climatol-
ogical Data, Florida. Vol. 52-70, U. S. Government Printing Office,
Washington, D. C.

U. S. DEPARTMENT OF COMMERCE, Weather Bureau. 1962. Climatography
of the United States No. 60-8, Climates of the States, Florida. U. S.
Government Printing Office, Washington, D. C., 24 pp.

RieHL, HerRBerT. 1954. Tropical Meteorology. McGraw Hill Book Co., New
York, 392 pp.

7280 SW 128th Street, Miami, Florida 33156. Miami Amateur
Meteorological Society, Contribution no. 1 (read March, 1967).

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

Two Ancient Florida Dugout Canoes
Riptey P. BULLEN AND HARotp K. Brooks

THis paper covers the discovery, investigation, and discussion
of two pre-Columbian canoes found in peat deposits of Central
Florida and subsequently dated by radiocarbon methods. The
first canoe, found in Lake Apopka near Zellwood, will be referred
to as the Zellwood canoe to distinguish it from other Lake Apopka
fragments as well as from the canoe subsequently found at Orange
Park near Lakeland. The account of the Lakeland canoe follows
the report on the work at Lake Apopka.

THE ZELLWOoD CANOE

Late in January 1961, Henry Swanson of Orlando, then county
agent, telephoned the Florida State Museum and reported that
Arch Hodges, Superintendent of the Zellwood Drainage District,
had located an Indian canoe under five feet of peat and muck in a
drainage ditch in Lake Apopka. A museum field party visited
the site but excessive rains prevented immediate excavation.

Excavation was accomplished 9 February with the assistance
of labor and necessary machinery kindly supplied by the Zellwood
Drainage District. Continued clearing and slight deepening of
the drainage ditch disclosed two more portions of dugout canoes.
The site was revisited 21 February and the provenience of these
fragments ascertained. Owing to their fragmentary condition and
the time and expense which would have been involved, they were
not removed. W. H. Sears, then at the Florida State Museum,
was in charge of the removal of the first or Zellwood canoe while
Bullen took notes, photographs, and peat samples. On the third
trip, Brooks was also present and took a series of peat samples.

Today, the eastern part of Lake Apopka consists of an exten-
sive peat area which is under intensive cultivation. A dike holds
back the water of the lake while drainage ditches maintain a
proper underground water level. The farm of C, L. Clonts is
located near the dike a considerable distance west of what would
normally be the eastern edge of Lake Apopka. It is separated
from the dike by a drainage ditch which parallels the dike.

The Zellwood canoe was first encountered during cleaning
operations in this ditch. Probing indicated that it extended from

98 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

the ditch northeasterly into the Clonts farm. Excavation showed
it to have been 15 or more feet long, to have been split lengthwise
into two pieces which were lying side by side, and to have been
damaged at one end by the ditch-cleaning machine. As shown in
Fig. 1, it lay in a narrow sapropelic or plastic peat zone which

idee Danae i ene ah naib __ Assumed surface
| of lake if no dyke

Present surface
of peat

O

2 Fibrous peat
3
7 Humic or plastic
- 5 Wo Canoe peat
@
- 6
Cc
ey te
IS
a 8
@ :
02 9 Fibrous peat
10
lI
l2

Marl (?)

Fig. 1. Section at site of Zellwood canoe.

BULLEN AND Brooks: Ancient Dugout Canoes 99

was both overlaid and underlaid by thick deposits of fibrous peat.
From the present surface of the peat down to the highest part
of the canoe was 54 inches. The canoe was entirely within and
surrounded by the zone of plastic peat.

Fig. 2 illustrates the excavation and removal of the Zellwood
canoe. It was necessary to construct coffer dams to isolate the
canoe area from the drainage ditch and then to pump water out
of the canoe excavation. Owing to the weakness of the canoe
(lack of tensile strength) it was necessary to support it by a cradle.
Sides of the canoe were undercut and then supported by wooden
studs placed underneath. The studs were lashed to the canoe by
cloth and ropes passed under the canoe through lateral tunnels
cut in the peat. Then these studs were tied to a third which was
suspended above and distributed the pull from the hoisting equip-
ment. A small boat trailer proved very satisfactory for hauling the
wrapped canoe to the museum.

Samples of peat were taken at a depth of 3 feet 11 inches from
the lower part of the superior zone of fibrous peat, from inside
the canoe itself, and at a depth of 5 feet 2 inches from the upper
part of the inferior zone of fibrous peat. These samples, plus a
piece of the canoe itself, were sent on 22 February 1961 to Dr.
Charles H. Fairbanks, then at Florida State University, Tallahassee,
for transmittal to the Oceanographic Institute of that institution
for radiocarbon dating. At that time an arrangement existed
whereby samples for radiocarbon analysis which pertained to
changes in the sea level would be dated by the Exploration
Department of the Humble Oil and Refining Company. Unfor-
tunately these samples arrived as the arrangement was _ being
terminated and, hence, were never tested.

In 1965 Brooks, in connection with his research on Florida
peat deposits, arranged for a radiocarbon determination which
would date the Zellwood canoe. Efforts were made in vain to
retrieve the samples delivered to the Oceanographic Institute in
1961. Fortunately untreated parts of the canoe were still available
at the Florida State Museum. These were taken by Brooks to the
University of Miami and dated by the Institute of Marine Science
under the direction of Dr. Gote Ostlund. The date (Sample ML-
324) was 1185 +75 years before present or around A.D. 765.

Some of the vegetable matter found within the Zellwood canoe

Fig. 2. Excavating the Zellwood canoe.

BULLEN AND Brooks: Ancient Dugout Canoes 101

was examined by Dr. John H. Davis of the University of Florida
and by Dr. Paul B. Sears, Department of Biology, University of
Louisville, who happened to be in Gainesville at that time. This
material consisted chiefly of rods indentifiable as bottlebrush.
Leaves, which were in actual contact with the canoe, they identi-
fied as water lily leaves. It was their opinion that the sapropelic
or plastic zone in which the canoe was found represented a time
of extremely high water—high enough to prevent the continued
growth of peat—and that after an undetermined length of the time
the water level fell and peat started growing again.

That this zone of plastic peat covered a substantial area is
indicated by the subsequent discovery of parts of two more canoes.
The second fragment was found in the lake side of the same
drainage ditch some 38 feet south of the Zellwood canoe. Probes
indicated this fragment to be about 2 by 4 feet in size. The dis-
tance from the surface of the peat to the highest part of the canoe
was 54 inches as before. Here the dark plastic zone was rather
thick, 20 inches instead of 11, but its top at a depth of 54 inches
was at the same elevation as at location 1. Examination of the
profile indicated that the top 24 inches of the upper peat was
dark, oxydized, and fibrous while the lower 26 inches was striated
into light and dark fibrous zones. This situation was probably also
present at location 1 but was not noted by us because of the
extremely wet conditions under which we operated there.

Fragment 3 was found in the bottom of the same drainage
ditch 40 feet south of location 2 (78 feet south of location 1).
The ditch-cleaning machine had removed the sides of this frag-
ment which extended longitudinally along the ditch for a distance
of about 5 feet. It is possible that fragments 2 and 3 came from
the same canoe.

As fragment 3 was badly damaged and lay below the water
level, we did no work at location 3 except to measure the depth
and to check that it was, like the others, in the same plastic peat
zone. The distance from the present surface of the peat down to
the remaining portion of the sides was 60 inches. This dimension
is 6 inches greater than for either of the other canoe fragments,
possibly because the ditch-cleaning machine had cut off the sides
or, by its weight, forced the fragments a little deeper into the
plastic peat zone.

102 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Cultivation of peat lands, as on the eastern part of Lake Apopka,
results in a gradual loss of peat so that the surface becomes lower
over the years. Enquiry elicited the estimate that the surface in
the Clonts farm area, where the canoe fragments were found,
had lowered about 2 feet since the building of the dike.

THE LAKELAND CANOE

Early in the spring of 1964 Mr. Joe Larned of Brewster,
Florida, informed the Florida State Museum that a dragline in the
American Cyanamid Company’s Orange Park mine had uncovered
three dugout canoes. Of these, two were destroyed during dis-
covery and the third, about 18 inches wide and 20 feet in length,
was removed by Larned. The depth at which these canoes were
found suggested respectable antiquity but their removal prevented
the recording of a profile or other provenience data and the site
was not visited at that time.

However, the potential value of a similar future discovery was
readily recognized, particularly in view of the previous records at
Lake Apopka. It was suggested to Larned that, if such an event
should occur again, we would appreciate being told about it
before the canoe was removed so we could make a scientific
investigation at the site.

Nearly a year later, early in January 1965, Mr. W. E. Stephen-
son, Community Relations Coordinator at the American Cyanamid
Company's Brewster Plant telephoned Bullen at the Florida State
Museum that Larned had located another canoe under some 6 feet
of mud at the Orange Park mine, northeast of Lakeland, Florida.
Ripley P. and Adelaide K. Bullen immediately visited the site.

The Lakeland canoe was located in the side of a drainage
ditch dug by a dragline at the Orange Park mine in 1960 as part
of the American Cyanamid Company’s phosphate mining opera-
tions. While the dragline’s bucket damaged one end of the
canoe, loose dirt, always present during these operations, hid the
damaged end from view. Subsequently erosion and slumpage
exposed a small portion of the canoe and, as discovered by exca-
vation, broke it off about 4 feet into the bank.

The canoe extended into the eastern bank of the drainage
ditch with which it formed an approximately right angle. We
excavated the broken 4-foot front (or back?) end which, after

BULLEN AND Brooks: Ancient Dugout Canoes 103

data were recorded and photographs taken, was then removed.
Later Larned, Stephenson, and others returned to the site and
exposed the rest of it, an additional section 15 feet long. Un-
fortunately the other end was missing. The mid-portion of the
canoe was reburied pending chemical preservation and ultimate
disposition.

The Lakeland canoe was made, or at least hollowed out, by
the traditional method of using fire and water. No tool marks
were observable. Made of a coarse grained wood (Fig. 4), it was
27 inches wide at the top of the sides and 10.5 inches deep with
walls 2.5 inches thick. Its length would be in excess of 19 feet.
Unfortunately, the Zellwood canoe, also of a coarse grained wood,
has no comparable dimensions.

Fig. 3 shows the stratigraphic situation for the Lakeland canoe
while pictures (Fig. 4), taken by Stephenson during excavation,
show details of the front end. The deposits were more compli-
cated than those at Lake Apopka.

The Lakeland canoe lay in a zone of fibrous peaty muck which
was about 12 inches thick. This zone was divided into an upper

0 Present surface

ly Fibrous peaty muck

NM

Black muck

Depthin feet
h WwW

on

) 4 wu + Fibrous

White sand peaty

Canoes, Newb” t__—_—— muck
Black muck

Gray yellowish fine sand

oO

7 Black sand

Fig. 3. Profile at site of Lakeland canoe.

104 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

and a lower part by a thin layer of white sand (Fig. 3). The base
of this zone was approximately horizontal as were those of the
underlying black muck and gray-yellowish fine sand zones. We
did not dig very deep into the basal black sand but from the
appearance of the sides of the drainage ditch it continued down-
ward a substantial distance.

The top of the fibrous peaty muck which contained the canoe,
and also the thin dividing layer of whate sand, curved upward
and downward in conformity with the cross-section of the canoe
(Fig. 3). Another thin layer of white sand was found lining the
inside of the canoe (Fig. 4). Over all was a zone of muck about

Fig. 4. End of Lakeland canoe
during excavation. Left, End in
situ. Upper, Close-up showing
sand.

4 feet thick the upper half of which was rather peaty. There was
not a sharp line but a gradual transition between the black muck
and the fibrous peaty muck portions of this overlying deposit.

The gray-yellowish sand appeared to be an old beach surface.
It sloped downward in the drainage ditch face towards the north
in the probable direction of deep water. Apparently there was
a rise of water which resulted in black muck intrusion upon an
old lake beach surface. Shortly peat started growing on this muck.

BULLEN AND Brooks: Ancient Dugout Canoes 105

A derelict Indian canoe became marooned in this growing peat
deposit and was eventually covered by it. The white sand in the
bottom of the canoe presumably blew in after the canoe became
marooned but possibly was present in the canoe earlier. The thin
zone of white sand which divided this peaty muck into two por-
tions must have been deposited by wind. It must represent a
pause in the building up of this peaty deposit. Other thinner and
less well delineated “layers” of white sand suggest a series of
minor halts in the deposition of the lower fibrous peaty muck zone
after the canoe became entrapped.

A sample of the fibrous peaty muck was taken from immediately
above the thin sand layer (X in Fig. 3) for radiocarbon dating.
It should be a little younger than the canoe itself. This sample
and a piece of the canoe were sent to Isotopes, Inc., Westwood,
New Jersey, for dating. Science is indebted to the American
Cyanamid Company for sponsoring this phase of the investigation.

Sample 1-1662 (peat) gave an age of 2,600 +130 b.p. (about
650 B.C.). Sample 1-1661 (wood) was dated at 3,040 +115
(about 1090 B.C.). These dates are internally consistent. Al-
though 400 years may seem overlong to bury the Lakeland canoe, it
is consistent with the theory that the thin layers of white sand found
immediately above the canoe reflect pauses in the pow of the
peaty enveloping deposit.

GEOLOGICAL IMPLICATIONS

That the age of the Lakeland canoe is about 1800 years older
than the Zellwood canoe is not surprising. Each represents an
interval of time of rising water in the lake basins of Florida.
However, Pirkle and Brooks (1959) have established that normal
climatic fluctuations bring about changes in the water table of
ten or more feet in the porous Floridian aquifer. Both the
Zellwood and the Lakeland sites are lakes controlled by these
climatic and sub-surface hydrological factors.

The general level of the water table and of the lakes in the
karst terrains of peninsular Florida are graded to sea level. Thus
fluctuations due to droughts or excessive precipitation have been
superimposed upon a general rise in water level correlated to a
ten foot rise in sea level during the last 4,500 years (Scholl,
1964). The fibrous peat represents growth and accumulation of
marsh plants in relationship to the rising water level. Interruption

106 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

of deposition of fibrous peat is not always readily distinguishable
as shown by the occurrence of the Lakeland canoe within a marsh
type peat zone. The thin sand stringers are evidence of changing
conditions. However, it is possible in this case that the canoe was
drawn up on a marsh encroaching upon an open lake. The
sapropelic plastic peats represent open lake conditions that existed
intermittently. Study of the peats in several lake basins in Florida
suggests that the interval, 3,000 to 1,000 b.p. (circa 1,000 B.C. to
A.D. 900), was the period of most rapid rise in water level.

CONCLUSIONS

A great many dugout canoes have recently been found in
Florida lakes and streams during long periods of drought and low
water. They fall into three groups, namely: (a) fairly delicate
Seminole canoes with pointed bows, blunt overhanging sterns,
and, frequently, thwarts; (b) rather massive thick-walled dugouts,
pointed at both ends, with no overhang but usually with a trans-
verse hole at one or both ends; and (c) the much more rare,
traditional, Indian dugout canoe with blunt overhanging ends.
The first two were made with iron tools, the last hollowed out by
fire which left irregular bumpy interior surfaces.

Seminoles started settling in Florida after 1715 and their canoes
could have been made any time after that date, but most are less
than 100 years old. The heavy, double pointed, dugouts were
made by early settlers or soldiers and are probably around 100 to
150 years old. One of their chief functions was to serve as ferry
boats.

The “type c” or pre-Columbian dugout canoes were made at
least until 1564 as one is illustrated by Le Moyne (Lorant 1946:
29, 119), and post-Columbian examples may exhibit axe marks.
How long ago Indians made such canoes, and hence had available
water transportation, is of considerable anthropological interest.
The Lakeland and Zellwood finds are the only instances known in
which dugouts have been found under conditions suggesting re-
spectable antiquity. That the radiocarbon dates support the im-
plications of antiquity suggested by the depositional data is grati-
fying. These dates, 1090 B.C., and A.D. 765 respectively, make
the Lakeland and Zellwood canoes the earliest water craft known
for America.

BULLEN AND Brooks: Ancient Dugout Canoes 107

During the last phase of the Orange or fiber-tempered ceramic
period of Florida, steatite vessel sherds are found in Indian mid-
dens. Such sherds have been dated at the J-5, Summer Haven,
and Cotten sites at 1195, 1365, and 1065 B.C. respectively (Bullen
1961, P. 105). As the nearest sources for this material are the
mountainous or hilly regions of Alabama and Georgia and as stea-
tite vessels are both bulky and very heavy, the introduction of
these containers has been considered by some to indicate the
presence of water-transport (Bullen and Bullen 1961, p. 14). It
will be noted the dates given above (1000-1400 B.C.) are about
the same as that given earlier for the Lakeland canoe (1090 B.C.).
Intercommunication and the diffusion of ideas (and/or people)
which occurred towards the end of the Archaic period may have
been to a large extent water borne.

On a continental basis, Coe (1960, p. 384) has suggested
communication and trade between La Victoria, Guatemala, on
the western coast of Central America and the Guayas coast of
Ecuador in South America around 750 B.C. and Ford (1966) is
suggesting early formative influences moved across the Gulf of
Mexico to Florida and Georgia around 2000 B.C. The date of
the Lakeland canoe would make such movements entirely feasible.

LITERATURE CITED

BULLEN, ADELAIDE K., AND RIPLEY P. 1961. The Summer Haven site, St.
Johns County, Florida. Florida Anthr., vol. 14, no. 1-2, pp. 1-15.
BULLEN, RipLey P. 1961. Radiocarbon dates for southeastern fiber tempered

pottery. Amer. Antiq., vol. 27, no. 1, pp. 104-106.

Cor, MicuaEt D. 1960. Archeological linkages with North and South Amer-
ica at La Victoria, Guatemala. Amer. Anthr., vol. 62, no. 3, pp. 363-393.

Forp, JAMEs A. 1966. Early formative cultures in Georgia and Florida. Amer.
Antiq., vol. 31, no. 6, pp. 781-799.

LORANT, STEFAN. 1946. The New World. Duell, Sloan and Pearce, New
York, 292 pp.

PrrKLe, E. C., anp H. K. Brooxs. 1959. Origin and hydrology of Orange
Lake, Santa Fe Lake, and Levys Prairie Lake of northcentral peninsular
Florida. Jour. Geol., vol. 67, pp. 302-317.

ScHOLL, D. W. 1964. Recent sedimentary record in mangrove swamps and
rise in sea level over the southeastern coast of Florida. Marine Geology,

vol. 2, pp. 343-364.

Florida State Museum, Gainesville, Florida 32601

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

Variation in Plantar Tubercles in Peromyscus polionotus

MicHAEL H. SmirH

In the cricetine genus Peromyscus the plantar tubercles on the
hind foot number 5 in the subgenus Podomys, whereas all other
subgenera of Peromyscus are characterized by 6 pads on the
hind foot (Blair et al., 1957; Hall and Kelson, 1959). Podomys
contains a single species, Peromyscus floridanus (Chapman), whose
distribution is restricted to Florida. Data presented below, how-
ever, indicate that the beach mouse, Peromyscus (Peromyscus )
polionotus (Wagner ), sometimes has only 5 plantar tubercles.

The feet of approximately 300 live or recently killed P. poliono-
tus were examined with the unaided eye. Forty-six specimens
came from the Ocala National Forest, Marion County, Florida,
and the rest came from other localities in Florida, Georgia, and
South Carolina. Any specimen suspected of having 5 tubercles
was also examined by dissecting microscope.

All of the plantar tubercles were variable in size but the
outer posterior one was completely lacking on some specimens.
Without magnification, 30 of the 46 specimens from the Ocala
National Forest appeared to have 6 tubercles on both feet; ten had
6 on one foot and 5 on the other, and six had 5 on both feet.
Microscopic examination showed the presence of a slight protuber-
ance in the approximate position of the posterior tubercle on some
of the feet which appeared to have only 5 tubercles. Of the 46
animals thus examined, three had only 5 tubercles on each foot,
four others had 6 on one foot and 5 on the other, and the remain-
ing specimens showed 6 on each foot.

Of the 254 remaining animals from other localities, 50 were also
examined by another person who was equally aware of the vari-
ation in this character. It was found that we disagreed on the
number of tubercles on 20 per cent of the specimens. The geo-
graphical variation of this trait cannot be determined objectively
until a quantitative measure is devised so that the minimum size
of what constitutes a tubercle can be well defined. However,
some specimens from other localities in Florida, Georgia, and
South Carolina seemed to have only 5 plantar tubercles even
when examined microscopically. Thus it is clear that P. polionotus
cannot be distinguished from P. floridanus by the number of tuber-

SmitH: Plantar Tubercles of Beach Mouse 109

cles on the hind feet. It is surprising that this character was
ever used as a diagnostic tool, since variation in the sixth plantar
tubercle was first pointed out by Osgood in his revision of the
genus Peromyscus (1909).

Peromyscus polionotus is the smallest Peromyscus in the United
States and thus can be easily distinguished from P. floridanus,
which is one of the largest species. No overlap occurs in the size
of the ear, hind foot, body, or tail of adult mice of the two species.
For example, the hind foot ranges from 25-27 mm in P. floridanus,
15-19 mm in P. polionotus (Blair et al., 1957). The bacula of the
two species are also easily distinguished (Blair, 1942). A clear
and concise key to the species of Peromyscus should be constructed
without reference to the number of plantar tubercles as a key
character to separate these two species. A taxonomic revision
is not needed because the subgeneric status of Podomys is pri-
marily based upon morphological differences other than the num-
ber of plantar tubercles (Hooper and Musser, 1964).

ACKNOWLEDGMENTS

I would like to thank Dr. Emmet T. Hooper for suggesting
this study and Miss Mary E. Glenn for her assistance in collecting
the data. Thanks are also due to Mr. Robert W. McFarlane and
Drs. E. G. Franz Sauer and Pierce Brodkorb for reading the manu-
script. Preparation of the manuscript was aided by AEC grant
AT (38-1)-310.

LITERATURE CITED

Buair, W. F. 1942. Systematic relationships of Peromyscus and_ several
related genera as shown by the baculum. Jour. Mammal., vol. 23,
no. 2, pp. 196-204.

Buair, W. F., A. P. Baim, PieRCE Bropxors, F. R. CAGLE, AND G. A. MOORE.
1957. Vertebrates of the United States. McGraw-Hill Book Co., Inc.,
New York. 819 pp.

Hai, E. R., anp K. R. Kelson. 1959. Mammals of North America. Ronald
Press, New York, Vol. 2, pp. 547-1083.

Hooper, E. T., anp G. G. Musser. 1964. Notes on classification of the
rodent genus Peromyscus. Occ. Pap. Mus. Zool. Univ. Michigan, no.
GSS pp:

110 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Oscoop, W. H. 1909 Revision of the mice of the American genus Peromyscus.
North Amer. Fauna, no. 28, 285 pp.

Savannah River Ecology Laboratory, U. S. Atomic Energy Com-
mission, SROO, Bldg. 772-G, Box A, Aiken, S.C. 29801.

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

A New Gill Trematode from Georgia
CHARLES E.. PRICE

DeEscripTiIvE research in North American Dactylogyrus began
with an account of D. extensus by Mueller and Van Cleave
(1932), from the gills of the carp, Cyprinus carpio L. With nearly
90 species now known in this genus, it is becoming more difficult
to establish affinities and dissimilarities by reference to the hap-
toral armament. A few species possess anchors, bars, and hooks
which are morphologically distinct, and can be used as guide-
lines in comparative morphology. In the majority of cases, how-
ever, the sclerotized structures of the haptor are of a somewhat
generalized nature.

The copulatory complex seems to afford the best structures
for comparison of species. The cirri of many North American
forms also tend to be patterned around a generalized architec-
ture, however, so the cirrus is many times unreliable as a feature
for establishment of species. The accessory pieces seem to offer
almost unlimited variations.

Five host specimens of the silverjaw minnow, Ericymba buc-
cata Cope, were obtained by trapping. The fresh hosts were
frozen as recommended by Mizelle (1938). The branchial ma-
terial and recovered parasites were treated as prescribed by Price
and Mizelle (1964), and measurements made as outlined by
Price (1966). Appropriate measurements and illustrations were
accomplished with the aid of a calibrated filar micrometer ocular
and a camera lucida, respectively. All measurements are ex-
pressed in microns.

Dactylogyrus jaini sp. n.

Host and locality. Ericymba Luccata Cope, the silverjaw
minnow; Milner’s Branch, three miles SE of Hollanville, Georgia.

Number of specimens studied. Twelve.

Types. Holotype deposited in the helminthological collection,
U. S. National Museum, Washington, D. C. Accession no, 61352.
Paratypes in author’s collection.

Etymology. Named in honor of S. L. Jain, a leader in the
study of the Monogenea of India.

Location of parasites on host. Gill filaments.

Description. A dactylogyrid of moderate size, provided with

112 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

a thin, smooth cuticle. A prominent constriction present in most
specimens, at a level just posterior to copulatory complex; length
482 (457-510), greatest width of body 65 (57-73), just anterior
or posterior to constriction. Anterior cephalic border cleft medi-
ally, resulting in prominent anterior cephalic lobes; lateral cephalic
lobes moderately developed. Eyespots four, members of pos-
terior pair closer together than members of other pair. Very
little tendency toward dissociation of eyespots. Pharynx sub-
circular in both dorsal and ventral views, somewhat elongate
longitudinally; transverse diameter 20 (18-23). Peduncle in most
narrow and elongate, tending to become short and stout in three
specimens. Haptor well set off from body proper, wider than
long; length 54 (50-58), width 72 (65-81).

One pair of anchors, dorsal in position (Figs. 1-2). Each anchor

| 0.05 mm |

Fig. 1. Dactylogyrus jaini sp. n. 1,2, anchors; 3,4, dorsal bars; 5, ventral
bar; 6,7,8, hooks; 9,10, cirri; 11,12, accessory pieces.

composed of (1) a solid base provided with well-defined deep
and superficial roots, the superficial much longer; (2) a solid
shaft; and (3) a solid point; shaft and point meet at a definite
angle. Length of anchor 29 (27-32), width of base 13 (11-14).
Anchor wings prominent. Dorsal bar simple, variable in shape
(Figs. 3-4); length 19 (18-22). Ventral bar in form of a wide
“V”, the lateral arms asymmetrical, with a built-up central portion.
Hooks 16 in number (8 pairs), arranged in essential agreement
with Mizelle and Crane (1964). Members of hook pair 4A ob-
served (Mizelle and Price, 1963). All hooks similar in shape and
subequal in size, except for members of pair 4A (Figs. 6-8).
Each hook composed of (1) a solid elongate base, well differen-
tiated from (2) a relatively narrow elongate shaft, and (3) a

Price: A New Gill Trematode a3

sickle-shaped termination provided with an opposable piece. Hook
lemethissemo. Ih) 16 (15-17); nos. 2 and 6, 18 (17-19); no. 3, 22
(21-24); nos. 4, 5, and 7, 19 (18-20); no. 4A, 10 (9-12).

Copulatory complex composed of a cirrus and an accessory
piece. Cirrus tubular, expanded at point of attachment of tube
and base (Figs. 9-10); length, including base, 35 (33-39). Ac-
cessory piece complex (Figs. 11-12), consisting of a sclerotized
main shaft and an accessory ramus. Main shaft ends in a mildly
recurved point. The accessory ramus, arising at the approximate
midlength of main shaft, curves partially over cirrus tube. At-
tached to distal aspect of accessory ramus is an additional sclero-
tized structure of lesser density than other parts of accessory
piece. Additional sclerotized structure of variable morphology.
Length of accessory piece 20 (19-22). A single prostatic reservoir,
in most cases folded back upon itself. Vagina not observed with
certainty, but an unsclerotized atrium apparently opens near left
body margin, just posterior to copulatory complex. Gonads glo-
bose-ovate in outline, the post-ovarian testis tending to become
somewhat more elongate in some. Seminal vesicle formed by di-
latation of vas deferens.

Vitellaria well-developed, composed of large granules; granules
evenly disseminated, with tendency toward clumping in a few
specimens. Vitellaria does not exhibit strong tendency toward
formation of lateral bands. Intestinal crura confluent posteriorly.

Comparisons. The present form possesses a copulatory com-
plex quite similar to that of D. microphallus Mueller, 1938, re-
covered from Semotilus atromaculatus. The accessory pieces of
the two species are similar except for the lightly sclerotized
accessory appendage present in D. jaini. The cirri of the two
forms are similar, each being rather small in the tubular portion;
the cirrus base, however, is larger and more complex in D. jaini
and additionally possesses a small digitiform process, missing in D.
microphallus. In this particular comparison, there exist enough
dissimilarities in the parts of the haptoral armament to separate
the species by comparing these structures. Although many small
to moderate differences can be detected, there is little doubt that
D. microphallus is the nearest morphological relative of D. jaini.

LITERATURE CITED

MizELLe, J. D. 1938. Comparative studies on trematodes (Gyrodactyloidae )

114 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

from the gills of North American freshwater fishes. Illinois Biol.
Monogr., vol. 17, pp. 1-81.

MIzELLE, J. D., aND J. W. CRANE. 1964. Studies on monogenetic trematodes.
XXIII. Gill parasites of Micropterus salmoides (Lacepede) from a
California pond. Trans. Amer. Micr. Soc., vol. 83, pp. 343-348.

MizELLE, J. D. anp C. E. Price. 1963. Additional haptoral hooks in the
genus Dactylogyrus. Jour. Parasit., vol. 49, pp. 1028-1029.

MUELLER, J. F.. anp H. J. Van CieEave. 1932. Parasites of Oneida Lake
fishes. Part II. Descriptions of new species and some general taxo-
nomic considerations, especially concerning the trematode family Hetero-
phyidae. Roosevelt Wild Life Annals, vol. 3, pp. 79-137.

Pricre, C. E. 1966. Urocleidus cavanaughi, a new monogenetic trematode
from the gills of the keyhole cichlid, Aequidens maroni (Steindachner ).
Bull. Georgia Acad. Sci., vol. 24, pp. 117-124.

Price, C. E. In press. A revised definition of the monogenetic trematode
genus Dactylogyrus, with descriptions of four new species. Proc. Hel-
minth. Soc. Washington.

Price, C. E., AND J. D. Mizetie. 1964. Studies on monogenetic trematodes.
XXVI. Dactylogyrinae from California with the proposal of a new
genus, Pellucidhaptor. Jour. Parasit., vol. 50, pp. 572-578.

Department of Biology, Millersville State College, Millersville,
Pennsylvania 17551.

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

A New Centrolenid Frog From Guyana
COLEMAN J. GOIN

WHILE investigating the status of some frogs from the Guyanas
recently I had occasion to examine two specimens from Guyana
(formerly British Guiana) in the British Museum of Natural His-
tory. They differ from all other species now known from North-
eastern South America and are apparently without a name. I
propose that they be called

Centrolenella taylori, n. sp.

Type. British Museum (Nat. Hist.) 1939.1.1.65, adult male;
at an elevation of 750 ft. along the New River, Guyana; collected
by C. A. Hudson.

Paratypes. British Museum (Nat. Hist.) 1939.1.1.64, adult
male; same data as the type; Rijksmuseum v. Naturrlijke Historie,
11472, Marowijne River, Suriname, 11473, Nassaugebergte, Maro-
wijne, Suriname; and 11474 from Langa Soela on the Paleomeu
River at the base of Grensgebergte, Suriname.

Diagnosis. A toothless centrolenid with a lavender dorsum
(in preservative), white visceral peritoneum, one and one-half
phalanges of fingers three and four free of web» aud a rather
distinct tympanum.

In the absence of teeth it is similar to C. fleischmanni (Boet-
tger, 1893) but it differs from that species in its dorsal pigmen-
tation, more slender build and in having the fingers more ex-
tensively webbed. In pattern and, general build it is similar to
C. antisthenesi (Goin, 1963) but it may be distinguished from that
species by its absence of vomerine teeth.

Description. Vomerine teeth absent; the small, rounded cho-
anae separated by about 3 times their own diameter; tongue
one-half as wide as mouth-opening, rounded, its posterior border
free and unnotched. Snout short, rounded when viewed from
above, truncate in profile, the upper jaw extending hardly at all
beyond lower; nostrils more lateral than superior, not projecting,
their distance from end of snout about equal that from eye,
separated from each other by an interval equal to their distance
from eye. Canthus rostralis slightly defined; loreal region slightly
concave and slightly oblique, the upper lip not flaring out ap-
preciably below it. Eye moderate, very prominent, its diameter

116 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

greater than its distance from nostril; palpebral membrane not
reticulate; interorbital distance slightly greater than width of up-
per eyelid, which is relatively wide and about equal the distance
between nostrils. Tympanum very indistinct, about one-third the
diameter of eye, separated from eye by a distance about equal to
one-half its own diameter. Fingers webbed from base of penulti-

Fig. 1. Type of Centrolenella taylori, BM(NH) 1939.1.1.65. Paul Laessle,
del.

Goin: A New Frog from Guyana IE Th

mate phalanx of fourth to middle of penultimate phalanx of
third, a web at the base between fingers two and three, no web
between fingers one and two. Fingers 3-4-1-2 in order of de-
creasing length; fourth considerably longer than second, just reach-
ing to middle of penultimate phalanx of third; disk of third
covers the tympanic area; no projecting rudiment of a_pollex;
no ulnar ridge; toes nearly completely webbed, the web on fourth
toe reaching the base of the penultimate phalanx, third and fifth
subequal, disk of fourth covering about one-half the tympanic
area; indistinct metatarsal tubercles; no tarsal ridge, no dermal
appendage on heel. Body not elongate, in post-axillary region a
little narrower than greatest width of head; when hind leg is
adpressed, heel reaches to nostril; when limbs are laid along the
side, knee and elbow considerably overlap; when hind legs are
bent at right angles to body, heels overlap slightly. No _ pata-
gium extends from the back of the upper arm to the side of the
body. Skin of upper parts smooth; a very indistinct narrow
glandular ridge passing above tympanic area; skin of throat, chest,
belly and lower surface of thigh smooth; traces of a skinfold
across chest; adult male, vocal sac wrinkled but without pigment.
Skin of head not coossified with skull, roof of skull not exostosed.

Dimensions. Head and body 18.5 mm; head length 5.5 mm;
head width 7.6 mm; femur 9.5 mm; tibia 10.00 mm; heel-to-toe
14.5 mm.

Color in alcohol. Top of head, dorsum, lower arm and outer
fingers, legs and feet lavender although the lavender pigmenta-
tion is restricted to a rather narrow band on the dorsal surfaces
of the thighs. This dorsal pigmentation beset with fairly num-
erous, small rounded pigmentless clear areas giving a pattern of
light or clear spots on a dorsum of lavender.

DIscussION

With the recognition of C. taylori the number of species of
this family in the Guianas and in Venezuela east of the Andes be-
comes four. Two of these are forms with vomerine teeth and
with the first finger shorter than the second. One of these, C. anti-
sthenesi is known only from the Coastal Range in Venezuela and
the other, C. geijskesi (Goin 1966) is known only from the Guyana
highlands in Suriname.

118 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

The remaining two are species without vomerine teeth and
with the first finger longer than the second. They are also similar
in that they have opaque white visceral peritoneum but trans-
parent, unpigmented parietal peritoneum. One of them C. fleisch-
manni, fades to white in preservative while the other, C. taylori,
has a lavender dorsum in preservation. The former species, fleisch-
manni, extends along the coastal region of Venezuela and the
Guianas while C. taylori is at present known only from the Guyana
highlands. It is not at all improbable that the form Rivero (1961:
p. 152) described but did not name from the Guyana highlands
in Venezuela is taylori. He described it as similar to fleischmanni
but with a pigmented dorsum.

ACKNOWLEDGMENTS

I am indebted to Miss Alice G. C. Grandison for making the
specimens in the British Museum available to me; to Dr. Leo D.
Brongersma and Dr. M. Boeseman for making the specimens in
the Rijksmuseum van Natuurlijke Historie in Leiden available
to me; to Mr. Paul Laessle for the figure which accompanies this
description; and to the National Science Foundation for a grant
(GB-3644) in support of my studies on South American tree-frogs.

LITERATURE CITED

BoETTGER, O. 1893. Ein neuer Laubfrosch aus Costa Rica. Ber. Naturf.
Gesellsch., Frankfort am Main, 1892-1893 (1893), pp. 251-252.

Gorn, C. J. 1963. A new centrolenid frog from Venezuela. Acta Biologica
Venezuelica. Vol. 3, no. 18, pp. 283-286.

Gorn, C. J. 1966. A new frog of the genus Centrolenella from Suriname.
Studies on the Fauna of Suriname and other Guyanas. Vol. VIII, no.
32, pp. 77-80.

Rivero, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool. Vol.
L265 no; pphel- 20K

Department of Zoology, University of Florida, Gainesville, Flor-
ida.

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

Behavior of Iguana During Rain Storms
CoLEMAN J. GoIN

WHILE in Paramaribo, Surinam, during July 1966 I made some
observations of the behavior of an iguana (Iguana iguana) that
seem to be worth recording.

In the yard of our house was a pine tree (Pinus carribea),
approximately eight inches in diameter at the base and thirty feet
tall. On July 15, it began to rain and blow about 2:30 in the
afternoon. As soon as the first drops started to fall, a large
iguana came to the tree, climbed until it had reached one of the
uppermost branches, crawled out to the end which was hardly
three-fourths an inch thick, wrapped its fore limbs around the
branch, braced its hind feet against tufts of needles and “held on
for dear life” while the rain pelted and the tree swayed in the
wind. After the storm abated about 5:00 o’clock PM the lizard
settled down on the same limb and so far as we know did not
move until it began to rouse a little after 10:00 the next morning.
It finally made its descent shortly after noon.

After the above observations were made we started to keep
more precise records and found that on afternoons in which we
had even a light shower the iguana spent the night in the pine,
climbing to: its perch before sundown, but if there was no shower
it spent the night elsewhere. For a period of about a week
during which we had no rain the big iguana was not seen
although a small one did feed in broad-leaved trees in the yard
from time to time.

On July 27, the day before our departure, another heavy storm
came up at 3:00 PM and once again the lizard came running
across the lawn, climbed to one of the uppermost branches, and
hung on while the rain pelted and the tree swayed as before.
It stayed in the tree after the storm was over. Since we had to
leave at 8:00 AM on the 28th we were not able to note when it
finally descended.

These observations are not fully in agreement with those of
Swanson (1950) in Panama, where he saw no iguanas in trees
during the periods of cool, wet weather, although he added they
seem to prefer more open trees when it is raining. It would
be interesting to know whether the behavior noted above is

120 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

typical or whether it simply represents the idiosyncratic behavior
of this individual.

My work in Surinam was supported by a grant from the Na-
tional Science Foundation GB-3644.

LITERATURE CITED

Swanson, Paut. 1950. The Iguana Iguana iguana iguana (L). Herpetolozica,
vol. 6, pp. 187-193.

Department of Zoology, University of Florida, Gainesville, Flor-

ida.

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968 )

Additional Miocene Anurans from Florida
J. Atan HoLtMaAn

THE exceptionally large anuran fauna of the early Miocene beds
of the Thomas Farm of Gilchrist County, Florida has been re-
ported on by Tihen (1951), Auffenberg (1956), Holman (1961),
and most recently by Holman (1965). In this last paper the new
material of leptodactylid, ranid, and brevicipitid anurans was
discussed, but the pelobatid, bufonid, and hylid frogs were not
covered, mainly because of the lack of recent comparative ma-
terial. Since that time more comparative material, as well as
some new fossil material has become available. Thus, the present
and final report details the pelobatid, bufonid, and hylid frogs
(as well as new material of other anuran groups), and summarizes
the knowledge of the early Miocene anurans of the Thomas Farm
of Florida.

Anuran skeletons at Michigan State University and material
borrowed or received as gifts from persons in the acknowledg-
ments section have been used as comparative material. The
following abbreviations are used: M.C.Z.—Museum of Compara-
tive Zoology; U.F.—University of Florida; F.G.S.—Florida Geologi-
cal Survey. Measurements are in millimeters.

Thanks go to Bryan Patterson (M.C.Z.), Stanley J. Olsen (F.
G.S.); and S. David Webb (U.F.) for the privilege of studying
fossils in their care. Several people have generously loaned or
given comparative material used in this study. These individuals
include Pierce Brodkorb, Gainesville, Florida; James A. Peters and
William J. Riemer, Washington, D. C.; Robert S. Simmons, Balti-
more, Maryland; Charles F. Walker, Ann Arbor, Michigan; George
B. Rabb and Robert Inger, Chicago, [linois; and John D. Lynch,
Lawrence, Kansas. Donna Rae Holman made the drawings. The
National Science Foundation supported the study through Grant
GB 5988.

FAMILY PELOBATIDAE
Pelobatids are known from the early Oligocene to the recent

of North America. Zweifel (1956) and Tihen (1962) provide
good accounts of the pre-Pleistocene forms.

22 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Scaphiopus ct. Scaphiopus holbrooki ( Harlan)

Previously, Auffenberg (1956) reported on four fragmentary
ilia (U.F. 9896), one maxilla (U.F. 9897), three fragmentary
frontoparietals (U.F’. 9898), and one presacral vertebra (U.F. 9899)
from the Thomas Farm of Florida.

New material. Two left and six right ilia (two left and four
right F.G.S. V- 6063, and two right M.C.Z. 3445); one fragmentary
sacrococcyx (M.C.Z. 3446), two left and three right maxillary
fragments (F.G.S. V- 6084), and two skull fragments (F.G.S. V-
6085 ).

Remarks. Auftenberg discussed ilial characters of fossil and
recent Scaphiopus. He reports that the subgenus Scaphiopus
may be separated from the subgenus Spea on the basis that in the
former group the dorsal prominence is either slightly developed or
absent, and that when it is present it is a small rounded pro-
tuberance that is directed dorsolaterally and lies about halfway
between the base and the end of the acetabular expansion. Auffen-
berg states that in the latter group this prominence is usually
ridge-like, directed more dorsally, and that it contributes to the
height and to the length of the dorsal portion of the acetabular
expansion.

In the present study these characters were re-examined in the
light of additional fossil and recent material. The prominence that
Auftenberg refers to is for the tendinous origin of the vastus exter-
nus head of the M. triceps femoris as determined by the dis-
section of a recent Scaphiopus h. holbrooki from Putnam County,
Florida. A prominence for the fleshy origin of this muscle occurs
in Rana pipiens and in Leptodactylus melanonotus (Holman,
1965, p. 71, Hg. 1, and p. 74, fig, 2): Im ‘thesemspeerecmatre
prominence is flatter and more extensive than in Scaphiopus h.
holbrooki. The following recent pelobatid skeletons were studied
by me. Scaphiopus (Scaphiopus) h. holbrooki (15), S. (Scaphio-
pus) couchi (5), S. (Spea) bombifrons (15), S. (Spea) h. ham-
mondi (2), and S. (Spea) h. multiplicatus (3). Based on this
material, most specimens of Spea and Scaphiopus may be dis-
tinguished from one another on the basis of their ilial prominences,
but there is some overlap in a few large specimens of Scaphiopus
h. holbrooki which have the ilial prominences as well developed as

Hou~MAN: Miocene Frogs from Florida 123

in Spea. Nevertheless, if ilia of the same size are compared be-
tween the two subgenera the prominence is usually more de-
veloped in Spea than in Scaphiopus. Moreover, Scaphiopus hol-
brooki has this prominence more highly developed than in S.
couchi. The eight new ilia (Fig. la) as well as Auffenberg’s four
fragmentary ilia resemble Scaphiopus holbrooki in this character.

Fig. 1. A, left ilium of Scaphiopus cf. S. holbrooki F.G.S. V- 6063; B,
sacrococcyx of Scaphiopus cf. S. holbrooki M.C.Z. 3446; C, maxillary fragment
of Scaphiopus cf. S. holbrooki F.G.S. V- 6084; D, right ilium of Eleuthero-
dactylus sp. indet. F.G.S. V- 6086. Each line equals two millimeters.

The greatest height of the acetabulum of the eight measurable
Thomas Farm Scaphiopus ilia (including Auffenberg’s specimens )
ranges from 1.8 to 3.1 with a mean of 2.41, thus the spadefoots
were rather small. A recent adult male Scaphiopus couchi from
San Luis Potosi, Mexico with a snout-vent length of 63 has a
greatest height of the acetabulum of 4.0.

Fortunately, a pelobatid sacrococcyx (Fig. Ib) is among the
new material from the Thomas Farm. Tihen (1960) has sum-
marized the works of Taylor (1942), Chrapliwy (1956) and
Zweifel (1956), and has added his own comments on the osteologi-
cal variability of the pelobatid sacrococcyx. He indicates that
although some variation exists, there are recognizable subgeneric
and in some cases specific differences in the sacrococcyx of the
genus Scaphiopus. The Thomas Farm fossil is similar to Scaphiopus
holbrooki in having the bony webbing between the sacral dia-

124 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

pophyses and coccygeal shaft limited in extent and regular in
outline, and in having an ovaloid sacral cotyle. In material
available to me, Scaphiopus holbrooki is readily separable from
S. couchi in that it has the bony webbing between the sacral
diapophyses and coccygeal shaft much more limited in extent than
in S. couchi. The fossil clearly resembles S. holbrooki in this
character.

The maxillae of Scaphiopus and Spea are quite distinct from
one another. In Scaphiopus the maxilla in lateral view, is sculp-
tured, flat or slightly to moderately concave, and the dorsal border
of the bone is partly formed by a smooth, somewhat laterally
produced shelf. In Spea the maxilla, in lateral view, is smooth,
moderately to strongly convex, and the dorsal shelf is absent.
The five new maxillae (Fig. lc) are definitely referrable to the
subgenus Scaphiopus on the basis of these characters. Moreover,
in available material, S. holbrooki differs from S. couchi in that in
S. couchi the anterior part of the shelf that forms part of the
dorsal border of the bone is interrupted by a sculptured portion
anteriorly, whereas this portion of the bone is smooth in S. hol-
brooki. The fossils resemble S. holbrooki in this character. Thus,
I would follow Auffenberg in designating all of the Thomas Farm
pelobatid material as Scaphiopus cf. S. holbrooki.

FAMILY LEPTODACTYLIDAE

Leptodactylids are known from early Miocene and from Pleis-
tocene deposits in North America (Holman, 1965). But lepto-
dactylids may have been present in North America as early as
upper Cretaceous (Estes, 1965).

Genus Eleutherodactylus Dumeril and Bibron

Based on the rather limited number of leptodactylid skeletons
available (add the following specimens to the list of specimens
studied in Holman, 1965, p. 69: Eleutherodactylus podociferus
2, Leptodactylus bolivianus 1, Odontophrynus americanus 1, Pleu-
rodema brachyops 3, and Syrrhophus campi 1) the ilium of Eleu-
therodactylus may be separated from Ceratophrys, Odontophyrnus,
and Pleurodema in that the prominence for the origin of the

HouMAN: Miocene Frogs from Florida 125

vastus externus head of the M. triceps femoris is moderately
produced laterally rather than being a highly produced dorsal
spike as in the latter three genera. Eleutherodactylus may be
distinguished from Leptodactylus in that this prominence is less
extensive, more knob-like, and usually less strongly bevelled.
Eleutherodactylus usually has a better developed dorsal ilial crest
than in Syrrhophus and Tomodactylus. Moreover, Eleutherodacty-
lus has the prominence for the vastus externus head of the M.
triceps femoris making a more oblique angle (posteroventral )
to the shaft than in Syrrhophus in which this prominence is
more nearly parallel to the long axis of the shaft. The following
fossil represents the first record of the genus previous to the
Pleistocene.

Eleutherodactylus sp. indet.

Material. Right ilium F.G.S. V- 6086 (Fig. 1d). Collected by
Pierce Brodkorb. The fossil represents a small Eleutherodactylus
that is of about the same size of recent E. ricordi. The little fossil
is very similar to the ilia of several small Caribbean species of
Eleutherodactylus, but because of the large number of described
species of this genus and because of my lack of skeletal material
of these forms I have not made a specific assignment for the
fossil.

FAMILY BUFONIDAE

Bufonids are known from the early Miocene to the recent of
North America. Tihen (1962) summarizes the occurrence of the
known fossil New World bufonids.

Bufo praevius Tihen

Bufo praevius was described from the Thomas Farm by Tihen
(1951) on the basis of a right ilium (M.C.Z. 1991). In addition,
he referred the following material to this species. Eight ilia
(M.C.Z. 1992); two tibio-fibulae (M.C.Z. 2000); five humeri (M.
C.Z. 2002); three urostyles (M.C.Z. 1995); one femur (M.C.Z.
2005) and four vertebrae (M.C.Z. 1996). Auffenberg (1956)
discussed B. praevius and reported on a few additional elements.

126 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Fig. 2. A. top, right ilium of Bufo praevius, bottom, left ilium of Bufo
praevius: B. top, left ilium of Bufo terrestris, bottom, right ilium of Bufo ter-
restris. Each line equals two millimeters.

Finally, Tihen (1962) thoroughly reviewed B. praevius in the
light of his intensive study of the osteology of recent and fossil
Bufo. Tihen placed the affinities of B. praevius with the B.
valliceps group and suggests that it belongs with the Caribbean
section of this group.

New material. Because of the hundreds of new bones available
it was decided to study the large number of new ilia in an
attempt to ascertain if indeed a single species of Bufo is repre-
sented by the Thomas Farm bufonid material as previously sug-
gested by Tihen. This new material includes 175 left and 216
right ilia (135 left and 164 right F.G.S. V- 6087, 32 left and 49
right U.F. 10207, and three left and three right M.C.Z. 3447).
In addition, hundreds of postcranial bones and fragments prob-
ably belong to this species.

Remarks. The series of 391 Thomas Farm Bufo ilia was
compared with series of several recent Bufo species in an attempt
to ascertain whether or not a single species of fossil was repre-
sented. I feel that Tihen was correct in assigning the Thomas Farm
bufonid material to a single species. But the ilial prominence is
more variable in shape than he indicated. Tihen (1962) states
that this prominence is quite low, having a consistent height of
about 20 per cent of the length of its base. Actually, although
many individuals have a prominence with a height of about 15 to
20 per cent of the length of its base, some forms, usually those
of larger size, have a much higher prominence (Fig. 2a). The

HotMan: Miocene Frogs from Florida 127

same variation in the shape of the ilial prominence obtain in
recent Bufo terrestris (Fig. 2b).

FAMILY HYLIDAE

North American hylid fossils have previously been reported
from the lower Miocene of Colorado (Chantell, 1965), Florida
(Auffenberg, 1956 and Holman, 1961); from beds that are transi-
tional between the uppermost Miocene and lowermost Pliocene in
Nebraska (Chantell, 1964 and 1966); from upper Pliocene deposits
in Kansas (Tihen, 1960 and Chantell, 1966); and from numerous
Pliestocene deposits. Estes (1964) indicates that there are “sug-
gestions’ that hylids may have been present in the upper Cre-
taceous of the Lance formation of Wyoming. Remarks on the
structure of recent hylid skeletons in the following sections are
based on the following comparative material: Acris crepitans (15),
A. gryllus (6), Anotheca coronata (2), Diaglena reticulata (2),
Gastrotheca marsupiata (1), Hyla arenicolor (7), H. californiae
(2), H. cinerea (4), H. crucifer (3), H. ebraccata (1), H. elaeo-
chroa (5), H. eximia (2), H. femoralis (2), H. gratiosa (3),
H. miotympanum (5), H. phaeocrypta (2), H. regilla (6), H.
septentrionalis (1), H. squirella (10), H. versicolor (10), H. wrigh-
torum (1), Limnaoedus ocularis (3), Pseudacris nigrita (8), P.
ornata (1), P. streckeri (14), P. triseriata (10), Phrynohyas spi-
lomma (2), Phyllomedusa dacnicclor (3), Pternohyla fodiens (3),
Smilisca baudini (9), and S. phaeota (1). The terminology
used by Chantell (1965, fig. 1) to describe features of the hylid
ilium is used here. As has been pointed out by Chantell, the
ilium is probably the best element to use to ascertain the relation-
ships of fossil hylids. In the study of the above material I find
that in some cases genera are quite satisfactorily separated from
one another on the basis of ilial characters, especially when these
genera consist of only a few or of a single species. Thus, Acris
and Anotheca may be quite easily defined on the basis of the
ilium. But in the large genus Hyla the species are so variable
that Hyla cannot be satisfactorily separated from Diaglena, Phry-
nohyas, Pternohyla, and Smilisca. Nevertheless, in many cases
identifications can be made at the specific level in these groups.

I would like to make a few comments at this point on the four

128 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Hyla “groups” of Lynch (1965). On the basis of material I have
studied the Hyla veriscolor group (H. versilcolor, H. phaeocrypta,
H. gratiosa, and H. andersoni) and the Hyla cinerea group (H.
cinerea, H. squirella, and H. femoralis) are not distinct from one
another. The H. versicolor group is defined as having a spine-
like prominence, whereas the H. cinerea group supposedly has an
oval-shaped prominence. I find only a few individuals of species
of the versicolor group with spine-like prominences, indeed, most of
them have ovaloid prominences. Moreover, quite a few smaller
individuals in the cinerea group have round prominences.

Genus Acris Dumeril and Bibron

Several workers (Chantell, 1964 and 1965, Holman, 1962 and
1964, and Lynch, 1962 and 1966) have pointed out characters
that are diagnostic for the ilia of the genus Acris. These charac-
ters include the anterior position of the dorsal protuberance; the
shape and narrowness of the ventral acetabular expansion, and
the presence of a thin ridge along the top of the ilial shaft.

Acris sp. has previously been reported from the lower Miocene
of the Pawnee Creek formation of Colorado (Chantell, 1965), and
Acris cf. A. crepitans has been reported from the Miocene-Pliocene
boundry of the Valentine formation of Nebraska (Chantell, 1964).
The genus has been reported from numerous Pleistocene localities.
New material from the Thomas Farm contains the ilia of a
distinctive new species of Acris.

Acris barbouri sp. nov.

Holotype. Right ilium U.F. 10208 (Fig. 3a). From Hawth-
orne formation, lower Miocene, Arikareean; Thomas Farm, Gil-
christ County, Florida. Collected by Clayton E. Ray.

Paratype. Right ilium (F.G.S. V- 6088). From the same lo-
cality as the holotype. Collected by Pierce Brodkorb.

Diagnosis. A Miocene Acris that is readily distinguished from
the recent species Acris crepitans Baird and Acris gryllus Le
Conte in that its dorsal protuberance is relatively smooth and has
its long axis about parallel to the long axis of the shaft, and in
that its dorsal prominence is relatively short. The recent species

Houtman: Miocene Frogs from Florida 129

Fig. 3. A, holotype right ilium of Acris barbouri sp. nov.; B, holotype left
ilium of Hyla miofloridana sp. nov.; C, right ilium of Hyla sp. indet. Each
line equals one millimeter.

have the dorsal protuberance roughened or deeply notched, and
with its long axis posteroventrally oblique to the long axis of the
shaft. The dorsal prominence is longer in the recent species.

Etymology. The species is named in honor of the late Dr.
Thomas Barbour in recognition of his successful efforts in securing
the Thomas Farm as a study area.

Description of holotype. The dorsal acetabular expansion is
broken just posterior to the dorsal protuberance. The dorsal
prominence is not highly distinct and is shorter than in recent
species of Acris studied. The dorsal protuberance is anterior in
position, having its posterior border even with the anterior border
of the acetabular fossa. The dorsal protuberance is ovaloid in
shape, smooth, and has its long axis about parallel to the long
axis of the shaft. The acetabular fossa has its posterior portion
broken. It is moderately excavated and has its lower border well
extruded. The tip of the ventral acetabular expansion is broken.
The anterior border of the ventral acetabular expansion makes an
angle of about 90 degrees with the shaft. The ventral acetabular
expansion is quite narrow. The ilial shaft is compressed and has a
moderately excavated area on its lateral surface just anterior to the
dorsal protuberance.

Variation in the paratype. The paratype is from a somewhat
larger individual and shows some slight differences that may be

130 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

attributed to individual variation. The dorsal protuberance is a
little rougher and the angle between the ilial shaft and the
anterior border of the ventral acetabular expansion is slightly
greater than in the holotype. The ilial shaft of the paratype is
more complete and shows the beginning of the thin dorsal ridge
that is found in recent Acris species.

Remarks. The Thomas Farm ilia are easily separable from
recent Acris species. This is interesting in that the recent species
are almost impossible to separate from one another and in that
Chantell states that in his Acris sp. from the lower Miocene of
Colorado “The differences between the fossil ilium and the modern
ilia of both species appear so slight that I believe the specimen
is simply a lower Miocene representative of population lines lead-
ing to modern A. crepitans or A. gryllus”. If this is indeed the
case, then two species of Acris were present in the lower Miocene
of North America.

Genus Proacris Holman

This extinct genus was described by Holman (1961) and re-
mains the most bizzare element of the Thomas Farm anuran
fauna and the only currently recognized extinct anuran genus of
the late Tertiary in North America. Unfortunately, although much
additional anuran material has accumulated from the Thomas
Farm, none of it is assignable to Proacris. The ilium of Proacris
(F.G.S. V- 5950) is easily separable from all of the hylid species
I have seen in having its acetabular cup much enlarged and
almost entirely encroaching the much reduced ventral acetabular
expansion, and in having a pronounced ridge just posterior (not
anterior as it states in Holman, op. cit.) to the dorsal protuberance.

Proacris mintoni Holman

Remarks. Holman suggested that Proacris might either be
ancestral to Acris (the fossil resembles Acris somewhat in the size
of its ventral acetabular expansion and in the anterior position of
its dorsal protuberance) or that possibly Proacris represented an
archaic hylid line that is not particularly close to the ancestry
of living forms. With the occurrence of Acris barbouri in the
same deposit the latter thesis becomes acceptable. Possibly, both

HoLtMAN: Miocene Frogs from Florida 131

Acris and Proacris have a common ancestor that lived quite
early in the Tertiary.

Genus Hyla Laurenti

North American fossils of the genus Hyla have been reported
from the lower Miocene of the Thomas Farm of Florida (Auffen-
berg, 1956); from beds that are transitional between uppermost
Miocene and lowermost Pliocene times in Nebraska (Chantell,
1964) and from numerous Pleistocene deposits.

The ilium of Hyla differs from Acris in that the angle between
the anterior border of the ventral acetabular expansion and the
ilial shaft is always greater than 90 degrees, where in Acris it is
about 90 degrees. In most Hyla the dorsal protuberance is more
posterior with respect to the anterior edge of the acetabulum than
in Acris (Chantell, 1964, Holman, 1964, Lynch, 1966). In most
Hyla the anterior edge of the dorsal protuberance lies about even
with or posterior to the anterior edge of the acetabulum, whereas
in Acris at least half of the length of the dorsal protuberance
projects anterior to the anterior edge of the acetabulum. Hyla
californiae, H. crucifer, H. eximia, and two of six.H. regilla are
similar to Acris in this character. In Hyla the ventral acetabular
expansion is much wider than in Acris. Finally, with the exception
of one of seven Hyla arenicolor and two of three H. crucifer,
the thin dorsal ridge that is present on the ilial shaft of Acris
(Lynch, 1962) is absent in Hyla.

Hyla differs from Anotheca in that the dorsal protuberance of
the ilium, although quite variable in shape, is always better
developed. In Anotheca the dorsal protuberance is obsolete and
quite irregular in shape. But in other ilial characters the two
genera are quite similar.

The ilium of Hyla differs from Gastrotheca in that the angle
between the anterior border of the ventral acetabular expansion
and the ilial shaft is always more than 90 degrees in Hyla and is
about 90 degrees in Gastrotheca. The ilia of these genera other-
wise are similar.

Hyla may be separated from Limnaeodus in that the angle
between the anterior border of the ventral acetabular expansion
and the ilial shaft is always more than 90 degrees in Hyla, whereas
in Limnaeodus this angle is about 90 degrees. Moreover, Hyla

132 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

usually lacks the thin dorsal ridge that occurs on the ilial shaft of
Limnaeodus.

In Hyla the dorsal protuberance is rounded, ovaloid, triangular,
or very occasionally irregular in shape, and is produced dorso-
laterally, whereas in Phyllomedusa the dorsal protuberance is
spike-shaped and dorsally produced, much as in some species of
Bufo. Moreover, in Hyla the ventral acetabular expansion is
wider and has its ventral border much less broadly truncate than
in Phyllomedusa.

Hyla and Pseudacris are quite similar in ilial structure, the
most consistent difference between the two genera being the wider
ventral acetabular expansion of Hyla.

I am unable to distinguish the ilia of Hyla from Diaglena,
Phrynohyas, Pternohyla, and Smilisca on any single character.
Thus, the identification of hylid fossils of the above genera must
be made at the specific level.

Hyla miofloridana sp. nov.

Holotype. Right ilium U.F. 10209 (Fig. 3b). From Hawthorne
formation, lower Miocene, Arikareean; Thomas Farm, Gilchrist
County, Florida. Collected by Clayton E. Ray.

Diagnosis. A moderately large Hyla showing similarities to
Recent Hyla cinerea (Schneider), H. gratiosa Le Conte, and H.
versicolor Le Conte, but differing rather strongly from these forms
in having the dorsal protuberance less produced and less distinct
from the ilial prominence, and in having a groove with a strong
ventral border on the lateral face of the ilial shaft just anterior
to the acetabulum.

Description of holotype. The tip of the dorsal acetabular
expansion is broken and the part remaining has its broken surface
worn rather smooth. The dorsal protuberance is ovaloid in shape,
rather weakly produced, and it is not highly distinct from the
dorsal prominence. The protuberance is slightly roughened in
shape. The anterior edge of the dorsal protuberance lies even
with the anterior edge of the acetabulum. The distance of the
protuberance to the acetabular border is less than one-third the
length of the protuberance. The acetabular fossa is only moder-
ately excavated and its border is rather weak. The ventral

HouMaANn: Miocene Frogs from Florida 133

acetabular expansion is well developed and wide. The anterior
edge of the ventral acetabular expansion makes an angle of much
greater than 90 degrees with the ilial shaft. The extreme tip of
the ventral acetabular expansion is broken. The ilial shaft is
compressed and lacks a ridge or crest along its dorsal surface.
Just anterior to the anterior edge of the ventral acetabular ex-
pansion, the shaft has a pronounced groove on its lateral face.
This groove has a rather strong ventral border. Measurements:
greatest height of shaft 1.8, height of acetabular fossa 2.8, length
of dorsal protuberance 1.4.

Remarks. The fossil is most similar to recent H. cinerea, H.
versicolor, and especially to H. gratiosa, but it shows differences
that I believe indicate it is at least specifically distinct. The
grooved lateral surface of the ilial shaft is a very strong character
that I have found duplicated in only one other hylid, a single
specimen of H. septentrionalis. But H. septentrionalis differs
from the fossil in several characters as follow: the dorsal pro-
tuberance is triangular in shape and strongly produced laterally;
the acetabular cup is more strongly excavated and has a stronger
border; and finally, the anterior edge of the ventral acetabular
expansion is more highly curved and makes a wider angle with

the shaft.

Hyla sp. indet.

An ilium (F.G.S. V- 6089, Fig. 3c) may represent a second
moderately large species, but unfortunately, the bone is too frag-
mentary for a specific identification. In fact, part of the fossil
in the area of the ventral part of the acetabular region is so worn
that many important features are obscured. A description of the
fossil is as follows. There is no fossa posterior to the dorsal
protuberance. The anterior edge of the dorsal protuberance lies
slightly anterior to the anterior edge of the acetabular fossa. The
dorsal protuberance is ovaloid and strongly produced laterally.
The shaft is compressed and lacks either a dorsal ridge or a crest.
There is no groove on the lateral face of the ilial shaft. The
dorsal acetabular expansion, almost all of the ventral acetabular
expansion, and about the posterior one-half of the acetabular fossa
are broken.

134 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Hyla goini Auffenberg

Hyla goini was described from the Thomas Farm by Auffen-
berg (1956) on the basis of a right ilium (M.C.Z. 2277) and
three paratype ilia (U.F. 9900).

New material. Fourteen left and 20 right ilia (11 left and 14
right F.G.S. V- 6090, three left and five right U.F. 10210, and
one right M.C.Z. 3448); four left and one right scapulae (three
left F.G.S. V- 6091, and one left and one right M.C.Z. 3449)
one radioulna (M.C.Z. 3450); and one coracoid (M.C.Z. 3451).

Remarks. In the light of the additional fossil material and
much more comparative material it was decided to re-examine
Hyla goini. On the basis of this re-study it appears that all of the
small Hyla ilia of the Thomas Farm represent one variable species
of Hyla, namely Hyla goini. The variation encountered in the 38
Hyla goini ilia is very similar to that one finds in a series of ilia
of recent Hyla squirella, a small species that occurs in Florida
today. Some of these variations appear to be ontogenetic. In
small Hyla goini and H. squirella the dorsal protuberance tends
to be small, round, and relatively far away from the dorsal border
of the acetabulum (Fig. 4a and b). But in larger individuals of
the fossil and of the recent species the protuberance is larger,
ovaloid in shape, and relatively near the dorsal border of the
acetabulum. Individual variation in H. goini and H. squirella
includes such features as slight differences in the shape of the
dorsal protuberance, small differences in the angle between the
anterior border of the ventral acetabular expansion and the shaft,
and slight differences in the width of the ventral acetabular
expansion.

A redescription of Hyla goini in the light of the new ilia that
are available is as follows. The dorsal acetabular expansion is
pointed and moderately long. The dorsal protuberance is well-
developed and is produced dorsolaterally. It ranges in shape
from round in small individuals to ovaloid in larger forms. The
anterior edge of the dorsal protuberance is usually even with the
anterior edge of the acetabular border, but in some individuals
it lies somewhat anterior to this border. The drawing of the
holotype of Hyla goini in Auffenberg (1956, p. 9, fig. 3f) does
not accurately show the appearance of the dorsal protuberance in
this species. In the figure the dorsal prominence is produced

HoutMAN: Miocene Frogs from Florida 135

Fig. 4. A, top, left ilium of Hyla goini, bottom, right ilium of Hyla goini;
B, top, right ilium of Hyla squirella, bottom, left ilium of Hyla squirella. Each
line equals two millimeters.

above the dorsal protuberance. In the ilia of Hyla goini the
protuberance is produced above the prominence. The distance of
the dorsal protuberance from the acetabular border varies from
that of a little under the length of the protuberance to about one-
third the length of the protuberance. The ilial shaft is compressed
and lacks a dorsal crest or ridge. Its lateral side is usually smooth,
but some individuals have a rather indistinct lateral groove or
depression. The anterior border of the ventral acetabular ex-
pansion always makes an angle of well over 90 degrees with the
ilial shaft, but there is some variation in this angle. The ventral
acetabular expansion ranges in width from about one-third as wide
as the length of the acetabulum to about one-half as wide as the
acetabulum. In the few specimens that have an almost complete
ventral acetabular expansion, the tip appears to have been quite
pointed.

Hyla goini is quite similar to recent Hyla squirella and H.
femoralis. Of these two species, the fossil is closer to H. squirella
than to H. femoralis on the basis of the shape of the dorsal
protuberance which is never as elongate as in H. goini and H.
squirella. In fact, I can find no consistent characters by which
to separate H. goini from H. squirella on the basis of the ilium,
and indeed, H. goini may well be directly ancestral to H. squirella.
I can see no resemblance of H. goini to Pseudacris as suggested
by Auffenberg (1956).

FamiLy RANIDAE

Ranid fossils are known from the early Miocene to the recent

136 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

of North America. Holman (1965) summarizes the occurrences of
known ranid fossils in North America.

Rana cf. Rana pipiens Schreber

Two ilia (one left U.F. 10203 and one right M.C.Z. 1994)
that Holman (1965) was unable to distinguish from recent Rana
pipiens have previously been reported from the Thomas Farm.

New material. Three right ilia (F.G.S. V- 6092). These ilia
are also indistinguishable from recent Rana pipiens and are thus
tentatively assigned to this species.

CHECK LIST OF THOMAS FARM ANURAN

Extinct species are marked by an asterisk, extinct genera by
two asterisks. Numbers refer to minimum number of individuals
represented of each species )

Family Pelobatidae
Scaphiopus cf. Scaphiopus holbrooki, 6

Family Leptodactylidae ©

*Leptodactylus abavus, 4
?* Eleutherodactylus sp. indet., 1

Family Bufonidae
*Bufo praevius, 216

Family Hylidae
**Proacris mintoni, 1
*Acris barbouri, 2
*Hyla miofloridana, 1
?*Hyla sp. indet., 1
*Hyla goini, 20
Family Ranidae
*Rana miocenica, 1
*Rana bucella, 1
Rana cf. Rana pipiens, 4
Family Brevicipitidae
Gastrophryne cf. Gastrophryne carolinensis, 1

Hou“MaAn: Miocene Frogs from Florida 137
PALEOENVIRONMENT

Estes has supported the more or less widely accepted idea
that the Thomas Farm deposits represent the filling of a sinkhole in
a porous, eroded limestone terrain. Estes further points out that
“This situation is most plausibly interpreted as a sinkhole with
internal drainage (probably spring-fed, since connection with other
drainage systems would be indicated by the presence of fresh-
water fishes) which, as a result of the available water, drew to it
animals from diverse habitats.

The anuran fauna represents an assemblage that could have
existed in the immediate vicinity of a spring-fed, sinkhole pond.
It is difficult to suggest the exact nature of the pond, but it seems
quite possible that it could have been a rather temporary one.
With the possible exception of a few ranids, all of the Thomas
Farm anurans should have been able to breed in such situations.
It is interesting to note in this regard that 83 per cent of the
minimum number of individual anurans in the Thomas Farm fauna
(216 out of 259) belong to a single species of toad, Bufo praevius.
Toads of the genus Bufo typically breed in small ponds, often ones
of a temporary nature. The lack of large “bullfrogs” of the genus
Rana has been noted before (Holman, 1965). Possibly the
absence of large ranid frogs is correlated with the fact that the pond
was a temporary one, for most large Rana require a relatively long
time for larval development.

PHYLETIC RELATIONSHIPS

The most striking aspect of the Thomas Farm anuran fauna is
that it is an essentially modern one. All of the anuran families
that occur in Florida today are present and there are no extinct
families. All of the genera that are indiginous to Florida today
are present with the exception of Pseudacris and Limnaeodus,
and only one genus (Proacris) is extinct. Two genera (Eleu-
therodactylus and Leptodactylus) that occur in the Thomas Farm
fauna do not occur naturally in Florida at present, but are found
in the western Gulf States and in the Caribbean region today.

Three species (Scaphiopus cf. S. holbrooki, Rana cf. R. pipiens,
and Gastrophryne cf. G. carolinensis) are indistinguishable from

138 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

species living in the area today, and indeed may represent these
species. Other species are possibly directly ancestral to species
living today. These include Leptodactylus abavus which is similar
to recent L. melanonotus and Hyla goini which is similar to recent
Hyla squirella. Other forms (Eleutherodactylus sp., Bufo prae-
vius, and Hyla miofloridana) are of less certain specific relation-
ships. Eleutherodactylus sp. is similar to several small species of
the West Indian region. Bufo praevius is similar to the Caribbean
section of the Bufo valliceps species group of Tihen (1962). Hyla
miofloridana shows similarities to H. cinerea, H. gratiosa, and H.
versicolor. Finally, there are several quite distinct species that
probably became extinct without replacement. These forms in-
clude Acris barbouri, Rana miocenica, and Rana bucella.

ZOOGEOGRAPHIC AFFINITIES

Because of the fragmentary and localized nature of the anuran
fossil record it is difficult to make zoogeographical interpretations
based on the Thomas Farm anuran fauna. Nevertheless, some
of the fossil anuran taxa can be grouped in zoogeographic cata-
gories relative to the distribution of related living forms. Species
of unknown geographic affinities such as the extinct species Acris
barbouri, Proacris mintoni, Rana miocenica, and Rana bucella;
and forms that are of very widespread occurrence today such as
Rana cf. R. pipiens cannot be grouped zoogeographically. The
other Thomas Farm anurans are grouped under the following
tentative catagories.

West Indian.—These include Eleutherodactylus sp. and Bufo
praevius. Eleutherodactylus sp. is similar in structure to several
small Caribbean species of the genus. Eleutherodactylus ricordi
occurs in Florida today, but it is said to have been introduced
from the West Indies (Schmidt, 1953). Bufo praevius is placed
in the Caribbean section of the Bufo valliceps species group, and
it is suggested that B. praevius may have arrived in Florida
from Cuba (Tihen, 1962). It is interesting to note that Estes
(1963) identified Leiocephalus, a lizard with definite West Indian
affinities, from the Thomas Farm. Estes suggests that this lizard
represents a sweepstakes occurrence.

Southeast Coastal Plain—The present day anuran fauna of

HoL~MAN: Miocene Frogs from Florida 139

Florida is one that has largely been derived from the Southeast
Coastal Plain of the United States. Thomas Farm anurans that
have recent relatives that are typical of the Southeast Coastal
Plain are Scaphiopus cf. S. holbrooki, Hyla miofloridana, H. goini,
and Gastrophryne cf. G. carolinensis.

Western Gulf States.——The fossil Leptodactylus abavus is quite
similar to recent L. melanonotus which today ranges from southern
Sonora and San Luis Potosi southward along both Mexican coasts
to Costa Rica (Smith and Taylor, 1948). The genus Leptodacty-
lus is absent from Florida today, but species of this genus occur in
southern Texas and in a few islands of the West Indies. Again,
the possibility exists that the fossil Leptodactylus has Caribbean
affinities, but at present, I feel that this possibility is a slight one.

LITERATURE CITED

AUFFENBERG, W. 1956. Remarks on some Miocene anurans from Florida,
with a description of a new species of Hyla. Breviora, no. 52, pp. 1-11,
figs. 1-3.

CHANTELL, C. J. 1964. Some Mio-Pliocene hylids from the Valentine forma-
tion of Nebraska. American Midl. Nat., vol. 72, no. 1, pp. 211-225,
figs. 1-4.

. 1965. A lower Miocene Acris (Amphibia:Hylidae) from Colorado.
Jour. Paleont., vol. 39, no. 3, pp. 507-508, 1 fig.

. 1966. Late Cenozoic hylids from the Great Plains. Herpetologica,
vol. 22, no. 4, pp. 259-264.

Curapiiwy, P. S. 1956. Taxonomy and distribution of the spadefoot toads
of North America (Salientia:Pelobatidae). Master’s thesis, University
of Kansas, Department of Zoology, Lawrence, Kansas, May, 1956.

Estes, R. 1963. Early Miocene salamanders and lizards from Florida. Quart.
Jour. Florida Acad. Sci., vol. 26, pp. 234-256, figs. 1-4.

. 1964. Fossil vertebrates from the Cretaceous Lance formation of
eastern Wyoming. Univ. California Publ. Geol. Sci., vol. 49, pp. 1-180,
(omneseo) pls:

Hotman, J. A. 1961. A new hylid genus from the lower Miocene of Florida.
Copeia, 1961, pp. 354-355, 1 fig.

. 1962. A Texas Pleistocene herpetofuana. Copeia, 1962, pp. 255-
261, 1 fig.

140 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

. 1964. Pleistocene amphibians and reptiles from Texas. Herpetologica,
Vole 20NnOw2-sppauio-Gosiesupl=A

. 1965. Early Miocene anurans from Florida. Quart. Jour. Florida
Acad. Sci., vol. 28, no. 1, pp. 68-82, figs. 1-2.

Lyncu, J. D. 1962. An osteological character on the ilia of Acris crepitans
Baird, Acris gryllus Le Conte, and Hyla crucifer Wied. Copeia, 1962,
p. 434, figs. 1-2.

. 1965. The Pleistocene amphibians of Pit II, Arredondo, Florida.
Copeia, 1965, pp. 72-77, figs. 1-4.

. 1966. Additional treefrogs (Hylidae) from the North American
Pleistocene. Annals Carnegie Mus., vol. 38, art. 1, pp. 265-271, 1 fig.

ScuHMipT, K. P. 1953. <A check list of North American amphibians and rep-
tiles. Univ. Chicago Press, i-viii plus 280 pp.

SmiTH, H. M., anp E. H. Taytor. 1948. An annotated checklist and key to
the amphibia of Mexico. Bull. U. S. Nat. Mus., no. 194, i-vi plus 118 pp.

Taytor, E. H. 1942. Extinct toads and frogs from the upper Pliocene de-
posits Meade County, Kansas. Univ. Kansas Sci. Bull., vol. 28, no. 10,
pp. 199-235, 20 pls.

THEN, J. A. 1951. Anuran remains from the Miocene of Florida, with the
description of a new species of Bufo. Copeia, 1951, pp. 230-235, 2 pls.

. 1960. On Neoscaphiopus and other Pliocene pelobatid frogs. Copeia,
1960, pp. 89-94, 1 fig.

. 1962. A review of New World fossil bufonids. American Midl. Nat.,
vol. 68, pp. 1-50, 62 figs.

ZWEIFEL, R. G. 1956. Two pelobatid frogs from the Tertiary of North
America and their relationships to fossil and recent forms. American
Mus. Novitates, no. 1762, pp. 1-45, figs. 1-25.

Museum, Michigan State University, East Lansing, Michigan
48823

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

Variation in Bovine Cholinesterase Levels
C. J. Witcox anp H. H. Heap

One of the factors upon which the design of efficient experiments
is dependent is knowledge of the experimental error (in terms of
variance) to be expected. In experiments with large animals such
as the cow, the number of animals to be assigned, the duration of
the experiment and the number of samples to be taken become im-
portant because of the expense involved per animal. Our studies
of the effects of application of organic phosphorus insecticides
upon cholinesterase levels in the cow have provided us with data
from which estimates of variance of this physiologically important
trait can be obtained ( Wilcox and Head, 1966).

Of the esterases, cholinesterase is the most widely known and
among the more important. The enzymes which catalyze the hy-
drolysis of acetylcholine are known as cholinesterases and are found
in red blood cells (RBC) and plasma, as well as in most of the
body tissues. They are particularly active in nervous tissue. The
normal function of the true cholinesterases, such as acetylcholines-
terase, is to hydrolyze acetylcholine to acetic acid and choline
(Fukuto, 1957 and Oser, 1965).

The true cholinesterases are found principally in RBC and in the
central nervous system. These enzymes hydrolyze acetylcholine
more rapidly than they do other choline esters such as butyrylcho-
line and benzoylcholine. The normal function of these enzymes,
catalyzing the hydrolysis of acetylcholine, is extremely important
physiologically since acetylcholine is the transmitting agent of the
nerve impulse to effector cells in the neuromuscular junction and
in cholinergic synapses of the nervous system. The pseudocholin-
esterase enzymes are found principally in the blood plasma. Their
physological importance is not clear but they are classified as
such because they hydrolyze choline esters other than acetylcholine
more rapidly than they do acetylcholine (Gage, 1961 and Oser,
1965).

MetTHops AND RESULTS

Eight Jerseys were sampled eight times each during a 17-day
period, there being five 2-day and two 3-day intervals between the
eight samples. After collection, blood samples were heparinized
and refrigerated overnight. The following morning they were

142 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

shipped to the laboratory. Plasma was separated by centrifugation
and the chemical analysis performed on the two blood fractions.
The method of Michel (Oser, 1965) was used without modifica-
tion to determine the cholinesterase levels of plasma and RBC
hemolyzates. Acetylcholine in a standard buffer solution was used
as the substrate. The action of the enzyme on acetylcholine pro-
duces acetic acid which lowers the pH and is a measure of enzyme
activity expressed as ApH units per hour. Additional details of
the experimental material, sampling procedures, and method of
chemical analysis have been described previously (Wilcox and
Head, 1966). For this report, the data have been subjected to
standard statistical analyses, mainly analysis of variance and corre-
lation.

Mean cholinesterase levels, expressed as ApH units per hour,
were 0.411 and 0.123 for RBC and plasma, respectively; corres-
ponding sample (total) standard deviations were 0.072 and 0.037.
The value 0.411 was within the range for RBC reported by Radeleff
and Woodard (1956); they reported no significant cholinesterase
activity present in the plasma of cattle or sheep. Our data show
clearly that dairy cattle have considerably lower cholinesterase
levels than do men. The difference between plasma levels of the
two is especially dramatic. However, the biological meaning of
this is not known. Similarly, RBC cholinesterase levels of cattle
were only about 50 percent of those observed in man.

TABLE 1
Variance analysis of blood cholinesterase levels
RBC Plasma
Source d.f.b E(MS) MS V V% MS V V%

Animals o2+ 802A .013406** .001564 30 .000479 .000014 1
Days 1 on So2D 023199" 20027188 08 -008377 = .00nOOn 72
Error 49 o? 000892 000892 17 .000366 .000366 27
Total 63 .005244 100 .001381 100

a Units of measure: ApH units per hour.
b d.f.=degrees of freedom, E(MS)=—expectation of mean squares, MS=mean
squares, V—variance.

=P <0:01

Variability was partitioned as shown in Table 1. Error variance
represented a combination of sampling and measurement errors

Witcox AND Heap: Cholinesterase in Cattle 143

and animal by day interactions, since only single samples were taken
each day. Repeatabilities of measurements, as represented by the
oA
intraclass correlation, r=——— , were 0.30 and 0.1 for RBC
GhinGaD oro A & Gee ae
and plasma, respectively. Hence, sensitive characterization of dif-

ferences among animals was not accomplished by the procedure
used for plasma, but was for RBC.

The value of additional measurements can be estimated by the

change in the variance (gain in accuracy) of a mean based on
varying numbers of measurements with
V, I1*r(n-1)
a a ae where V= the variance, r=repeatability, and
n=the number of measurements represented in the mean. A gain
in accuracy in the characterization of an animal level, by taking
two measurements rather than one, of 49.5 percent would be ex-
pected for plasma and 35 percent for RBC. Measurements beyond
the second would result in gains with diminishing increments.

Day effects were of major importance, accounting for 53 per-
cent of the variability in RBC levels and 72 percent in plasma levels.
Further, comparisons among animals when determinations are
made on different days (or perhaps at different locations ) would
be subject to considerable error and would have limited value.
Doubtless any number of environmental and nutritional factors can
result in variation in cholinesterase levels.

Cholinesterase levels of RBC and plasma are essentially inde-
pendent of each other. Correlations obtained were as follows: for
total, 63 d.f., —.10; among animals, 7 d.f., +.04; among days, 7 d.f.,
—,.21; for error, 49 df., +.14. None of these were statistically
significant.

SUMMARY

Cholinesterase levels in RBC and plasma were determined in
eight samples from each of eight Jerseys over a 17 day period. Mean
levels, expressed as ApH units per hour, were 0.411 and 0.123 for
RBC and plasma, respectively; corresponding sample (total) stand-
ard deviations were 0.072 and 0.037. Animals accounted for 30
percent of the variability in RBC cholinesterase levels, days 53 per-
cent; corresponding percentages for plasma levels were 1 and 72.
Correlations between RBC and plasma levels were close to zero and
not significant.

144 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

ACKNOWLEDGMENTS

The authors wish to express appreciation to the Shell Chemical
Company for partial financial support, and to C. H. Van Middelem,
B. B. Byham, H. L. Somers, M. Kersey, and C. Sheffield for techni-
cal assistance. Chemical analyses were performed by Patterson and
Coleman Laboratories, Tampa, Florida.

LITERATURE CITED

Fuxuto, T. R. 1957. The chemistry and action of organic phosphorus insec-
ticides. Adv. in Pest Control Res., Interscience Publ., Inc., New York,
VO pps lA Ol:

Gacg, J. L. 1961. Residue determination of cholinesterase inhibition analysis.
Ady. in Pest Control Res., Interscience Publ., Inc., New York, vol. 4,
pp. 183-210.

OserR, B. L. (ed.) 1965. Hawk’s Physiological Chemistry, 1965. McGraw-
Hill, New York, 14th ed., pp. 1128-1132.

RADELEFF, R. D., AND G. T. Woopwarp. 1956. Cholinesterase activity of
normal blood of cattle and sheep. Vet. Med., vol. 51, pp. 512-514.

Witcox, C. J., anp Heap, H. H. 1966. Effect of organic phosphorus insecti-
cide application upon blood cholinesterase levels of dairy cattle. Florida
Agr. Exp. Sta. Dairy Sci. Mimeo Rpt. DY67-1, December 1, pp. 1-5.

Department of Dairy Science, University of Florida Institute of
Food and Agricultural Sciences, Gainesville, Florida. Florida Agri-
cultural Experiment Stations Journal Series No. 2629.

Quart. Jour. Florida Acad. Sci. 30(2) 1967 (1968)

FLORIDA ACADEMY OF SCIENCES

COUNCIL FOR 1967
President: JAcKson P. SICKELS
President Elect: CLARENCE C CLARK
Secretary: JoHN D. McCronE
Treasurer: JAMES B. FLEEK
Past President: Marcarer L. GILBERT
Past President: O. E. FRYE
Chairman, Charter and By-Laws Committee: ELMER C. PRICHARD
Finance Committee: JoHn S. Ross
Honors Committee: CLARENCE C CLARK
Quarterly Journal Committee: I. G. Foster
Talent Search Committee: ALFRED P. MILs
Program Committee: JAcKson P. SICKELS
Editor, Quarterly Journal: PrercE BropkKoRB
Chairman, Biological Sciences Section: RopErT W. LONG
Physical Sciences Section: RicHarp A. RHODES II
Social Sciences Section: GEORGE OsBORN
Medical Sciences Section: WiLL1AM K. BARTON
Science Teaching Section: ALBERT A. LATINA
Conservation Section: ELBERT A. ScHory, SR.
AAAS Council and Conference Representative: LAUREN GILMAN
State Coordinator of the Junior Academy: Louise V. AsH
Councilor at Large, Elected: R1icHARD GARRETT
Councilor at Large, Elected: James D. Ray, Jr.
Councilor at Large, Appointed: Earn Ricu
Councilor at Large, Appointed: Kerra HaNnsEN

ADDITIONAL COMMITTEE CHAIRMEN
Auditing: Joun S. Ross
Awards and Grants: PAUL VESTAL
Future Annual Meetings: MARGARET GILBERT
Local Arrangements: FRED E. CLARK
Membership: CLARENCE C CLARK
Resolutions: O. E. FRYE
Necrology: D. C. Swanson
Nominating: Grorce K. Rem
Visiting Scientist Program: CLARENCE C CLARK
News Letter: ALFRED P. MILLs

146 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MEMBERS OF THE ACADEMY

December 31, 1967

Sectional membership is indicated as follows: B, Biological
Sciences; C, Conservation; M, Medical Sciences; P, Physical Sci-
ences; S, Social Sciences. Members are requested to inform the
Editor of their sectional preferences.

Alexander, Taylor R., Dept. Botany, Univ. Miami, Coral Gables, Florida
S242 3

Allabough, Edwin D., Jr., 2111 Howard Drive, Orlando, Florida 32803 B

Allen, Ross, Ross Allen’s Reptile Inst., P. O. Box 367, Silver Springs, Florida
32688 B

Allen, Dr. Ted T., Jacksonville Univ., Jacksonville, Florida 32211 B

Almodovar, Dr. Luis R., Box 286, San German, Puerto Rico B

Alt, David, Dept. Geology, Univ. Montana, Missoula, Montana 59801 P

Anderson, William D., Jr., Dept. Biology, Univ. Chattanooga, Chattanooga,
Tennessee 37403 B

Andrews, Donald H., 21 S.E. Third St., Boca Raton, Florida 33432

Arnold, Dr. Luther A., 171 Norman Hall, Univ. Florida, Gainesville, Florida
32601 B

Ash, Mrs. Louise V., 2926 Swan Lane, Pensacola, Florida 32504 T

Ash, Dr. Willard O., 2926 Swan Lane, Pensacola, Florida 32504 P

Ashford, Dr. Theodore A., Univ. South Florida, Tampa, Florida 33620 B

Auffenberg, Walter, Florida State Museum, Gainesville, Florida 32601 B

Baker, Robert A., III, 2004 Romeria, Austin, Texas 78757

Banerjee, Shib Das, Roswell Park Mem. Inst., Cancer Prevention Research
Center, Orchard Park, New York 14127

Banks, Joseph E., 1411 Marion Ave., Tallahassee, Florida 32303

Barkuloo, James M., Fishery Biologist, Game and Fresh Water Fish Com.,
P. O. Box 576, Panama City, Florida 32402 B

Barton, Dr. Maurice A., 1900 Almeria Way S., St. Petersburg, Florida 33712
M

Barton, Dr. William K., 861 Sixth Ave. S., St. Petersburg, Florida 33701 M

Beck, William M., Jr., 1621 River Bluff Road, Jacksonville, Florida 32211 B

Becknell, Dr. G. G., 6900 Dixon Avenue, Tampa, Florida 33604 P

Beery, Dr. John R., Dean, School of Education, Univ. Miami, Coral Gables,
Florida 33124 §

Bellamy, R. E., Rural Route 1, Carrying Place, Ontario, Canada

Bellamy, Dr. Raymond F., 326 DeSoto, Tallahassee, Florida 32301 S

Beller, H. E., 743 duPont Bldg., Miami, Florida 33132

Bennett, Frank A., P. O. Box 3, Olustee, Florida 32072

Beohm, E. E., 324 W. 7th St., Jacksonville, Florida 32206

Membership List 147

Berne-Allen, Allan, 144 Washington Drive N., St. Armands Key, Sarasota,
Florida 33577

Betts, Joseph M., Jr., Topaz Home Apt. 224, 4400 East-West Highway,
Bethesda, Md. 20014

Biber, David, 725 S.W. 4th Ave., Gainesville, Florida 32601

Bieber, Theodore I., Dept. Chemistry, Florida Atlantic Univ., Boca Raton,
Florida 33432 P

Bird, L. C., 303 S. 6th Street, Richmond, Virginia 23219 B

Birdsey, M. R., Dept. Biology, Miami-Dade Junior College, Miami, Florida
3o156) ~B

Bissland, Howard R., 810 N.W. 19th Terrace, Gainesville, Florida 32601 C

Blanchard, Dr. Frank N., Dept. Geology, Univ. Florida, Gainesville, Florida
32601 P

Block, W. F., 2616 Fairway Ave., S., St. Petersburg, Florida 33700 P

Bodman, John W., 18 Wedgemere Rd., Winchester, Massachusetts 01890

Bonninghausen, Russel A., 313 E. Lakeshore Dr., Tallahassee, Florida 32303

Bonniwell, Bernard, Univ. Villanova, Villanova, Penn. 19085

Borders, Huey I., 1121 Citrus Isle, Fort Lauderdale, Florida 33315

Boss, Dr. Manley L., Florida Atlantic Univ., Boca Raton, Florida 33432 B

Boulware, Joe W., Univ. South Florida, Tampa, Florida 33620

Breder, Dr. C. M., Jr., R.F.D. 1, Box 452, Englewood, Florida 33533

Breen, Dr. Ruth S., 1806 Croydon Drive, Tallahassee, Florida 32303

Brey, Dr. Wallace S., Jr., 800 N.W. 37th Drive, Gainesville, Florida 32601 P

Brodkorb, Dr. Pierce, Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Brooker, Dr. Ralph H., Dept. Physics, University of South Florida, Tampa,
Florida 33620 P

Brooks, Dr. Karl M., Dept. Psychology, Florida A & M, Tallahassee, Florida
32307 S

Browder, James S., 1018 N.W. 10th Ave., Gainesville, Florida 32601

Brown, Charles P., 641 N.W. 34th Terrace, Gainesville, Florida 32601 B

Brown, Dr. Relis B., Dept. Biol. Sci.,° Florida State Univ., Tallahassee,
Florida 32306 B

Broyles, Dr. Arthur A., Dept. Physics, Univ. Florida, Gainesville, Florida
326 0leeee

Buckland, Charlotte B., c/o Mrs. Alexander W. Winkler, Spring Valley Rd.,
Woodbridge, Conn. 06525 B

Bullen, Mrs. Adelaide K., 103 Seagle Bldg., Gainesville, Florida 32601 B

Bullen, Ripley P., 103 Seagle Bldg., Gainesville, Florida 32601 B

Burgess, William D., Florida College, Tampa, Florida 33617 B

Buri, Dr. Peter, Div. Natural Sci., New College, Sarasota, Florida 33570 B

Burlingham, Kenneth, 5106 Memory Lane, El Paso, Texas 79932

Burns, Russell M., Box 900, Marianna, Florida 32446

Burton, R. H., 5121 S.W. 93rd Ave., Fort Lauderdale, Florida 33314

Byrd, I. B., U. S. Bureau of Commercial Fisheries, 144 Ist Ave. S.,
St. Petersburg, Florida 33701 B

Byron, C. W., 1034 Samar Rd., Cocoa Beach, Florida 32931

148 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Cain, Walter L., Box 12, Jacksonville Univ., Jacksonville, Fla. 32211

Caldwell, David K., Marineland Research Lab., Rt. 1, Box 122, St. Augustine,
Florida 32084 B

Calvin, D. Bailey, Univ. Miami School of Med., P. O. Box 875, Biscayne
Annex, Miami, Florida 33152

Camacho, Dr. E. Oliver, 740 N.E. 18lst St., North Miami Beach, Florida
33162

Campbell, James A., 2707 Bronte Ave., Nashville, Tenn. 37216 B

Carlisle, Dr. Victor W., 223 McCarty Hall, Univ. Florida, Gainesville, Florida
32601 P

Carmichael, Billy E., Chipola Jr. College, Marianna, Florida 32446

Carr, Dr. A. F., Jr., 14 Flint Hall, Univ. Florida, Gainesville, Florida
32601 B

Carr, Joseph A., Jr., 3402 Riverview Drive, Tampa, Florida 33604 P

Carr, Dr. T. D., Dept. Physics, Univ. Florida, Gainesville, Florida 32601 P

Carroll, Dr. Edward J., 8201 Ponce de Leon Road, Miami, Florida 33143

Carter, William J., P. O. Box 247, Yankeetown, Florida 32698

Chalmers, Dr. E. L., Jr., College Arts & Sci., Florida State Univ., Tallahassee,
Florida 32306

Chamelin, Dr. I. M., 615 Phelps Ave., Winter Park, Florida 32789 B

Chapman, H. L., Range Cattle Experiment Station, Ona, Florida 33865 B

Chen, Dr. Kwan-Yu, Dept. Physics-Astronomy, Univ. Florida, Gainesville,
Florida 32601 P

Chernetski, Dr. Kent E., Dept. Zoology, Univ. Florida, Gainesville, Florida
326019 2

Chew, Victor, 380 Greenway Ave., Satellite Beach, Florida 32935

Christensen, Robert F., 19435 N.W. 24th Ave., Opa Locka, Florida

Clapper, Russell B., 2072 Braman St., Fort Myers, Florida 33901

Clark, Dr. Clarence C., Univ. South Florida, Tampa, Florida 33620 P

Clark, Fred E., Dept. Biology, Steston Univ., DeLand, Florida 32720 B

Clark, Samuel F., Dept. Chemistry, Florida Atlantic Univ., Boca Raton,
Florida 33432 P

Clarke, Walter V., 1195 S.E. 17th St., Fort Lauderdale, Florida 33316

Clary, J. D., 1608 Meadowbrook Ave., Lakeland, Florida 33803

Claus, Dr. Edward P., Ferris State College, Big Rapids, Michigan 49307

Clugston, James P., Bur. Sport Fisheries & Wildlife, Div. Federal Aid,
Peachtree-Seventh Bldg., Atlanta, Georgia 30323 B

Collier, Albert W., 1622 Seminole Dr., Tallahassee, Florida 32301

Conard, Dr. Henry S., Lake Hamilton, Florida 33851 P

Craighead, Frank C., Box 825, Homestead, Florida 33030 B

Crawford, Mrs. Eleanor L., P. O. Box 16, Jacksonville Univ., Jacksonville,
Florida 32211

Creager, Don B., Prof. Biology, Jacksonville Univ., Jacksonville, Florida
SPPALIL 183

Crossman, Roy A., Jr., 106 4th St., Jar-Phyl Village, Winter Haven, Florida
33880

Membership List 149

Crouch, Dr. G. E., Jr., Physics Dept., Florida State University, Tallahassee,
Florida 32306 P

Culbertson, Jon R., Div. Nat. Sci., New College, Sarasota, Fla. 33578

Cunha, Dr. T. J., 252 McCarty Hall, Univ. Florida, Gainesville, Florida
32601 B

Dambaugh, Dr. Luella N., Univ. Miami, Coral Gables 33124 S

Dana, Allan H., 6606 S.W. 60th St., South Miami, Florida 33143

Davis, Fanny-Fern, P. O. Box 475, Valparaiso, Florida 32580

Davis, Dr. George K., Director Biological Sci., Gainesville, Fla. 32601 B

Davis, Gordon, 10545 N.W. 28th Ave., Miami, Florida 33147

Davis, Jefferson C., Jr., Dept. Chemistry, Univ. South Florida, Tampa, Florida
33620 P

Davis, Robert H., Dept. Physics, Florida State Univ., Tallahassee, Florida
32306 P

Davis, Robert L., 11525 82nd Avenue N., Largo, Florida 33540

Dauer, Dr. Maxwell, Dept. Radiology, Univ. Miami, School of Medicine,
Miami, Florida 33136 M

Deichmann, Dr. Wm. B., School of Medicine, Univ. Miami, Coral Gables,
Florida 33134 M

Denman, Dr. Sidney B., College Med., Univ. Florida, Gainesville, Florida
32601 S

Dequine, John F., Southern Fish Culturists, Box 251, Leesburg, Florida
32748 B

Derr, Dr. Vernon E., 3410 Everett Drive, Boulder, Colorado 80302 P

Dew, Dr. Robert J., Jr., P. O. Box 18243, Tampa, Florida 33609 P

Dickinson, Dr. J. C., Jr., Florida State Museum, Gainesville, Florida 32601 B

Dijkman, Dr. Marinus J., 6767 S.W. 112th St., Miami, Florida 33156 B

Dobkin, Dr. Sheldon, Dept. Biological Sciences, Florida Atlantic University,
Boca Raton, Florida 33432 B

Doyle, Dr. Laura M., 365 Hawthorne Avenue, Palo Alto, California 94301

Driver, Paul J., 1347 West 9th St., Jacksonville, Florida 32209

Drysdale, Taylor, 5526 Parkdale Drive, Orlando, Florida 32809

Dubbeldam, Dr. Pieter S., 506 Magnolia Ave., Melbourne Beach, Florida
32951

Dudley, Frank M., Div. Physical Sci., Univ. South Florida, Tampa, Florida
33620 P

DuMond, Frank V., P. O. Box 246, Goulds, Florida 33170

Dunathan, Jay P., 452 86th Ave. N., St. Petersburg, Florida 33702

Dunning, Wilhelmina F., U. Miami Cancer Res. Lab., Box 8215, Coral
Gables, Florida 33124

Dyrenforth, Dr. L. Y., 3885 St. Johns Ave., Jacksonville, Florida 32205 M

Eoff, K. M. 450 Little Hall, U. Florida, Gainesville, Florida 32601
Edwards, Dr. Richard A., Dept. Geology, Univ. Florida, Gainesville, Florida
S26018 2

150 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Emmerton, Ernest E., 213 N.W. 34th Terr., Gainesville, Florida 32601

Ehrhart, Llewellyn, Research Park Bldg. Cornell Univ., Ithaca, N. Y. 14850
Evans, Dr. Charles A., 208 S. Renellie Dr., Tampa, Florida 33609

Evans, Dr. Elwyn, 500 E. Colonial Dr., Orlando, Florida 32803 M

Evers, Mrs. Rebecca Carr, 2110 N.W. 204th Street, Opa-Locka, Florida 33054
Ewing, Upton C., 362 Minorca Ave., Coral Gables, Florida 33134

Faber, John L., 1242 N. Palm Ave., Sarasota, Florida 33577

Faloona, Res. Div., V. A. Hosp., 4500 Lancaster, Dallas, Texas 75216

Faust, Burton, 2178 Harbor View Dr., Dunedin, Florida 33528

Ferguson, Cecil R., P. O. Box 128, Fort Pierce, Florida 33450

Ferguson, Dr. John C., Dept. Biology, Florida Presbyterian College, St.
Petersburg, Florida 33733 B

Fernandez, Dr. Jack E., Univ. South Florida, Tampa, Florida 33620

Field, Dr. Henry, 3551 Main Highway, Coconut Grove, Florida 33133 S$

Finucane, John H., 1320 58th St. S., Gulfport, Florida 33707 B

Fleek, Dr. James B., 518 Patricia Lane, Jacksonville Beach, Florida 32050 P

Fly, Prof. Lillian, 4060 Battersea Road, Coconut Grove, Florida 33133 B

Foote, Dr. Perry A., 525 N.E. 9th Ave., Gainesville, Florida 32601 P

Foraker, Dr. Alvan G., 800 Miami Road, Jacksonville, Florida 32207 M

Ford, Dr. Ernest S., Dept. Botany, Univ. Florida, Gainesville, Florida
32601 B

Forman, Guy, Dept. Physics, Univ. South Florida, Tampa, Florida 33620 P

Foster, Dr. I. G., Florida Presbyterian College, St. Petersburg, Florida 33733
P |

Foster, Dr. Virginia, Box 87, Pensacola College, Pensacola, Florida 32504

Foster-Quick, Mrs. Joyce, P. O. Box 464, Jensen Beach, Florida 33457

Fox, Dr. S. W., Institute of Molecular Evolution, Univ. Miami, 521 Anastasia,
Coral Gables, Florida 33134 B

Foxman, David A., 1521 Garcia Ave., Coral Gables, Florida 33146

Friedland, Bernard, 1010 Manati Ave., Coral Gables, Florida 33146

Frye, Dr. O. E., Jr., Game and Fresh Water Fish, Comm., Tallahassee,
Florida 32304 B

Fuller, Dorothy L., P. O. Box 418, DeLand, Florida 32720 B

Gardner, Elizabeth Ann, 1000 Spring Garden Rd., Apt. 13, Miami, Florida
33136 M

Garrard, Dr. Leon A., 1617 N.W. 10th Terr., Gainesville, Florida 32601 B

Garrett, James R., 245 Nicholson Dr., Moorestown, New Jersey 08057

Garrett, Richard E., Dept. Physics and Astronomy, Univ. Florida, Gainesville,
Florida 32601 P

Gathman, C. A., 501 21st Avenue N., Lake Worth, Florida 33460 B

Gilbert, Dr. Margaret, Dept. Biology, Florida Southern College, Lakeland,
Florida B

Gilman, L. C., Dept. Biology, Univ. Miami, Coral Gables, Florida 33124 B

Girard, Murray, 5900 Devonshire Blvd., Miami, Florida 33155 B

Goin, Dr. Coleman J., Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Membership List 151

Golightly, Jacob F., Jacksonville Univ., Jacksonville, Florida 32211

Gordon, Thomas E., Jr., D.D.S., 2517-A E. Colonial Drive, Orlando, Florida
32803 M

Gramling, Dr. L. G., Coll. Pharmacy, Univ. Florida, Gainesville, Florida
32601 M

Gray, Oscar S., Gray Industries, Inc., 948 N.W. 44th Street, Fort Lauderdale,
Florida 33309

Green, Alex E. S., 2900 N.W. 14th Pl., Gainesville, Florida 32601

Gresham, W. B., Jr., P. O. Box 18525, Tampa, Florida 33609

Gut, H. J., P. O. Box 700, Sanford, Florida 32771 B

Haines, Charles E., Amer. Con. Gen-Recife IRI, APO New York, N. Y.
09676

Hall, Everette E., Jr., P. O. Box 12976, Univ. Station, Gainesville, Florida
32601 M

Hanley, James R., Jr., 6446 Anvers Blvd., Jacksonville, Florida 32210

Hansen, Dr. Keith L., Biology Dept., Stetson Univ., DeLand, Florida 32720 B

Hansel, Paul G., 1374 Monterey Blvd., St. Petersburg, Florida 33704

Harlow, Richard, 31 Colonial Pl., Ashville, North Carolina 28804

Harms, Dr. R. H., Dept. Poultry Sci., Univ. Florida, Gainesville, Florida
32601 B

Harrington, Dr. Robert W., Jr., Florida State Board of Health, P. O. Box 308,
Vero Beach, Florida 32960 B

Hatala, Dr. Robert J., 2167 Vivian Way S., St. Petersburg, Florida 33712

Harris, Herbert H., 610 Ampton Ave., Tallahassee, Florida 32304

Hausman, Daniel W., 615 Paul Russell Road, Tallahassee, Florida 32301

Heemstra, Phillip C., Box 70, Univ. Miami, Inst. Marine Sci., 1 Rickenbacker
Causeway, Miami, Florida 33149 B

Heinrich, Dr. Edwin P., Orange Park, Florida 32073

Hellwege, Dr. Herbert, Rollins College, Winter Park, Florida 32789 P

Hentges, Dr. James F., Jr., 253 McCarty Hall, Univ. Florida, Gainesville,
Florida 32601 B

Hetrick, Daniel L., 7420 S.W. 19 Terr., Miami, Florida 33155

Heuser, Dr. Gustave F., 608 Hillside Dr., Lakeland, Florida 33803

Highsmith, John Lewis, St. Johns River Jr. College, Palatka, Florida 32077

Hill, William P., Studio 9, 275 W. Magnolia Ave., Merrit Is., Florida 32952

Hilmon, J. B., P. O. Box 2570, Asheville, North Carolina 28802

Hobbs, Dr. Horton H., Jr., U. S. National Museum, Washington, D. C.
20560 B

Holman, Dr. J. Alan, The Museum, Michigan State Univ., East Lansing,
Michigan 48832 B

Hopkins, E. F., 1207 Biercliff Drive, Orlando, Florida 32806

Hopkins, Dr. Thomas S., Div. Nat. Sci., New College, Sarasota, Florida
33578

Houser, James G., Tech. Director, Martin Co., P. O. Box 5837, Orlando,
Florida 32805 P

152 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Howard, Fred E., Jr., 306 Gardner Dr., N. E., Ft. Walton Beach, Florida
32548

Hsia, S. L., Dept. Dermat., Univ. Miami School of Medicine, 1600 N.W.
10th Ave., Miami, Florida 33136 M

Hubbell, Dr. David S., 861 6th Ave. S., St. Petersburg, Florida 33101

Hubbs, Dr. Carl L., Scripps Institution of Oceanography, La Jolla, California
92037 B

Hughes, Dr. William E., Box 1337, Stetson Univ., DeLand, Florida 32720

Hull, Robert W., Dept. Biol. Sci., Florida State Univ., Tallahassee, Florida
32306 B

Hunt, Burton P., Dept. Biology, Univ. Miami, Coral Gables, Florida 33124 B

Hunter, Dr. George W., III, Dept. Microbiology, Univ. Florida, Gainesville,
Florida 32601 B

Idyll Dr. C. P., Inst. Marine Sci., Univ. Miami, Miami, Florida 33149

Ingle, Robert M., Florida State Board of Conservation, Tallahassee, Florida
S2o0I1— 1B

Ivey, Dr. Marvin L., Dept. Natural Sciences, St. Petersburg Junior College,
P. O. Box 13489, St. Petersburg, Florida 33733

Jenkins, George L., Dept. Physics, Stetson Univ., DeLand, Florida 32720 P

Jerris, S. R., Roehr Products Co., Inc., P. O. Box 880—2010 New Daytona
Rd., DeLand, Florida 32720

Jodrey, Louise H., 1200 S. Ocean Blvd., Apt. 6-H, Boca Raton, Florida
33432

Jones, Dr. E. Ruffin, Jr., Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Joyce, Edwin A., Jr., 1943 Mound Place, S., St. Petersburg, Florida 33712

Kane, Howard L., 1264 Cleburne Drive, Fort Myers, Florida 33901

Kaplan, Sherman R., 333 41st St., Miami Beach, Florida 33140

Katz, Edith E., R. D. 2, Box 788 E, DeLand, Florida 32720

Kendall, Harry W., Dept. Physics, Univ. South Florida, Tampa, Florida
33620 P

Kerman, Herbert D., Halifax Dist. Hosp., Daytona Beach, Florida 32015 M

Keuper, Dr. Jerome P., Florida Inst. Tech., P. O. Box 1150, Melbourne,
Florida 32901 P

Kilby, Dr. John D., Dept. Biology, Univ. Florida, Gainesville, Florida 32601

Kilgore, Dr. Charles F., Florida Jr. College, Jacksonville, Florida 32207

Kinser, B. M., P. O. Box 158, Eustis, Florida 32726 P

Kirk, Dr. W. G., Range Cattle Exp. Sta., Ona, Florida 33865

Knowles, Robert P., 2101 N.W. 25th Ave., Miami, Florida 33142 M

Koenig, Dr. Duane, Dept. Hist., Univ. Miami, Coral Gables, Fla. 33124

Koger, Dr. Marvin, Dept. Animal Science, Univ. Florida, Gainesville, Florida
32601 B

Kolipinski, Dr. Milton C., U. S. Geological Survey, 51 Southwest First Ave.,
Miami, Florida 33130 P

Membership List 153

Krivanek, Dr. Jerome O., Univ. South Florida, Tampa, Florida 33620 B
Kromhout, Robert A., Physics Dept., Florida State Univ., Tallahassee, Florida
32306

LaCava, Dr. Frederick W., 264 South Atlantic Ave., Ormond Beach,
Florida 32074

Lacy, Dr. Burritt S., Box 267, Keene, New York 12942

Lackey, Dr. James B., Box 36, Melrose, Florida 32666 B

Laessle, Dr. Albert M., Dept. Zoology, Univ. Florida, Gainesville, Florida

32601 B

Larson, Dr. Edward, Dept. Biology, Univ. Miami, Coral Gables, Florida
33124 B

Latina, Albert A., 311A Science Bldg., Univ. South Florida, Tampa, Florida
33620 B

Lawrence, Dr. John M., Dept. Zoology, Univ. South Florida, Tampa, Florida
33620 B

Laxson, D. D., 231 41st St., Hialeah, Florida 33012

Layne, Dr. James N., Archbold Bio. Sta., Rt. 2, Box 380, Lake Placid,
Florida 33852 B

Leavitt, Dr. Benjamin B., Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Lee, Mrs. Annette J., P. O. Box 1233, Lake City, Florida 32055

Lee, Clarence E., 2833 N.E. 26th Ave., Lighthouse Point, Pompano Beach,
Florida 33064

Leigh, Dr. W. Henry, Dept. Zoology, Univ. Miami, Coral Gables, Florida
33124 B

Leto, Frank P., Jr., 4713 Leila Ave., Tampa, Florida 33616 T

Lilly, Dr. John C., 3430 Main Highway, Miami, Florida 33133

Little, Dr. William Asa, P. O. Box 875, Biscayne Annex, Univ. Miami,
Miami, Florida 33152 M

Long, Dr. William Hendren, Dept. Meteorology, Florida State Univ., Talla-
hassee, Florida 32306 P

Long, Dr. Robert W., Dept. Botany & Bact., Univ. South Florida, Tampa,
Florida 33620

Lorz, Dr. Albert, 409 Newell Hall, Univ. Florida, Gainesville, Florida
32601 B

Lovell, William V., Route 2, Box 18, Sanford, Florida 32771 P

Lutz, Nancy E., Box 114, Mandarin, Florida 32064

Lyons, William G., Florida Board of Conserv. Marine Lab., P. O. Drawer F,
St. Petersburg, Florida 33731

McBee, Ethelyne L., 4191 S.W. 97th Ct., Miami, Florida 33165

McClure, J. S., Jacksonville Univ., Jacksonville, Florida 32211 P

McCrone, Dr. John D., Dept. Zoology, Univ. Florida, Gainesville, Florida
32601

McDarment, Corley P., Route 1, Box 205, Eau Gallie, Florida 32935

Macgowan, Prof. Robert, Florida Southern College, Lakeland, Florida 33803 S

154 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

McKenzie, Dr. Doris, Dept. Med., Univ. Miami, Med. School, Miami, Florida
33136
McMahan, Mary Ruth, 6070 6th Ave., North, St. Petersburg, Florida 33710

Mann, Eloise Ann, 5748 Merrill Rd., Jacksonville, Florida 32211

Marks, Dr. Meyer B., 333 41st St., Miami Beach, Florida 33140 B

Martz, Roger A., 3994 S.W. 12th Terr., Ft. Lauderdale, Florida 33315

Masters, Dale R., Gulf Coast Jr. College, Panama City, Florida 32401

Maxwell, Lewis S., 6230 Travis Blvd., Tampa, Florida 33610

Meck, Mervin E., M. D., 225 N. Causeway, New Smyrna Beach, Florida
32069 M

Meisel, Max, 4444 Post Ave., Miami Beach, Florida 33140

Menzel, Dr. R. W., Oceanographic Institute, Florida State Univ., Tallahassee,
Florida 32306 B

Metcalf, Zubie W., Jr., 1601 Hernando Dr., Tallahassee, Florida 32304

Miles, E. P., Computing Center, Florida State Univ., Tallahassee, Florida
32306 P

Miller, Dr. E. Morton, 1212 Manati Ave., Coral Gables, Florida 33146

Miller, Pat H., Box 279, Everglades Nat’l. Park, Homestead, Florida 33030

Miller, J. E., Florida Inst. Tech., P. O. Box 1150, Melbourne, Florida 32901

Mills, Alfred P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida
Cola ae

Miner, Dr. Helen I., 770 Lake Road, Bay Point, Miami, Florida 33137

Minto, Wallace L., 1242 North Palm Ave., Sarasota, Florida 33577

Moe, Martin A., Jr., Board of Conservation, Marine Lab., P. O. Drawer F,
St. Petersburg, Florida 33731 B

Montgomery, Harley A., 1700 N.E. 105th St., Miami Shores, Florida 33138

Montgomery, Joseph G., Dept. Biology, Manatee Jr. College, Bradenton,
Florida 33505 B

Moody, Harold L., Game & Freshwater Fish Comm., 545 N. Woodland,
Winter Garden, Florida 32787

Moore, McDonald, 562 Leamore Ct., Mobile, Alabama 36617

Morrison, Thomas F., 1491 Summerland Ave., Winter Park, Florida 32789

Morton, Mrs. Julia F., Univ. Miami, Box 8204, Coral Gables, Florida 33124

Morton, Richard K., Jacksonville Univ., Contract Branch, Jacksonville, Florida
SpPAnL &

Moya, Frank, Dept. Anesthesiology, Jackson Memorial Hosp., Miami, Florida

33136 M

Mulson, Joseph F., Dept. Physics, Rollins College, Winter Park, Florida

32789 P

Murray, Mary Ruth, 1326 S.W. Ist St., Miami, Florida 33135 S

Mustian, William R., Jr., 529 Bonnie Drive, Lakeland, Florida 33803

Nagelsen, Mrs. J. C., Orlando Jr. College, 901 Highland Ave., Orlando,
Florida 32803

Nation, Dr. James L., 537 N.W. 35th Terrace, Gainesville, Florida 32601

Neithamer, Richard W., Florida Presbyterian College, St. Petersburg, Florida
8388

Membership List 155

Nelson, E. Lee, Florida Jr. College at Jacksonville, Cumberland Campus,
Jacksonville, Florida 32205

Nelson, Dr. Gid E., Jr., Univ. South Florida, Tampa, Florida 33620 B

Nichols, Cecil B., Miami Dade Jr. College, N. Campus, 11380 N.W. 27th
Ave., Miami, Florida 33167

Niven, Dr. Jorma I., 1803 E. Lakeview Ave., Pensacola, Florida 32503

Noble, Dr. Nancy L., 1550 N.W. 10th Ave., Miami, Florida 33136

Nordlie, Dr. Frank G., Dept. Zoology, Univ. Florida, Gainesville, Florida
32601 B

Ober, Lewis D., 1235 N.E. 204th St., North Miami Beach, Florida 33162 B

O’Brien, Robert E., P. O. Box 39, Rollins College, Winter Park, Florida
32789

O’Hara, Dr. James, Dept. Biology, Central State College, Wilberforce, Ohio
45384

Orgell, Dr. Wallace H., 19240 Christmas Rd., Miami, Florida 33157

Osborn, Dr. George, Little Hall 487, Univ. Florida, Gainesville, Florida 32601

Owens, Clarence B., Box 8, Florida A & M Univ., Tallahassee, Florida
323000) S

Palmer, Dr. A. Z., Dept. Animal Husbandry & Nutrition, Univ. Florida,
Gainesville, Florida 32601 B

Pan, Huo-Ping, U. S. Dept. Interior, 3700 E. University Ave., Gainesville,
Florida 32601

Park, Dr. Mary Cathryne, 450 Norwood St., Merrit Is., Florida 32952 S

Paul, John R., Illinois State Museum, Springfield, Ill. 62706

Paulson, Dr. Dennis R., Dept. Zoology, Univ. Washington, Seattle, Wash-
ington 98105

Peaslee, Dr. Margaret H., Dept. Biology, Florida Southern College, Lake-
land, Florida 33801

Penner, Dr. Lawrence R., Dept. Biol. Sci. Syst. & Environ. U-42, Univ.
Connecticut, Storrs, Connecticut 06268 B

Phillippy, Clayton, 2202 Lakeland Hills Blvd., Lakeland, Florida 33801

Phipps, Dr. Cecil G., 1245 E. 8th St., Cookeville, Tennessee 38501 P

Pierce, Dr. E. Lowe, Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Pirkle, Dr. E. C., 2219 N.W. 17th Ave., Gainesville, Florida 32601 P

Plendl, Dr. Hans S., Dept. Physics, Florida State Univ., Tallahassee, Florida
32306 P

Plyler, Earle K., Dept. Physics, Florida State Univ., Tallahassee, Florida
32306 P

Poitras, Dr. Adrian W., Div. Natural Sci., Miami-Dade Jr. College, 11380
N.W. 27th Ave., Miami, Florida 33167

Polskin, Dr. Louis J., 1401 S. Florida Ave., Lakeland, Florida 33803 M

Powell, Dr. Howard B., Dept. Chemistry, Univ. Miami, Coral Gables, Florida
Sole 12

156 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Powell, Dr. Leon W., Jr., Martin Mem. Hosp., Stuart, Florida 33494

Prichard, Elmer C., 605 N. Amelia, DeLand, Florida 32720 B

Prichett, Dr. Cavett O., 1 Davis Blvd., Suite 307, Tampa, Florida 33606

Provost, Dr. Maurice W., Box 308 Vero Beach, Florida 32960 B

Puryear, Dr. R. W., Florida Normal and Industrial Memorial College, St.
Augustine, Florida S

Rader, Dr. Luke, 7900 S.W. 104th St., Miami, Florida 33156

Rappenecker, Dr. Caspar, Dept. Geology, Univ. Florida, Gainesville, Florida
32601 P

Ray, Dr. James D., Jr., Univ. South Florida, Tampa, Florida 33620 B

Rehburg, Weldon Reed, 3820 2nd Ave. N., St. Petersburg, Florida 33733

Reid, Dr. George K., Dept. Biology, Florida Presbyterian College, St. Peters-
burg, Florida 33733 B

Rhodes, Richard A., II, 205 N.W. Monroe Circle, N., St. Petersburg, Florida
33702

Rice, Dr. Laurence B., 43 W. Bay Drive, Cocoa Beach, Florida 32931

Rich, Earl R., Dept. Zoology, Univ. Miami, Coral Gables, Florida 33124

Riemer, Mrs. Florence R., 9130 S.W. 100th St., Miami, Florida 33156

Rippy, W. C., Jr., 2525 Sunset Dr., Tampa, Florida 33609

Rivas, Luis Rene, Dept. Zoology, Univ. Miami, Coral Gables, Florida 33124
B

Roberts, Dr. Leonidas H., Dept. Physics & Astronomy, Univ. Florida, Gaines-
ville, Florida 32601 P

Robins, Dr. C. Richard, Institute Marine Science, Univ. Miami, 1 Ricken-
backer Causeway, Va. Key, Miami, Florida 33149 B

Roess, William B., Florida Presbyterian College, P. O. Box 12560, St.
Petersburg, Florida 33733

Rolfs, Herman E., 5513 Merrick Drive, Univ. Health Center, Coral Gables,
Florida 33134 M

Rooks, Herman, Gulf Coast Jr. College, Panama City, Florida 32401

Ross, Arnold, Dept. of Invert. Paleo., Nat. Hist. Mus., Balboa Park, Box
1390, San Diego, California 92112 P

Ross, Dr. John S., Dept. Physics, Rollins College, Winter Park, Florida
SSS) 12

Ryser, Dr. Phillip W., 1315 N.E. 4th Ave., Fort Lauderdale, Florida 33304

Sachs, K. N., Jr., U. S. Geological Survey, E-214 U. S. Nat. Museum,
Washington 25, D. C. 20242 P

Sall, Walter G., 605 Lincoln Road, Miami Beach, Florida 33139

Sanders, Dr. Murray, Gray Industries, Inc., 948 N. W. 44th St., Fort Lauder-
dale, Florida 33309 M

Sandstrom, Carl J., Dept. Biology, Rollins College, Box 53, Winter Park,
Florida 32789 B

Saslaw, Dr. Milton S., 1350 N.W. 14th St., Miami, Florida 33125 M

Sauer, Dr. E. G. Franz, Dept. Zoology, Univ. Florida, Gainesville, Florida
32601

Membership List 157

Sawyer, Dr. Earl M., Dept. Physics, Univ. Florida, Gainesville, Florida
S260) ve

Schneider, George H., 2292 S.W. 36th Ave., Miami, Florida 33145

Schory, Elbert A., Sr., P. O. Box 1468, Fort Myers, Florida 33902

Schrader, H. W., 128B Williamson Hall, Univ. Florida, Gainesville, Florida
32601 P

Schricker, John Adams, 4565 Duhme Rd., Apt. 203, St. Petersburg, Florida
33708

Schultz, Dr. Harry P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida
Sol Agee

Schwartz, Dr. Albert 10000 S.W. 84th St., Miami, Florida 33143 B

Schwarz, Guenter, Dept. Physics, Florida State Univ., Tallahassee, Florida
32306 P

Scolaro, Reginald J., Div. Invert. Paleo., Smithsonian Inst., U. S. Nat. Mus.,
Washington, D. C. 20560

Scott, Bruce Von G., 6201 Chapman Field Drive, Miami, Florida 33156

Seaman, Dr. Irvin, 470 Biltmore Way, Coral Gables, Florida 33134 M

Serfass, Dr. Earl J., Milton Roy Co., P. O. Box 12169, St. Petersburg, Florida
BOTS.)

Shanor, Leland, Dept. Botany, Univ. Florida, Gainesville, Florida 32601 B

Shepard, Dr. Weldon O., U. S. Forest Service, P. O. Box F, Ft. Myers,
Florida 33902

Sherman, Dr. H. B., 410 Howry Ave., DeLand, Florida 32720 B

Shirley, Dr. Ray L., Nutrition Lab., Univ. Florida, Gainesville, Florida 32601
B

Shor, Prof. Bernice C., Rollins College, Winter Park, Florida 32789

Sickels, Dr. Jackson P., 541 San Esteban Ave., Coral Gables, Florida 33146 P

Simon, Dr. Joseph L., Dept. Zoology, Univ. South Florida, Tampa, Florida
33620

Sims, Harold W., Jr., 7542—18th Ave. N., St. Petersburg, Florida 33710 B

Slack, Dr. Francis, P. O. Box 818, Hobe Sound, Florida 33455 M

Smith, Dr. Alex G., Dept. Physics and Astronomy, Univ. Florida, Gainesville,
Florida 32601 P

Smith, Earl D., 2309 Coventry Ave., Lakeland, Florida 33803 P

Smith, Dr. Fredrick B., Dept. Soils, Univ. Florida, Gainesville, Florida
32601 B

Smith, Marshall E., 418 W. Platt St., Tampa, Florida 33606 M

Smith Cmdr. Nathan L., 631 N.W. 34th Dr., Gainesville, Florida 32601 P

Soldo, Dr. Anthony T., V.A. Hosp., 1200 Anastasia Ave., Coral Gables,
Florida 33134 M

Sokoloff, Dr. Boris Th., Dept. Biology, Florida Southern College, Lakeland,
Florida 33803 B

Soule, Dr. James, 104 McCarty, Univ. Florida, Gainesville, Florida 32601 B

Starn, Charles H., 740 N.W. 65th Ave., Fort Lauderdale, Florida 33313

Steidinger, Karen A., P. O. Drawer F, St. Petersburg, Florida 33731

Stelz, William, 3129 Glenwood Ct., St. Ann, Missouri 63074
Stevens, Marion, 3924 Cleveland St., Hollywood, Florida 33021 B

158 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Stevenson, Dr. Henry M., Dept. Biological Sciences, Florida State University,
Tallahassee, Florida 32306 B

Stewart, Mrs. Violet N., Box 744, Crystal River, Florida 32629

Stubbs, Sidney A., P. O. Box 2066, Houston, Texas 770152

Swanson, Dr. D. C., Dept. Physics, Univ. Florida, Gainesville, Florida 32601
P

Sweat, Donald E., Marine Lab., P. O. Box 1207, Key West, Florida 33040

Sweigert, Ray L., 2761 E. Vina Del Mar Blvd., St. Petersburg Beach, Florida
33706

Swift, Camm C., Dept. Biol. Sci., Florida State Univ., Tallahassee, Florida
32306 B

Swindel, David E., Jr., 905 E. Park Ave., Tallahassee, Florida 32301 B

Taft, Dr. William H., Univ. South Florida, Tampa, Florida 33620

Tanner, W. Lee, Box 38, Lake Panasoftkee, Florida 33538

Tanner, William F., Dept. Geology, Florida State Univ., Tallahassee, Florida
eyo OLS) Je

Taylor, James R., 10922 52nd Ave. North, St. Petersburg, Florida 33708 B

Taylor, John L., 527 New York Ave., Dunedin, Florida 33528 B

Tebeau, C. W., Univ. Miami, 307 Aledo Ave., Coral Gables 33134 §S

Teas, Dr. Howard J., 6700 S.W. 130th Terr., Miami, Florida 33156

Thomas, Dr. Dan A., Dean of the Faculty, Jacksonville Univ., Jacksonville,
Florida 32211 P

Thomas, Dr. Garland L., Physics Dept., Florida Inst. Tech., Melbourne,
Florida 32901

Thomas, Richard, 10000 S.W. 84th St., Miami, Florida 33143 B

Ting, Dr. S. V., Citrus Experiment Station, P. O. Box 1088, Lake Alfred,
Florida 33850 B

Tinner, J. C., 1305 N. Montford Avenue, Baltimore, Maryland 21213 P

Tissot, Dr. A. N., Agricultural Exp. Sta., Univ. Florida, Gainesville, Florida
32601 B

Tocci, Dr. Paul M., P. O. Box 875, Biscayne Annex, Miami, Florida 33152

Totten, Henry R., Dept. Botany, Univ. North Carolina, Box 247, Chapel Hill,
North Carolina 27514 B

Toulmin, Dr. Lymon D., Dept. Geology, Florida State Univ., Tallahassee,
Florida 32306 P

Tricaro, Robert, 5890 S.W. 79th Ct., South Miami, Florida 33143

Tyrone, Victor, Box 1438, St. Petersburg, Florida 33731

Van Vleck, David B., Dept. Zoology, Univ. Miami, Coral Gables, Florida
BOISE 1B}

Van Wagtendonk, Dr. W. J., Research-Veterans Admin. Hosp., Coral Gables,
Florida 33134 M

Vance, Dr. Jimie A., 9885 N. Kendall Dr., Miami, Florida 33156

Vernon, Dr. Robert O., Box 631, Tallahassee, Florida 32302 P

Vestal, Dr. Paul A., Dept. Botany, Rollins College, Winter Park, Florida
32789 B

Membership List 159

Wadell, Dr. Glenn H., Dept. Biol. Sci., Florida Atlantic Univ., Boca Raton,
Florida 33432

Wade, Richard A., Box 2007 Flagler St., Key West, Florida 33041 B

Wallace, Dr. H. K., Dept. Biology, Univ. Florida, Gainesville, Florida
32601 B

Ward, Dr. Daniel B., 733 S.W. 27th St., Gainesville, Florida 32601 B

Warnick, Dr. Alvin C., McCarty Hall, Univ. Florida, Gainesville, Florida
32601 B

Warnke, D. A., Oceanographic Institute, Florida State Univ., Tallahassee,
Florida 32306 B

Warren, Joseph F., Jr., 16101 Ist St., E. Ridington Beach, Florida 33708

Wassell, Glenn H., 641 Shore Dr., Boynton Beach, Florida 33435

Webb, Dr. S. David, Florida State Museum, Gainesville, Florida 32601 B

Weber, Dr. George F., 1122 S.W. 3rd Ave., Gainesville, Florida 32601 B

Webster, W. C., Box 526, Cottage Hill, Florida 32533

Weidner, James P., 403 Reed Lab., Coll. Engineering, Univ. Florida,
Gainesville, Florida 32601

Weigel, Dr. Robert D., Biology, Illinois State Univ., Normal, Illinois
61761 B

Weithe, Dr. Rudolph G., 415 45th Ave., S., St. Petersburg, Florida 33705

Weisbord, Norman E., Dept. Geology, Florida State University, Tallahassee,
Florida 32306 P

Weise, Dr. Emil H., 1360 Avondale Ave., Jacksonville, Florida 32205 B

Weise, Gilbert N., 8601 Emerald Isle Circle N., Jacksonville, Florida 32216

Weiser, Dr. Josejf, Wiley College, Marshall, Texas 75670

Wellman, Wayne E., 967 S.W. 5th St., Miami, Florida 33130 P&S

Wells, Dr. Harry W., Dept. Biological Sciences, Univ. Delaware, Newark,
Delaware, 19711 B

West, Felicia E., 1906 N.E. 9th St., Gainesville, Florida 32601

Westfall, Dr. Minter J., Jr., Dept. Zoology, Univ. Florida, Gainesville, Florida
32601 B

Westgate, Dr. Philip J., Central Florida Experiment Station, P. O. Box 909,
Sanford, Florida 32771 B

Wheat, Dr. Myron W., Jr., Dept. Surgery, College Medicine, Univ. Florida,
Gainesville, Florida 32601 M

Whittaker, Edward, 1320 N.W. 14th St., #503, Miami, Florida 33125

Wiesler, Dr. Donald P., 149 Poe Dr., Winter Haven, Florida 33880

Wilcox, Dr. Charles J., Dept. Dairy Sci., Univ. Florida, Gainesville, Florida
32601 B

Wilkinson, Dr. Robert C., Dept. Entom., Univ. Florida, Gainesville, Florida
32601

Williams, Dr. James H., Route 1, Box 312, Avon Park, Florida 33825 S

Williams, Louise, 511 Wilson Ave., Lakeland, Florida 33801

Williams, Lovett E., Game & Fresh Water Fish Comm., Suite 21, 412
N.E. 16th Ave., Gainesville, Florida 32601

Williams, Dr. Robert H., Univ. Miami, P. O. Box 8263, Coral Gables,
Florida 33124 B

160 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Wilson, Druid, E-506, U. S. National Museum, Washington, D. C. 20560 B

Wilson, Dr. Henry R., Poultry Science Dept., Univ. Florida, Gainesville,
Florida 32601

Wilson, Dr. John L., Florida Mem. College, St. Augustine, Florida 32084 P

Wiltse, Dr. James C., Route 3, Box 180, Orlando, Florida 32811

Wirtz, William O., II, 451 N. Triphammer Rd., Ithaca, New York 14850

Wisner, Carl V., Jr., P. O. Box 260, Fort Lauderdale, Florida 33302

Witham, Ross, 10 Lake Point, North River Shores, Stuart, Florida 33494 B

Wolfenbarger, Dr. D. O., Sub.-Trop. Exp. Sta., 18905 S.W. 280th St., Home-
stead, Florida 33030 B

Wolfgang, Dr. Rebecca, Okaloosa-Walton Jr. College, Valparaiso, Florida
32580

Woolfenden, Dr. Glen E., Dept. Zoology, Univ. South Florida, Tampa,
Florida 33620 B

Wright, Shirley Jean, P. O. Box 4552, Miami, Beach, Florida 33141

Yaffa, Harold, Dept. Biology, Miami-Dade Jr. College, North Miami, Florida
33167 B

Yakaitis, Dr. Albinia, Dept. Anatomy, Univ. Miami, Coral Gables 33134

Yerger, Dr. Ralph W., Dept. Biol. Sci., Florida State Univ., Tallahassee,
Florida B

Young, Dr. Harold, P. O. Box 539, Monticello, Florida 32344

Zeiller, Warren, 910 Catalonia Ave., Coral Gables, Florida 33134

Zeppa, Robert, Dept. Surgery, P. O. Box 875, Biscayne Annex, Miami,
Florida 33152

Zinner, Dr. Doran D., 2017 Alhambra Circle, Coral Gables, Florida 33134 B

Zinober, Dr. M. R., 2105 Friley Road, Ames, Iowa 50012

Zirin, Benjamin O., 199 W. 24th St., Hialeah, Florida 33010

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1967

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Presbyterian College
Florida Southern College
Florida State University
Jacksonville University
Marymount College
Miami-Dade Junior College
Mound Park Hospital Foundation
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida
University of Tampa

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1967

President: Jackson P. SICKELS
Department of Physical Sciences, University of Miami
Coral Gables, Florida

President Elect: CLARENCE C CLARK
Department of Physical Sciences, University of South Florida
Tampa, Florida

Secretary: Joun D. McCRoNnE
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: JAMES B. FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: Pierce BRODKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges

should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

—_

a
\
up
eae

oT Quarterly Journal
of the
Florida Academy of Sciences

Vol. 30 September, 1967 No. 3

CONTENTS
Differential thermal analysis of wavellite Frank N. Blanchard 161
Effects of gamma irradiation on prickly pear cactus Carl D. Monk 168
Notes on Balanus humilis Conrad, 1846 Arnold Ross 173
Spawning season and sex ratio of echinoids John W. Brookbank 177

Sleep behavior in frogs
J. Allan Hobson, Coleman J. Goin, and Olive B. Goin 184

New records of fishes from the western Caribbean
Ray S. Birdsong and Alan R. Emery 187

Cuban lizards of the genus Chamaeleolis
Orlando H. Garrido and Albert Schwartz 197

Natural factors affecting deer movements
Richard F. Harlow and William F. Oliver, Jr. 221

Round-tailed muskrat in west-central Florida John R. Paul 227

Mating behavior of Peromyscus polionotus Michael H. Smith 230

Mailed September 20, 1968

aN
f ‘ 7

QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Editor: Pierce Brodkorb

The Quarterly Journal welcomes original articles containing
significant new knowledge, or new interpretation of knowledge, in
any field of Science. Articles must not duplicate in any substantial
way material that is published elsewhere.

INSTRUCTIONS TO AUTHORS

Rapid, efficient, and economical transmission of knowledge by means of
the printed word requires full cooperation between author and editor. Revise
copy before submission to insure logical order, conciseness, and clarity.

Manuscripts should be typed double-space throughout, on one side of
numbered sheets of 8% by 11 inch, smooth, bond paper.

A Carson Copy will facilitate review by referees.

Mazeins should be 1% inches all around.

TirLes should not exceed 55 characters, including spaces.

Footnotes should be avoided. Give ACKNOWLEDGMENTs in the text and
ADDRESS in paragraph form following Literature Cited.

LITERATURE CrTED follows the text. Double-space and follow the form
in the current volume. For articles give title, journal, volume, and inclusive
pages. For books give title, publisher, place, and total pages.

TaBLEs are charged to authors at $19.95 per page or fraction. ‘Titles
must be short, but explanatory matter may be given in footnotes. Type each
table on a separate sheet, double-spaced, unruled, to fit normal width of page,
and place after Literature Cited.

LecEnps for illustrations should be grouped on a sheet, double-spaced,
in the form used in the current volume, and placed after Tables. Titles must
be short but may be followed by explanatory matter.

ILLUSTRATIONS are charged to authors ($17.30 per page, $15.80 per half
page. Drawincs should be in India ink, or good board or drafting paper, and
lettered by lettering guide or equivalent. Plan linework and lettering for re-
duction, so that final width is 4% inches, and final length does not exceed 6%
inches. Do not submit illustrations needing reduction by more than one-half.
PHOTOGRAPHS should be of good contrast, on glossy paper. Do not write
heavily on the backs of photographs.

Proor must be returned promptly. Leave a forwarding address in case
of extended absence.

REPRINTS may be ordered when the author returns corrected proof.

Published by the Florida Academy of Sciences
Printed by the Storter Printing Company
Gainesville, Florida

QUARTERLY JOURNAL
of the
BLORIDA AGADEMY OF SCIENCES

Vol. 30 September, 1967 No. 3

Differential Thermal Analysis of Wavellite

FRANK N. BLANCHARD

THE general shape of the thermogram obtained from differential
thermal analysis of wavellite is known (Manly, 1950; Kauffman and
Dilling, 1950), but there appears to be no published interpretation
of the pattern. The purpose of this paper is to describe the changes
which take place in wavellite during a differential thermal analysis,
to describe the variations in DTA patterns among different samples
of wavellite, and to evaluate the use of differential thermal analysis
in detection of and quantitative determination of wavellite con-
tained in rocks.

MATERIAL STUDIED

Wavellite specimens used in this study are from the following
locations.

Sample 1. Siglo XX mine, Llallagua, Bolivia. White botryoidal
radiating aggregates of wavellite with cassiterite and quartz.

Sample 2. Highdown quarry, Filleigh, near Barnstable, Devon,
England. Yellowish gray radiating aggregates of wavellite on black
slate.

Sample 3. Ronneburg, Thuringia, German Democratic Repub-
lic. Green and blue-green radiating aggregates of wavellite on
black slate.

Sample 4. Forastara vein, Salvadore section, Llallagua, Bolivia.
Grayish white radiating encrustations of wavellite on quartz crys-
tals.

Sample 5. Bohemia, Czechoslovakia. Grayish white, with pale
orange staining, radiating aggregates of wavellite.

Sample 6. Near Ashville, Jefferson County, Florida (Blanchard

162 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

and Denahan, 1967). White to grayish white microcrystalline
radiating aggregates of wavellite in sandstone.

METHODS

Differential thermal analyses were made with a Fisher model
260P Differential Thermalyzer. The output from a platinel dif-
ferential thermocouple was amplified with a d.c. amplifier employ-
ing a Union Carbide operational amplifier (H6010), and the signal
from the amplifier was used to drive the X-axis of a Houston In-
strument Corp. Model HR-96 X-Y recorder. The Y-axis of the re-
corder was driven by the output from the furnace-temperature
thermocouple, one junction of which was maintained at 0 Centi-
grade, and special graph paper was used to correct for nonlinearity
of the thermocouple output. Samples were heated at 10° or 25°
per minute, or both. Preheated alumina was used as an inert refer-
ence material, All samples were sieved, and for most determina-
tions the fraction between 80 and 200 mesh was used.

Weight changes associated with DTA peaks were determined
on separate samples by weighing the sample (to the nearest 0.1
mg) before and after a thermal reaction.

RESULTS AND DISCUSSION

Fig. 1 shows representative thermograms for each of the samples
studied. Each determination was made with an 80 to 200 mesh,
100 mg sample heated at a rate of 10° per minute. The reactions
appearing between room temperature and 150° are attributed to
loss of absorbed water and to a combination of imperfect thermo-
couple placement and a difference in thermal conductivity between
the wavellite and alumina. Between 150° and 380°, there is clearly
a typical pattern for wavellite with a strong endothermic reaction
at about 260° and a weak to very weak endothermic reaction at
320°; however, there is also variation from sample to sample in the
peak-height ratio. Extremes in this variation are seen in samples |
and 6, while sample 2 represents a median. Because these three
samples represent extremes and median, and because of quantities
available, samples 1, 2, and 6 were chosen for more complete study
by x-ray, thermal, and optical techniques. From 380° to 1150°, no
prominant DTA peaks show up with the degree of amplification

BLANCHARD:

Thermal Analysis of Wavellite

163

DIFFERENTIAL TEMPERATURE

2 el

Be

ENDOTHERMIC

Spee
eee

| Peeled
Sore
5 abo 1000 0 50¢

TEMPERATURE ¢C

Fig. 1. Thermograms of Samples 1-6, showing both the similarity and the
variability of DTA patterns from wavellite.

1000

164 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

used; very weak exothermic reactions do occur (weaker and more
poorly defined than is obtained from the quartz inversion), but
there is little or no consistency from one sample to the next.

Weight losses between 150° and 380° (associated with the two
endothermic reactions) are nearly the same for all samples (24.95-
26.55 per cent), averaging 25.6 per cent. From this it is concluded
that all of the samples contain about the same percentage of water
(below 380°), about 25.6 per cent, which is slightly less than the
26.02 per cent computed from the ideal wavellite composition
[Al,(PO,)2(OH);.5H.O]. The percent of the total water which is
lost during each endothermic reaction corresponds roughly with
the ratio of the peak magnitudes. Factors such as grain size (in
ranges between 40—300 mesh), heating rate, and packing of sample
do not appreciably effect the relative heights of the two peaks.

From consideration of the wavellite structure, weight loss meas-
urements, x-ray diffraction patterns, and optical measurements, it
is concluded that as wavellite is heated it undergoes, successively,
partial to complete dehydration, dehydration and dehydroxylation
accompanied by partial recrystallization, and finally more complete
recrystallization (and possibly inversions) into a form or forms of
aluminum phosphate.

Water and hydroxyl are held structurally in wavellite in differ-
ent ways: 61.5 per cent of the total water (present as H,O plus
OH-) is held as H,O in octahedral coordination (along with OH-
and O~) about Al**, 15.4 per cent is held as H.O in large cavities
in the structure, and 23.1 per cent is held as OH~ in octahedral co-
ordination (along with O? and H.O) about Al** (computed from
structural data provided by Araki and Zoltai, 1966 and Zoltai, per-
sonal communication). The weight loss associated with the 260°
peak varies for different wavellite samples from about 58 per cent
(sample 1) to about 73 per cent (sample 6) of the total weight
loss. Seventy-three per cent corresponds approximately to the per-
centage of molecular H.O in the structure (76.92 per cent), and
the 260° peak is attributed, therefore, to loss of part to all of the
molecular water. In some wavellite (such as sample 1) as little
as 58 per cent of the total water is lost over the 260° peak, and
this corresponds (perhaps fortuitously, perhaps significantly ) with
the octahedrally coordinated water, X-ray diffraction patterns of
wavellite which has been heated to 290° (past the 260° peak, but

BLANCHARD: Thermal Analysis of Wavellite 165

before the 320° peak) show wavellite lines, reduced in intensity,
but no new lines (Fig. 2). Apparently the wavellite structure
still exists after loss of the molecular water of crystallization.
This might be expected because the octahedrally coordinated
water is not shared by adjacent octahedral units in the structure
and thus is not essential in holding the structure together. Opti-
cally, the heated wavellite has indices (approximately 1.48), bire-
fringence, and 2V angle considerably lower than for normal wavel-
lite.

The weight loss occurring over the second DTA peak (320°)
makes up the remainder (27 to 42 per cent) of the total weight
loss. This corresponds with the loss of any remaining H.O (if there
is any) and all of the OH. X-ray diffraction patterns of wavellite
heated to 380° (past the 320° peak) show the emergence of a
weakly developed new structure (Fig. 2) with d-spacings close to
those of AlPO,-cristobalite (D-Yvoire, 1961). Alumina tetrahedra

290° ae

380° —-——-—---p"

1100° |

OT

DEGREES 26

Fig. 2. X-ray diffractometer patterns for Samples 1, 2, and 6. Patterns
are for normal wavellite (top row) and for wavellite after heating to 290, 380,
and 1100 C. d-spacings for the well-defined lines showing in the bottom pat-
tern are 4.12, 4.33, 3.90, 2.524.

166 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

are linked together in the structure by sharing of OH~ and loss of
OH therefore results in collapse of the wavellite structure. Further
heating (to 1150° ) results in strengthening of the lines of aluminum
phosphate with some change in relative intensities (Fig. 2). The
higher temperature, weak exothermic reactions are probably pro-
duced as amorphous aluminum phosphate crystallizes, and, per-
haps, as minor inversions occur within the crystallized AIPO,.
Optically, the material which has been heated past 320° is isotropic
with a refractive index between 1.44-1.46. A slight increase in re-
fractive index occurs after the material has been heated to 1150°.

Differences from sample to sample in peak-height ratio (260°:
320°), and in the weak exothermic reactions at higher tempeya-
tures, may be related to differences in degree and nature of crystal-
linity among the wavellite samples. This suggestion is supported
from study of the material included in this report, and from pre-
vious work (Blanchard and Denahan, 1967, 1968). Microcrystal-
line to cryptocrystalline wavellite from certain localities in Florida
lack the 320° peak, microcrystalline material from Florida (such as
sample 6) shows a very weak 320° peak, and macrocrystalline
wavellites (samples 1-5) show well developed peaks at 320°. It is
also possible, however, that these differences are related to minor
elements in the wavellite.

Differential thermal analysis provides an easy, sensitive, and
relatively inexpensive technique for detection of wavellite in rocks.
It is particularly useful where the wavellite is microcrystalline to
cryptocrystalline and intergrown with other minerals (such as
clays), a condition which makes positive identification by optical
methods difficult, and a condition which is commonly encountered
in wavellite-bearing rocks from the aluminum phosphate zone of
Bone Valley and Hawthorn formations of Florida. Other hydrous
phosphates (such as crandallite, millisite, and variscite-strengite )
show distinctly different patterns, and there seems to be little inter-
ference from other minerals which commonly occur with wavellite.
In a matrix of quartz, as little as 5 per cent (by weight) is easily
detected in a 100 mg sample by the methods employed in this
study. Increasing the weight of the sample, the rate of heating,
and amplifier gain would likely increase the sensitivity even more.
Approximate quantitative determinations are also possible, using
the 260° peak height, or area under the 260° peak, as a measure of

BLaNcHARD: Thermal Analysis of Wavellite 167

concentration, An improvement in accuracy and precision is ac-
complished by using an internal standard to eliminate errors re-
sulting from differences in packing and thermocouple placement
from sample to sample. In quartz-free samples, quartz may be
added to the sample in constant proportion; relative concentration
of wavellite may then be related to the ratio wavellite peak-height
to quartz peak-height.

ACKNOWLEDGMENTS

For use of X-ray diffraction equipment, I am grateful to Dr. R.
W. Gould and to the Department of Metallurgy, University of
Florida. Stephen A. Denahan assisted in preparation of samples.

LITERATURE CITED

ARAKI, TAKAHARU, AND Trpor ZOLTAI. 1966. Crystal structure of wavellite
(abstract). G. S. A., Program 1966 Annual Meetings, p. 6.

BLANCHARD, FRANK N., AND STEPHEN A. DENAHAN. 1967. Variscite from
the Hawthorn Formation. Quart. Jour. Florida Acad. Sci., vol. 29, no.
3, pp. 163-170.

. 1968. Wavellite-cemented sandstones from northern Florida. Quart.
Jour. Florida Acad. Sci., vol. 29, no. 4, pp. 248-256.

Dp YvorRE, FERDINAND. 1966. Etude des phosphates d’aluminum et de fer
trivalent. L’orthophosphate neutre d’aluminum. Bull. Soc. Chim.
France, pp. 1762-1776.

KAUFFMAN, A. J., JR.. AND E. Don Dmuinc. 1950. Differential thermal
curves of certain hydrous and anhydrous minerals, with a description
of the apparatus used. Econ. Geol., vol. 45, pp. 222-244.

Manty, Roperr L., Jr., 1950. The differential thermal analysis of certain
phosphates. Amer. Mineral., vol. 35, pp. 108-115.

Department of Geology, University of Florida, Gainesville,
Florida 32601

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968)

Effects of Gamma Irradiation on Prickly Pear Cactus
Cart D. Monk

On May 11, 1965, a 12-year-old field on the AEC Savannah
River Plant was exposed for 400 hours to a 9200 curie "Cs irradi-
ator (Monk 1965). The field vegetation was typical of abandoned
farmland on the dry, excessively drained Lakeland sandy soils with
annuals predominant in number of individuals. Scattered through
the exposed portion of the community were forty-five plants of
prickly pear cactus (Opuntia compressa (Salisb.) Macbr.). At the
time of exposure most of the cactus plants were in a similar flower-
ing stage. Each plant had fully opened flowers at the beginning
of the exposure and many possessed flower buds which did not
open until the termination of irradiation. The precise stage of
micro- and megasporogenesis was unknown. Prior to and at vari-
ous periods following exposure a number of biological measure-
ments were made on each plant. This paper represents a report on
the effects of irradiation on prickly pear.

METHODS

Twelve days prior to radiation exposure, the number of stem
segments, new vegetative buds, and flower buds were recorded for
each plant in the population. On August 23, the number of stem
segments and fruits per plant were counted. In early November
all fruits were collected for measurement of diameter and seeds per
fruit were counted. The seeds were dried at room temperature for
one week and those of each plant were weighed collectively then
planted in the field for germination studies. Except for germina-
tion tests similar measurements were made in 1966.

The radiation exposure of each plant was monitored by Toshiba
Low Z phosphate glass rods. The rods were placed in opaque plas-
tic vials to minimize fading due to sunlight (Blaylock and Wither-
spoon 1965).

RESULTS AND DISCUSSION

The forty-five plants were arranged in descending order of radi-
ation exposure. The means of the biological data and rate of radi-
ation exposure were calculated for each successive group of five
plants (Tables 1-2). The basic form and position of the data were
not materially affected by the data treatment. The actual range in
R/hr was 31.3 to 0.1.

Monk: Irradiation of Prickly Pear 169

All flower and vegetative buds aborted on plants receiving in
excess of 22.5 R/hr. Plants with exposures greater than 27.5 R/hr
died within a month following irradiation. Subsequently, all plants
included in the 23.5 R/hr exposure level (Table 1) died except for
one plant exposed at a rate of 18.7 R/hr.

Fruit size, seed weight, and seeds per fruit were more radiation
sensitive than fruit and stem segment survival in 1965 (Table 1).
The former three exhibited a marked reduction between 2.9 and
5.9 R/hr. Fruit survival began to decrease at the same radiation
level, however, variability was too large for the trend to be signifi-

TABLE 1

Effects of 400 hours of high dose irradiation on a population of prickly pear
cactus. Each figure represents a mean for five plants with one standard error.

R/HR
30.4 23.5 13.0 9.3 5.9
=| xD) ate +0.9 +0.5 +0.5

1965
Fruit

Survival (%) 0 0 5NGs=16:3) 564-1015) 6 O21 7
Fruit

Diameter (Cm) — — OZ OW | e370 2 0:0
Seeds/ Fruit — — yet) Ore IE Gse 3 S52 91.6
Seed

Weight (Mg) — — NOVAS i 22. Aaa, Dio) P29 Ase 2
Stem Segment

Survival (% ) — Os OSG O14 1 Go.O2=8 ino. 166 9==. oF2
1966
Fruit

Survival (% ) 0 0 OO Soe So, o=s= oni o0.O==) Onl
Fruit

Diameter (Cm) — — ROSE OM AO =e Or 1G == 0:0
Seeds /Fruit — — BAe ae le O oD pata 2 Ol OA a OrO
Seed

Weight (Mg) = — 21-4, O52 3.7 21.332" 3.9
1966/1965

Flower Bud

Production (%) — 35.2200 25.3419.3 27.74 8.7 48.72:17.0
1966/1965

Vegetative Bud

Production

(ratio ) cs 0.51 1.67 7.00 1.00

170 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

cant. All of the non-aborted fruits contained seeds, many of which
possessed only one. This fact tended to extend the radiation toler-
ance as well as increase the variability of that biological parameter.
Stem segment survival was not materially affected until the 23.5
R/hr isodose line.

The germination response was similar to the data presented in
Table 1, however, germination percentages were too low in the
controls for the experiment to be conclusive. Seeds from plants
receiving 9.3 R/hr or more were generally less viable than those
from plants with less exposure.

TABLE 2

Effects of 400 hours of low dose irradiation on a population of prickly pear
cactus. Each figure represents a mean for five plants with on standard error.

R/HR
2.9 1.4 0.9 0.3
a5 ))58) +0.1 ==()),J1 +0.0 0.0
1965
Fruit

Survival (% ) 66.622 5.8 9-72.22 3:7 (81/222 5.8) 7 7si0 sa R oma O: toe
Fruit

Diameter ((Gm)- 1.62: O01 142 01 1.52.01 > 1920s eeleeae
Seeds/ Fruit 25.622 1.3 2742 1.6 27.622 1:8) 23.522 Osos aero
Seed

Weight (Me) 38.722 2:8) 29732 2:1 931.322 274 S218s2 0S 2p nU aeons
Stem Segment

Survival (%) 92:42: 44° 95.72: 3.1 98A== 1-6 S790S25S:6 go OG sare

1966
Fruit

Survival (%)) 65.72: 5137 69.52211.3 79,722 14.9) (625s 213 2 ale oa emonl
Fruit

Diameter (Cm) 1.6 00 - 17+ O01 1.5= 0.05) ees Olean

Seeds/ Fruit 20.522 1.1 87.92 2:5 18-72 4:3 2310s olee2 eal
Seed

Weight (Me) 35.022 4.3 3412 41 3054234 3322 73.0) olelese0
1966/1965

Flower Bud

Production(% ) 56.2+14.6 63.5+16.8 74.2+11.1 69.7+ 86 64.5=13.1
1966/1965
Vegetative Bud

Production
(ratio ) 0.71 0.19 0.50 0.75 0.19

Monk: Irradiation of Prickly Pear v/a

One year following irradiation, the prickly pear cactus popula-
tion still exhibited a response to exposure. All plants with less
than 17.5 R/hr produced flower buds in 1966. Flower bud pro-
duction in 1966 was consistently lower than in 1965. Those plants
exposed to 9.3 R/hr or more were less productive than the rest of
the population (Table 1).

In 1966, no fruits survived on plants receiving in excess of 13.6
R/hr. Fruit survival in 1966 was slightly less at lower exposures
than in 1965 and considerably lower at the higher radiation levels
(Table 1). Fruit size and seeds per fruit were not affected in 1966,
however, seed weight tended to be lower at the higher exposures.

Certain segments of the cactus population formed more vege-
tative buds in 1966 than in 1965 while the remainder produced less
(Table 1). The ratio of 1966/1965 new stem segments was higher
in those plants at the 13.0-5.9 R/hr levels than in those exposed at
higher or lower levels.

The enhancement of vegetative bud production at the 13.0-5.9
R/hr levels could be interpreted as a stimulatory effect for which
evidence exists both for (Sax 1963) and against (Skok et al., 1965).
However, in light of continued reduction of flower bud production,
fruit survival, and lighter seeds in 1966, the enhancement of vege-
tative bud formation may represent a radiation induced imbalance
in the normal development in cactus whereby the plant reserves
are channelled more into vegetative recovery. Survival from radi-
ation damage in a small segment of a plant population is depend-
ent upon vegetative recovery or the production of viable seeds. If
either of these recovery mechanisms is enhanced at the expense of
the other, it is difficult to view the enhancement in terms of a “true”
stimulatory response but rather as a compensatory recovery mech-
anism.

SUMMARY

The irradiation effects on prickly pear may be summarized as
follows:

(1) Exposures in excess of 18.7 R/hr administered over a 400
hour period are lethal.

(2) Exposures of 5.9 R/hr are sufficient to cause a major re-
duction in fruit size, seeds per fruit, and seed weight and a slight
reduction in fruit survival.

(3) Stem segment survival is reduced at 13.0 R/hr.

172 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

(4) One year following irradiation, flower bud production and
fruit survival are decreased in plants receiving in excess of 5.9 R/hr
while vegetative recovery is greatest in plants exposed at 13.0-5.9
R/hr.

(5) The enhancement of vegetative bud production in 1966
may be a recovery mechanism from radiation induced damage
rather than a “true” stimulatory response to radiation.

ACKNOWLEDGMENTS
Support for this project was provided by Contract AT (38-1)-310
between the AEC and the University of Georgia.

LITERATURE CITED

BiayLock, B. G., AND J. P. WitHERSPOON, JR. 1965. Environmental factors
affecting glass rod dosimetry. Health Phys., vol. 11, pp. 549-552.

Monk, C. D. 1966. Effects of short-term gamma irradiation on an old field.
Radiation Botany, vol. 6, pp. 329-335.

Sax, K. 1963. The stimulation of plant growth by ionizing radiation. Radi-
ation Botany, vol. 3, pp. 179-186.

SKOK, J., W. CHorNeEy, AND E. J. RAkosnik, Jr. 1965. An examination of
stimulatory effects of ionizing radiation in plants. Radiation Botany,
vol. 5, pp. 281-292.

Institute of Ecology and Department of Botany, University of
Georgia, Athens, Georgia 30601.

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968 )

Notes on Balanus humilis Conrad, 1846

ARNOLD Ross

Tue balanomorph barnacle Balanus humilis was described by
Timothy Abbott Conrad in 1846 in one of his many studies on the
organic remains from the Tertiary sediments of the southeastern
United States. Since the original publication of this taxon, it has
never been redescribed, refigured, nor has its true nature been
determined. An attempt to locate the type specimen in the col-
lections of both the Paleontological Research Institution, Ithaca
and the Academy of Natural Sciences of Philadelphia has proven to
be fruitless.

Darwin (1854, p. 495) was the first to discuss Conrad’s species,
succinctly stating, “The opercular valves are not described, and I
doubt whether the species could be recognized.” Pilsbry (1930,
p. 433), in referring to this taxon, merely noted that it was “un-
recognizable.” Both of these workers unquestionably based their
comments on the fact that the description of B. humilis was too
brief and incomplete to be of taxonomic value, and Conrad’s
illustration of this species, which measures less than 5 mm in
height, was reproduced too small to reveal diagnostic characters.
Mansfield (1937, p. 9) also cited Conrad’s taxon in a discussion
of the molluscan fauna of the Tampa Limestone.

Recently, the author in a discussion of the Eocene Balaninae
of the world pointed out that “. . . the Tampa Limestone from
which this species [B. humilis] presumably was collected, is as-
signed an early Miocene age” (Ross, 1965, p. 59; See Mansfield,
1937). No attempt was made in that study to discuss the morphol-
ogy or nature of this species, nor was this attempted in a subse-
quent paper (Ross and Newman, 1967).

The original description of B. humilis (Conrad, 1846, p. 400)
is as follows: “Suboval, not elevated; aperture large, somewhat
diamond shaped; valves with rather fine longitudinal approximate
sulci.” Conrad’s illustration (see Fig. 1) clearly shows that the
shell is divided into two distinct horizontal zones. The upper half
is for the most part smooth, with the exception of three, moderate-
ly broad, vertical sulci. The lower half of the shell is unequally
divided into three major zones by approximate, vertical sulci.

174 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Fig. 1. Balanus humilis Conrad, 1846; original figure reproduced natural
size (left), and same figure redrawn greatly enlarged (right).

Each of these basal zones is finely ribbed, the ribs slightly less
than the height of the lower zone.

On numerous occasions while collecting from Miocene sedi-
ments in Florida small, phosphatized steinkerns, or fillings of the
internal cavity of what appeared to be a barnacle were collected
from the sediments of both the Hawthorne and Tamiami Forma-
tions of Middle and Late Miocene ages, respectively. Determina-
tion of the taxonomic identity of these specimens was precluded
owing largely to their mode of preservation. In the cirriped col-
lections of the Department of Fossil Invertebrates at The Ameri-
can Museum of Natural History there is a specimen of what has
been identified tentatively as Balanus glyptopoma_ Pilsbry
(A.M.N.H. 11382/1; Holmes collection, specimen from question-
ably late Miocene of South Carolina) in which the body cavity of
the barnacle is filled, and the calcareous shell is partially eroded
away. Continued eroding of the calcareous shell would no doubt
have left just the phosphatized shell filling.

Based upon a critical comparison of Conrad’s figure, and the
available morphological evidence derived from a study of more
than 20 specimens that are similar to the one figured by Conrad,
the author has concluded that B. humilis is nothing more than
the phosphatized internal filling of a barnacle shell. It can be
demonstrated that the separation between the upper and lower
halves shown on Conrad’s figure, demarcates on specimens, the
lower edge of the depending sheath; on all specimens examined
this break is actually a sulcus. The vertical grooves are merely
those regions of the compartments where the radii or alae are
situated. The fine ribs of the lower zone represent the sulci be-

Ross: Notes on a Barnacle 175

tween the ribs on the exposed surface of the inner lamina of the
parietes. All the above mentioned features can be observed on
the majority of the complete steinkerns thus far examined. In addi-
tion, not infrequently the growth lines of the sheath can be seen
clearly.

Although it is demonstrable that B. humilis is only a phospha-
tized internal filling, there are a number of factors that serve to
obscure the taxonomic identity of this fossil. Foremost among
these is that there are undoubtedly several species in the Miocene
sediments of Florida, as well as elsewhere along the southeastern
Atlantic Coastal Plain, preserved in a manner analogous to that of
B. humilis. Furthermore, consideration must be given to whether
or not the specimen Conrad figured represents a mature individual
or a juvenile specimen. This is clearly brought out by the present
collections from Florida and South Carolina wherein there are
specimens ranging in height from 2.5 to more than 30 mm.

Although B. humilis was described initially as an Eocene
species, this age assignment was subsequently questioned (Ross,
1965, p. 59). The present collections tend to indicate that a Mio-
cene age for B. humilis is closer to being correct, but the age
within this epoch cannot be stated with any certainty.

Conrad reported the type locality of B. humilis to be the “falls”
of the Hillsborough River. However, there is some question on
the part of the writer as to whether or not this species actually
occurs in the Tampa Bay region, or was secondarily transported
there, if indeed the specimen was actually collected from the
Tampa Limestone.

On the basis of the information available the specific identity
of Conrad’s taxon must remain an enigma.

LITERATURE CITED

Conrap, TrmMoruy Apspotrr. 1846. Description of new species of organic re-
mains from the upper Eocene limestone of Tampa Bay. Amer. Jour.
Sci. Arts, ser. 2, vol. 2, no. 6, art. 37, pp. 399-400, 9 text-figs.

DARWIN, CHARLES RosertT. 1854. A monograph on the sub-class Cirripedia,
with figures of all the species. The Balanidae, (or sessile cirripedes );
The Verrucidae, etc., etc., etc. London, Ray Society, pp. i-viii, 1-684,
pls. 1-30.

176 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MANSFIELD, WENDELL CLay. 1937. Mollusks of the Tampa and Suwannee
Limestones of Florida. Bull. Florida Geol. Surv., no. 15, pp. 1-334, pls.
A-D, 1-21, text-figs. 1-2, tables 1-2.

Pitspry, Henry Aucust. 1930. Cirripedia (Balanus) from the Miocene of
New Jersey. Proc. Acad. Nat. Sci. Philadelphia, vol. 82, pp. 429-433,
figs. 1-2, pls. 36-37.

Ross, ARNOLD. 1965. A new cirriped from the Eocene of Georgia. Quart.
Jour. Florida Acad. Sci., vol. 28, no. 1, pp. 59-67, figs. 1-2.

Ross, ARNOLD, AND WILLIAM A. NEwMAN. 1967. Eocene Balanidae of Flor-
ida, including a new genus and species with a unique plan of “turtle-
barnacle” organization. Amer. Mus. Novitates, no. 2288, pp. 1-21, figs.
1-7.

Department of Invertebrate Paleontology, Natural History Mu-
seum, Balboa Park, San Diego, California 92112.

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968 )

Spawning Season and Sex Ratio of Echinoids

Joun W. BrooOKBANK

Durinc the past three years, various embryological projects in
this laboratory have required the use of large numbers of gametes
from large numbers of sea urchins, Lytechinus variegatus (La-
marck), and sand dollars Mellita quinquiesperforata (Leske). In
order to preserve valuable data, records were kept on the number
of these animals collected, and treated to induce spawning, since
May, 1964. These records are a by-product of the other work, and
thus do not include the precise locations of the collecting sites, the
temperature of these locations, the size of all individuals collected,
or histological preparations of the gonads. However, in view of the
large number of animals involved (thousands), and since accurate
temperature data are available for the Cedar Keys area from Coast
and Geodetic Survey records, it was decided that the information
should be made available for those wishing to work on these ani-
mals in Florida, either from the standpoint of their life history or
their embryology. The data also have a bearing on the problem of
sex determination in echinoids.

METHODS

The animals were collected by random dredging in the chan-
nels or on the shallow banks in the vicinity of the University of
Florida Marine Laboratory at Seahorse Key. Animals with test
diameters smaller than two inches (Lytechinus) or two and one-
half inches (Mellita)were discarded. The animals were kept in
fresh sea water (changing several times) on the collecting boat,
and were transported to Gainesville “dry” (in a bucket under a
layer of paper towels soaked in sea water). The trip to Gainesville
requires about one hour. Upon arrival, the animals were trans-
ferred to two-gallon sea water aquaria, and kept in a beverage
cooler at 17 C. The aquaria were supplied with a constant flow
of washed, compressed air. The animals remain viable for about
two weeks under these conditions. No attempt was made to feed
the animals, or to provide continued filtration of the aquaria.
Spawning was induced by the injection of 1-5 ml of 0.55 M KCl
(isotonic) into the coelom through the peristomial membrane

178 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

(Palmer, 1937; Tyler, 1949). “Ripe” animals shed gametes within
5 minutes. The sex of the animal is readily apparent on shedding
in either Mellita or Lytechinus. Other sex limited characteristics
are apparently lacking in both animals, though Lytechinus pictus
(Verrill) and Lytechinus anamesus Clark from Southern California
show sexual dimorphism of the gonopores, those of the female be-
ing statistically larger than those of the male (Tyler, 1944). No
information is available regarding gonopore size of L. variegatus.
The gametes were examined with phase contrast optics, and the
presence or absence of fertilized eggs, sperm motility, primary
oocytes, and eggless-jelly (Mellita) noted. The syringe used for
KC] injection as well as the animals to be injected were rinsed
with tap water to inactivate “stray” sperm. Such precautions, along
with gross observation of spawning gonopores, allowed for diag-
nosis of hermaphroditism (none was noted).

Monthly spawning and temperature data for each year have
been lumped into a three year total since no gross variation in
these figures was evident within a given month. The average
monthly temperature was taken as the average of the two diurnal
extremes recorded for that particular month at Cedar Keys, Florida.
The sampling frequency varied with the month and year between
no collections and one collection every week. Each sample was
assayed in its entirety for sex ratio and unresponsive individuals.
The accumulative (three year) monthly sample sizes ranged _ be-
tween extremes of 16-264 individuals. The average sample size
was 125 for both Lytechinus and Mellita. No collections of Mellita
were made during July and August, and only a small August
sample (16 animals) of Lytechinus is available. Totals for the
entire three year period are tabulated (Table 1) and the propor-

TABLE 1

Spawning data for Mellita quinquiesperforata and Lytechinus variegatus from
May, 1964, to May 1967

Number Male Female Unresponsive
Lytechinus 1504 Ole 205 952
Mellita LAI 381 294 446

tion spawning (per cent) is presented graphically along with a
plot of the temperature (Fig. 1).

BROOKBANK: Spawning in Echinoids 179

100

@
Oo

©
z Le
z= °
= 60 -70
QO =)
op) [=
<

Ee
Zz ta
W 40 a.
O a
uJ
ti =

20 50

o+— iit if Dba
D

Fig. 1. Annual spawning cycle and local temperature. Black bars, Mellita
quinquiesperforata. Open bars, Lytechinus variegatus. Per cent of animals
responding to injected KCl (see text), ordinate. Months of the year (1964

to 1967), abcissa. Black line, mean diurnal temperature at Cedar Keys, Flor-
ida 1964-67.

RESULTS

Aside from the data tabulated below, the following information
was noted. 1) There is no relationship between sex and size, in
either animal, above the size limits previously stated. 2) Males of
both populations tend to spawn spontaneously during the early
part of the shedding period. This usually occurs during the return
trip following collection. Females also shed spontaneously on
rare occasions. 3) Collections during the cooler months frequently
yield a high proportion of males, above 70 per cent. At no time
during the collection period included in this report did the num-
ber of spawning Lytechinus females equal or exceed the number
of males. The sex ratio (males:females) drops from a value of
5:1 in February to about 1.4:1 in April (Lytechinus). The same
holds true for Mellita, though the sex ratio is closer to unity in this
animal (1.3:1 overall). 4) Primary oocytes appear, immediately

180 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

following (two weeks) the initial observations of spawning males.
These are inter-mixed with fertilizable, mature ova in concentra-
tions approaching 20 per cent. 5) During the decline of the breed-
ing period, Mellita females release eggless-jellies, observable macro-
scopically since the jelly coat of these eggs contains red echino-
chrome granules. These jelly coats presumably represent mature
eggs which have been completely or partially absorbed following
the main spawning period. The same may hold true for Lytechinus,
since cytolyzing eggs are occasionally seen toward the end of the
spawning season, though the jelly surrounding the eggs is devoid
of pigment and of the same refractive index as sea water, thus
being invisible even with phase contrast optics.

It is also apparent that, during the period in question and with
the sampling methods used, males outnumber females by an over-
all ratio of 3:2 (Lytechinus) and 4:3 (Mellita). The apparent dis-
crepancy from a 1:1 ratio of sexes is discussed below. Maturation
of gametes unquestionably commences with warm weather, as one
would expect, and probably proceeds on a continuous basis
throughout the summer months in both animals. Synchronous
spawning (perhaps triggered by the females, as they approach
ripeness ) presumably occurs soon after the initial ripening period
in April and May. March of 1967 was the earliest that Lytechinus
was available in abundance in ripe condition.

DIscussIoNn

It seems clear from the evidence above, that here is a prepon-
derance of males in both populations, particularly Lytechinus. The
overall sex ratio may still be 1:1 if one could sex all animals col-
lected, ripe or not. One is, however, examining the populations
for the ratio of ripe males to ripe females. Since no relationship
between size and sex seems to exist, the explanation for the excess
of males observed in the population is probably not to be found
in the phenomenon of protandrous hermaphroditism (e.g., Moore
et al., 1963). Rather, it seems likely that the males mature first
and spawn in response to the shedding of the females when the
latter become ripe. Having spawned, the males regenerate mature
gametes faster than the spent females. Thus, at a given time the
population will consist of more ripe males than ripe females. At
the height of the breeding season, when the animals are found in

BROOKBANK: Spawning in Echinoids 181

aggregates (see Tennent, 1910 and Moore, et al., 1963), nearly all
the animals respond to the treatment. The preponderance of males
is still observed (Table 2). Since it has been shown (Moore et al.,

TABLE 2

Spawning data for Mellita quinquiesperforata and Lytechinus variegatus dur-
ing selected periods of maximum ripeness (more than 70% spawning )

Number Male Female Unresponsive
Lytechinus
April-May
1967 131 81 50 0
Mellita

May 1966 235 112 99 24

1963) that mature animals come together in beds, it is not sur-
prising to note the continued excess of males, since solitary (un-
ripe) animals would be least likely to be collected by dredging
in the more or less random manner described (hunting for the
area yielding the most animals per trial). The extent to which
immature animals aggregate in comparison to mature animals is as
yet undetermined. Corollaries of this theory would be: 1) That
most solitary Lytechinus (and, perhaps, Mellita) found during the
breeding season would be females. 2) The initial trigger for syn-
chronous spawning would, in the absence of artificial stimulation,
be the spawning of the female(s). The above statements apply
to Mellita as well as Lytechinus with the following reservations:
1) Bedding activity in sand dollars would seem to be less marked
than in Lytechinus, since sand dollars seem, in the author's ex-
perience, to be found mainly in rather restricted areas (or beds)
regardless of season. 2) Misclassification of females as unresponsive
individuals is less frequent in Mellita due to the presence of echino-
chrome granules in the jelly surrounding even degenerating eggs,
rendering them macroscopically visible. The latter condition tends
to even the sex ratio during the declining weeks of the breeding
season.

The cytology of sex determination in sea urchins and sand
dollars is not completely clear, mainly due to the small size and
considerable number of the chromosomes. Some genera (Trip-
neustes, for one) appear to have an “XO” (digametic male)

182 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

mechanism (Tennent, 1912). Others, including Lytechinus, may
have a similar mechanism (Tennent, 1912). Shapiro (1935) reports
the sex ratio of Arbacia punctulata at Woods Hole to be 1.03:1 in
favor of females (2,358 animals), a statistic compatible with an
XO mechanism. Sexing in this case was done by cutting the ani-
mals open around the circumference of the test and recording
gonad color (ovaries are red) as well as presence of eggs or sperm.
Hermaphrodites such as those described by Moore et al. (1963)
and others (numerous reports exist, see Harvey, 1956; Hyman,
1955; Boolootian and Moore, 1959) should be investigated cytologi-
cally to determine whether or not there is a chromosomal basis for
this condition, rather than a phenotypic explanation. Until further
information is available, it seems unnecessary and unwarranted to
assume anything different from a 1:1 sex ratio, in the total popu-
lation, of either Lytechinus or Mellita. The assumption of a dif-
ferential rate of gametogenesis, the males being more rapid than
the females in the production of gametes, is sufficient to account
for the observed departures from a 1:1 ratio in the breeding pop-
ulation.
ACKNOWLEDGMENTS

This work was supported in part by a research grant (GM
04659) from the National Institutes of Health. The author wishes
to thank Professor James H. Gregg for flying many of the speci-
mens from Cedar Keys to Gainesville.

LITERATURE CITED

BooLooTian, R. A. AND A. R. Moore. 1959. <A case of ovotestes in the sea
urchin Strongylocentrotus purpuratus. Science, vol. 129, pp. 271-272.

Harvey, E. B. 1956. The American Arbacia and other sea urchins. Prince-
ton University Press, Princeton, N. J., 298 pp.

Hyman, L. H. 1955. The Invertebrates. Vol. 4. Echinodermata. Mc-
Graw-Hill, New York, 746 pp.

Moore, H. B., T. Jurare, J. C. BAvER, AND J. A. Jones. 1963. The biology
of Lytechinus variegatus. Bull. Mar. Sci. Gulf and Carib., vol. 13, pp.
23-53.

PauMer, L. 1937. The shedding reaction in Arbacia punctulata. Physiol.
Zool., vol. 10, pp. 352-357.

SHaprro, H. 1935. A case of functional hermaphroditism in the sea urchin,

Arbacia punctulata, and an estimate of the sex ratio. Amer. Nat., vol.
69, pp. 286-288.

BROOKBANK: Spawning in Echinoids 183

TENNENT, D. H. 1910. Variation in echinoid plutei. Jour. Exp. Zool., vol. 9,
pp. 657-714.

TENNENT, D. H. 1912. The correlation between chromosomes and particular
characters in hybrid Echinoid larvae. Amer. Nat., vol. 46, pp. 68-75.

Tyxter, A. 1944. Sexual dimorphism in sea urchins. Anat. Rec., vol. 89, p.

573.

1949. A simple, non-injurious method for inducing repeated spawn-
ing of sea urchins and sand dollars. The Collecting Net, vol. 19, pp.
19-20.

Department of Zoology, University of Florida, Gainesville, Flor-
ida 32601.

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968)

Sleep Behavior of Frogs

J. AtLAn Hopson, CoLEMAN J. Gorn, AND OLIVE B. Gorn

Prrron (1913) established postural and threshold criteria of
sleep as major elements in his definition of that behavioral state. A
review of the literature and his own naturalistic laboratory observa-
tions led him to conclude that mammals, birds, and reptiles all sleep;
that insects, fish and amphibians clearly fulfill the postural, but not
so clearly the threshold, criterion of sleep; and that the behavior of
other invertebrates defies analysis by means of his definition. Re-
cent physiological work has added an electrographic criterion of
sleep, the application of which strengthens Pieron’s views on the
presence of sleep in mammals, birds, and reptiles and also indicates
considerable differentiation of phases of sleep within these classes
of animals.

A recent study (Hobson, 1966) has shown that the bullfrog,
Rana catesbeiana, does not fulfill either the threshold or the elec-
trographic criterion of sleep. In brief summary, though impressively
immobile and clearly at rest for long periods of time, the bullfrog
remains alert, reacting with equal speed and intensity to experi-
mental stimuli delivered at any phase of the rest-activity cycle.
During periods of rest, furthermore, the brain waves of the bull-
frog are of low voltage and fast electrical activity which is associa-
ted with alertness in other animals. To obtain complementary field
data for this species of frog and to examine two species of tree frogs
whose field behavior has been extensively studied and described
(Goin and Goin, 1957; Goin, 1958) we made observations of frogs
in the region of Gainesville, Florida, during the month of March,
1966.

Rana catesbeiana, normally active at night, proved impossible to
observe at rest by day. In a trip around a small pond where these
frogs are known to abound we saw only 6 specimens, all moving.
Despite our caution, a frog would be seen only after it had begun
a long leap from the edge of the pool into deeper water about 15
or 20 feet ahead of us. The extreme sensitivity of this animal to
stimulation is as remarkable in the field as it is in the laboratory
and it is confirmed that, in all observed cases at least, the state of
rest is characterized by alertness and not by the loss of reactivity

Hopson ET AL.: Sleep in Frogs 185

required by our definition of sleep. We cannot rule out the possi-
bility, of course, that the bullfrog sleeps unobserved but can state
that the observable behavior of this species is the same in the field
as it is in the laboratory. We therefore conclude that Rana cates-
beiana does not sleep.

Hyla squirella and Hyla cinerea, also nocturnally active, were
invariably found to be resting by day. In a mesic hammock area we
observed at least 20 specimens of each species of frog, all of which
were immobile, with eyes closed, and with heads pointing toward
the tips of the palmetto leaves where they rested about 2 feet above
the ground. Only after moderately intense and direct stimulation
such as shaking the plant, did the animals assume a crouching pos-
ture, open the eyes, and sometimes attempt to flee by leaping to a
neighboring plant. In every case this sort of crude clinical testing
indicated that the resting tree frogs had a clearly elevated thres-
hold to sensory stimulation and could thus be said to be alseep by
behavioral definition.

We speculate that the presence of clearcut sleep behavior in
Hyla and its absence in Rana may reflect an evolutionary modifica-
tion of behavior: with the adaptation of Hyla to a more terrestrial
and arboreal mode of life, there must have developed increasing
neuromuscular sophistication, as well as important changes in res-
piratory functions. Kleitman’s evolutionary theory of sleep asserts
that sleep behavior is not a primitive state but one which has evolved
in parallel with increasing complexity of waking behavior. We
would, therefore, expect to find correlative neurophysiological dif-
ferences between Rana and Hyla. It will be of interest now to study
the electrographic correlates of the putative state of sleep in tree

frogs.

LITERATURE CITED

Goin, C. J., AND O. B. Gorn. 1957. Remarks on the behavior of the squirrel
tree frog, Hyla squirella. Ann. Carnegie Mus., vol. 35, pp. 27-36.

Gorn, O. B. 1958. A comparison of the non-breeding habits of two tree frogs,
Hyla squirella and Hyla cinerea. Quart. Jour. Florida Acad. Sci., vol. 21,
no. 1, pp. 49-60.

Hosson, J. A. 1966. Electrographic correlates of behavior in the frog with
special reference to sleep. Electroencephalography and clinical neuro-
physiology, vol. 22, pp. 113-121.

186 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

KLEITMAN, N. 1963. Sleep and wakefulness. Univ. of Chicago Press, Chicago,
SSI ps

Preron, H. 1913. Le problem physiologique du sommeil. Masson, Paris, 520
pp.

Department of Psychiatry, Harvard Medical School, Boston,
Massachusetts, and Department of Zoology, University of Florida,
Gainesville, Florida.

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968 )

New Records of Fishes from the Western Caribbean

Ray S. BrrpsonG AND ALAN R. EMERY

INCREASING interest in the inadequately studied Central Ameri-
can marine fish fauna warrants the documentation of certain of the
new range records that have accumulated in the Institute of Marine
Sciences, University of Miami, fish collection during the past sev-
eral years. The material reported here is derived from two expedi-
tions to the area, one by Walter A. Starck II in 1961 and the other
by the senior author in 1966. Starck’s collections include 28 stations
in Yucatan and British Honduras and the senior author’s include
10 stations from Courtown and Albuquerque Cays near Nicaragua.
These collections are comprised of about 209 species in 860 lots.
New records have been reported only for species for which recent
authoritative ranges are available. The list of species probably
contains new records of which we are unaware.

The catalog numbers given below are all from the Institute of
Marine Sciences, University of Miami (—UMML) and are followed
by the number of specimens and the range of standard lengths in
parentheses. Material examined is given only for new records, but
all the species listed are cataloged in the museum at the Institute
of Marine Sciences, University of Miami.

COLLECTION AREAS

Collections were made in a variety of habitats but primarily on
patch reefs and reef top areas. Most of the collections were made
with rotenone derivations and small multibarb spears. Those made
at Courtown and Alburquerque Cays were made in tidepools, on
patch reefs within the atoll lagoons and on the inner edge of the
barrier reefs in depths to eight meters. The ecology of these reefs
is discussed by Milliman et al., in press. Collections in Yucatan
were made at Cozumel Island and Banco Chinchorro. At Cozumel,
stations were made on coral reefs to 25 meters, on patch reefs,
beaches, in a salt pond, and a cave. At Banco Chinchorro, poison
stations were made on patch reefs and beds of Thalassia. In one
case a small trawl and cast net were also used. Collections in

188 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

British Honduras were made at Belize, Turneffe Island, and Light-
house Reef. These included stations in mangrove habitats, but
most were made on rocky shores, patch reefs, and beds of Thalas-
sia in shallow water. Collection areas are shown in Fig. 1.

List or NEw RECORDS

The following list of species includes those which are new dis-
tributional records. For each, the material examined is noted and
the previously reported range recorded.

LABRIDAE

Halichoeres garnoti (Valenciennes). Nicaragua: Albuquerque Cays, UMML
23096 (9 specimens, 43.6 mm—101 mm SL). British Honduras: Turnefte
Island, 10312 (1, 15.0), 9842 (1, 21.5); Lighthouse Reef, 9277 (4, 28.2—
48.2), 9473 (1, 51.7). Yucatan: Cozumel Island, 9515 (2, 36:0—=54.5):
Banco Chinchorro, 9636 (4, 24.3—35.5). Reported previously from Ber-
muda, Florida. Bahamas, Greater and Lesser Antilles, Venezuela to Brazil
(Randall and Bohlke, 1965, p. 252; Cervigon, 1966, p. 606).

Halichoeres maculipinna (Miller and Troschel). Nicaragua: Albuquerque
Cays, 23082 (1, 53.8), 23080 (2, 50.0, 63.5); Courtown Cays, 23078 (24,
29.0—97.7 ), 23079 (11, 32:.9—74.5), 23083) (33:3) 2380S ar Giemoless)
Yucatan: Cozumel Island, 9205 (1, 43.3), 9712 (1, 18.5); Banco Chin-
chorro, 9307 (2, 23:9—25.8), 9394 (3, 16.2—28:8),, 9750NGi Zona) ie=
corded from North Carolina to Florida, Bermuda, Bahamas, Greater and
Lesser Antilles, Venezuela to Brazil (Randall and Bohlke, 1965, p. 241;
Cervigon, 1966, p. 605).

Halichoeres poeyi (Steindachner). British Honduras: Lighthouse Reef,
11457 (1, 35.9). Reported from Florida, the Bahamas, Greater and Lesser
Antilles, and Brazil (Randall and Bohlke, 1965, p. 250).

GoBIIDAE

Lythrypnus elasson Bohlke and Robins. Yucatan: Cozumel Island, 9524 (7,
10.5—12.5). Previously reported only from the Bahamas (Bohlke and
Robins, 1960, p. 79).

Quisquilius hipoliti (Metzelaar). Nicaragua: Albuquerque Cays, 23094 (1,
18.0), 23093 (1, 13.2), 23095 (4, 13—19); British Honduras: Turneffe
Island, 9572 (1, 18.5), 9802 (4, 11.1—19.3), Lighthouse Reef, 9419 (1,
10.3), 9463 (4, 9.4—14.0); Yucatan: Cozumel Island, 9218 (2, 9—12),
9706 (4, 9.8—18.3), 9519 (9, 10—19.5), Banco Chinchorro, 9620 (6,
9.0—15.5), 9369 (1, 13.0). Reported from Florida, the Bahamas, Puerto
Rico, Lesser Antilles, and Venezuela (Bohlke and Robins, 1960, p. 88;
Cervigon, 1966, p. 748).

! CHINCHORRO

'} TURNEFFE ISLAND

(1S LIGHTHOUSE REEF

Brea! Jt ]
, Ws:
13.

o

¢ COURTOWN
: CAYS

ALBUQUERQUE
CAYS

Fig. 1. Map of collection area. Nicaragua (Albuquerque and Courtown
Cays) 10 collections; British Honduras (Turneffe Island, Belize, Lighthouse
Reef) 13 collections; Yucatan (Cozumel Island and Banco Chinchorro) 15 col-

lections.

190 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SCORPAENIDAE

Scorpaena albifimbria Evermann & Marsh. Nicaragua: Albuquerque Cays,
23062 (1. 36.0). Previously recorded from Florida, the Bahamas, Greater
and Lesser Antilles (Eschmeyer, 1965, p. 118).

OPISTHOGNATHIDAE

Opisthognathus maxillosus Poey. Nicaragua: Albuquerque Cays, 23092 (2,
75.0, 103.5); Yucatan: Banco Chinchorro, 9300 (1, 28.6). Reported from
Florida, the Bahamas, and the Greater and Lesser Antilles (Randall, MS).

Opisthognathus whitehursti (Longley). British Honduras: Turneffe Island,
9860 (1, 28.4), 9839 (1, 33.6), Lighthouse Reef, 11463 (1, 42.2). Re-
ported from Florida, the Bahamas, and the Greater and Lesser Antilles
(Randall, MS).

BLENNIIDAE

Hypleurochilus springeri Randall. Nicargua: Albuquerque Cays, 23091 (1,
21.6). Reported from Florida, the Bahamas, the Greater and Lesser An-
tilles, Grand Cayman, and Venezuela (Randall, 1966, p. 66).

CLINIDAE

Acanthemblemaria aspera (Longley). British Honduras: Lighthouse Reef,
9481 (1), 9554 (1). Reported from Tortugas, the Bahamas, Haiti, and
Venezuela (Stephens, 1963, p. 35; Cervigon, 1966, p. 689).

Acanthemblemaria chaplini Bohlke. Yucatan: Banco Chinchorro, 9375 (1).
Previously reported from the Bahamas (Stephens, 1963, p. 34).

Enneanectes altivelis Rosenblatt. Nicaragua: Courtown Cays, 23055 (4,
11.5—24.7), 23059 (1, 21.0), Albuquerque Cays, 23058 (3, 13:5—17.7);
British Honduras: Lighthouse Reef, 9490 (3), Turneffe Island, 9566 (4),
10306 (2); Yucatan: Banco Chinchorro, 9361 (1), 9626 (2), 9795 (2).
Reported from the Bahamas (Rosenblatt, 1960, p. 21).

Enneanectes atrorus Rosenblatt. British Honduras: Turneffe Island, 9852
(2). Reported from the Bahamas (Rosenblatt, 1960, p. 11).

Enneanectes pectoralis (Fowler). Nicaragua: Courtown Cays, 23049 (4,
11.2—16.5); British Honduras: Turneffe Island, 9571 (1); Yucatan:
Banco Chinchorro, 9370 (1). Reported from Florida, the Bahamas, the
Lesser Antilles, and Venezuela (Rosenblatt, 1960, p. 15; Cervigén, 1966,
p. 681).

Emblemaria diaphana (Longley). British Honduras: Lighthouse Reef, 9285
(1), 9499 (1). Reported from the Bahamas (Stephens, 1963, p. 94).

Emblemaria bahamensis (Stephens). Nicaragua: Albuquerque Cays, 23051
(1, 16.0). Reported from the Bahamas (Stephens, 1963, p. 94). The

BiIRDSONG AND EMERY: New Records of Fishes 191

status of one new record included here is not clear: Emblemaria baham-
ensis may be a synonym of E. diaphana (Stephens, pers. comm.), but both
forms were taken in this series of collections, previously having been known
only from the Bahamas.

Labrisomus haitiensis Beebe and Tee-Van. British Honduras: Lighthouse
Reef, 9492 (2, 34.0, 35.6), 9412 (3, 29.0—38.5); Yucatan: Banco Chin-
chorro, 9646 (3, 32.6—37.0). Reported from Florida, the Bahamas, Haiti,
and Virgin Islands (Springer, 1959a, p. 430; 1959b, p. 291).

Labrisomus nigricinctus (Rivero). Nicaragua: Courtown Cays, 23052 (2,
41.1, 45.6); Yucatan: Banco Chinchorro, 9350 (2, 41.3, 44.2), 9298 (1,
40.8). Reported from Florida, the Bahamas, Cuba, Virgin Islands, Ja-
maica, Haiti, Puerto Rico, and Barbados (Springer, 1959a, p. 439; 1959b,
jos PASE).

Malacoctenus boehlkei Springer. Nicaragua: Albuquerque Cays, 23050 (3,
31.1—45.3); British Honduras: Turneffe Island, 9865 (1). Reported
from the Bahamas and Virgin Islands (Springer, 1959a, p. 444; 1959b, p.
Oi)

Malacoctenus erdmani Smith. Nicaragua: Albuquerque Cays, 23087 (3,
17.3—25.4); British Honduras: Lighthouse Reef, 9467 (1, 17.0); Yucatan:
Banco Chinchorro, 9366 (1, 10.5). Reported from the Bahamas, Cuba,
Puerto Rico, Jamaica, Haiti, and Barbados (Springer, 1959a, p. 448; 1959b,
p. 291).

Malacoctenus versicolor (Poey). British Honduras: Lighthouse Reef, 9996
(1, 62.4). Reported from the Bahamas, Cuba, Puerto Rico, Virgin Islands,
Jamaica, Barbados, and Tobago (Springer, 1959a, p. 456; 1959b, p. 271).

Starksia atlantica Longley. Nicaragua: Courtown Cays, 23061 (9, 12.0—
18.4), 23060 (1, 16.9); British Honduras: Turneffe Island, 10300 (1, 9.6),
9854 (6, 9.9—15.7), 9558 (7, 11.6—16.0), Lighthouse Reef, 9497 (1,
9.8); Yucatan: Banco Chinchorro, 9760 (2, 15.5, 16.2). Reported from
the Bahamas (Bohlke and Springer, 1961, p. 34).

Starksia lepicoelia Bohlke and Springer. Nicaragua: Albuquerque Cays,
23048 (1, 17.6); British Honduras: Turneffe Island, 9837 (4, 14.9—26.6),
9546 (4, 16.3—22.9), Lighthouse Reef, 9474 (14, 9.9—24.1), 9411 (1,
16.4); Yucatan: Banco Chinchorro, 9340 (1, 14.7), 9629 (9, 14.0—22.7),
9755 (1, 17.6). Reported from the Bahamas and Virgin Islands (Bohlke
and Springer, 1961, p. 40).

Starksia nanodes Bohlke and Springer. Nicaragua: Albuquerque Cays, 23046
(1, 10.5); British Honduras: Turneffe Island, 9576 (1, 19.0). Reported
from the Bahamas and the Virgin Islands (Bohlke and Springer, 1961, p.
44).

Starksia y-lineata Gilbert. Nicaragua: Courtown Cays, 23057 (1, 13.8).
Reported from Grand Cayman Island (Gilbert, 1961, p. 1).

192 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

List oF SPECIES COLLECTED

The following list compiles all the species collected on the two
expeditions. Some 209 species are listed; those listed as unidentified
are referable to an undescribed species. A code follows each
species name indicating the locations from which it was taken.
Collections from Albuquerque and Courtown Cays off Nicaragua,
are designated “A”; collections from Turnefte Island, Lighthouse
Reef and Belize, British Honduras, are designated “B”; collections
from Banco Chinchorro and Cozumel Island, Yucatan, are desig-
nated “C”. Identifications of Yucatan and British Honduras material
were largely made by C. R. Robins in 1961.

Abulidae; Albula vulpes (C). Moringuidae; Moringua edwardsi (A, B, C).
Xenocongridae; Chilorhinus suensoni (A, C), Kaupichthys atlanticus (A, B).
Muraenidae;Echidna catenata (B), Enchelycore sp. (A), E. nigricans Us )).
Gymnothorax moringa (A, B), G. vicinus (A, B), Uropterygius diopus (C).
Congridae; Ariosoma impressa (C). Ophichthidae; Ahlia egmontis (A, B),
Myrichthys acuminatus (A, B, C), Myrophis punctatus (B, C). Clupeidae;
Harengula humeralis (A). Synodontidae; Synodus synodus (A, B, C). Mycto-
phidae; Diaphus dumerili (A), D. elucens (A).

Batrachoididae; Opsanus astrifer (B). Gobiesocidae; Acyrtops beryllinus
(B), Acyrtus artius (B, C), Gobiesox punctulatus (B), Tomicodon fasciatus
(B). Antennariidae; Antennarius scaber (B). Ophidiidae; Otophidium dormi-
tator (C), Ogilbia sp. (A, B, C), Petrotyx sanguineus (B, C). Belonidae;
Platybelone argalus (A), Strongylura notata (B), S. timucu (B). Cyprino-
dontidae; Cyprinodon veriegatus (C), Floridichthys carpio (C), Garmanella
pulchra (C). Poeciliidae; Poecilia sphenops (B). Atherinidae; Allanetta har-
ringtonensis (C), Atherinomorus stipes (A, B). Holocentridae; Holocentrus
ascensionis (A), H. coruscus (B, C), H. marianus (A). H. rufus (A, B,C),
H. vexillarius (A, B, C), Myripristis jacobus (A, C), Plectrypops retrospinis
(A, B, C). Aulostomidae; Aulostomus maculatus (A, B, C). Syngnathidae;
Corythoichthys brachycephalus (B, C), Syngnathus floridae (C).

Scorpaenidae; Scorpaena albifimbria (A), S. brasiliensis (C), S. calcarata
(B), S. inermis (C), S. plumieri (A), Scorpaenodes caribbaeus (A, B, C).
Dactylopteridae; Dactylopterus volitans (C). Serranidae; Alphestes afer (A),
Cephalopholis fulvua (A, C), Liopropoma rubre (C), Epinephalus adscensionis
(A), E. guttatus (A), petrometopon cruentatum (A, B, C), Serranus tabacarius
(C), S. tigrinus (B, C). Grammistidae; Pseudogrammus gregoryi (A, B, C),
Rypticus brachyrhinus (B), R. saponaceus (A), R. subbifrenatus (B, C). Gram-
midae; Gramma loreto (A, B, C), G. melacara (B). Priacanthidae; Priacanthus
cruentatus (A). Apogonidae; Apogon alutus (C), A. binotatus (A, B), A.
conklini (A, B, C), A. lachneri (A, B,C), A. maculatus (A, B, C), A. pigmen-
tarius (A, B, C), A. planifrons (A, C), A. sp. (A, B), A. quadrisquamatus (A,
C), A. townsendi (A, B, C). Carangidae; Caranx bartholomaei (A), C. ruber

BiIRDSONG AND EMERY: New Records of Fishes 193

(A). Lutjanidae; Lutjanus apodus (A, B, C), L. griseus (B), L. mahogoni
(A). Gerreidae; Eucinostomus argenteus (B, C), E. havana (C), Gerres cin-
ereus (B). Pomadasyidae; Haemulon carbonarium (A), H. chrysargaereum
(A), H. flavolineatum (A, B, C), H, parrai (A, B), H. plumieri (A), H.
sciurus (A). Sparidae; Archosargus rhomboidalis (B). Sciaenidae; Bairdiella
ronchus (B), Equetus lanceolatus (B), E. punctatus (A, B, C). Mullidae;
Mulloidichthys martinicus (A), Pseudupeneus maculatus (A, B, C). Chaeto-
dontidae; Chaetodon capistratus (B, C), C. striatus (A), Holocanthus ciliaris
(A), H. tricolor (C), Pomacanthus arcuatus (A). Pomacentridae; Abudefduf
saxatilis (A, B, C), Chromis multilineata (B, C), C. cyanea (A, B, C),
Eupomacentrus sp. (A, B, C), E. fuscus (A, B, C), E. leucostictus (A, B, C),
E. partitus (A, B, C), E. planifrons (A, B, C), E. variabilis (B, C), Micro-
spathodon chrysurus (A, B, C). Cirrhitidae; Amblycirrhitus pinos (B, C).
Mugilidae; Mugil trichodon (B). Sphyraenidae; Sphyraena barracuda (C).
Labridae; Bodianus pulchellus (A), B. rufus (C), Clepticus parrai (C), Do-
ratonotus megalepis. (B), Halichoeres bivittatus (A, B, C), H. garnotti (A, B,
C), H. maculipinna (A, B, C), H. pictus (B, C), H. poeyi (B), H. radiatus
(A, C), Hemipteronotus splendens (A), Thalassoma bifasciatum (A, B, C).
Scaridae; Nicholsina usta (C), Scarus coeruleus (B), S. croicensis (A, B, C), S.
guacamaia (C), S. taeniopterus (A), S. vetula (C), Sparisoma aurofrenatum
(A,B; ©), S. radians (A), S. rubripinne (A, B, C), S. viride (A, B, C).
Opisthognathidae; Opisthognathus maxillosus (A, C), O. whitehursti (B).
Dactyloscopidae; Dactyloscopus tridigitatus (B, C), Heteristius rubrocinctus
(A, B, C). Uranoscopidae; Astroscopus y-graecum (B). Blenniidae; Ento-
macrodus nigricans (A, B, C), Hypleurochilus springeri (A), Ophioblennius
atlanticus (A, B, C). Clinidae; Acanthemblemaria aspera (B), A. chaplini
(C), A. spinosa (A), Emblemaria bahamensis (A), E. diaphana (B), En-
neanectes altivelis (A, B, C), E. atrorus (B), E. boehlkei (A, B, C), E. pec-
toralis (A, B, C), Labrisomus albigenys (B), L. bucciferus (A, B, C), L.
gobio (A, B, C), L. guppyi (A, B, C), L. haitiensis (A, B, C), L. kalisherae
(A), L. nigricinctus (A, C), L. nuchipinnis (B), Malacoctenus aurolineatus
(C), M. boehlkei (A, B), M. delalandei (B), M. erdmani (A, B, C), M. gilli
(A, B, C), M. macropus (A, B, C), M. triangulatus (A, B, C), M. versicolor
(A, B), Paraclinus fasciatus (B), P. nigripinnis (A, B), Starksia atlantica (A,
B, C), S. lepicoelia (A, B, C), S. nanodes (A, B), S. sluiteri (A, C), S. y-
lineata (A), Stathmonotus stahli (B, C). Gobiidae; Bathygobius mystacium
(A), B. soporator (B), Coryphopterus dicrus (B), C. glaucofraenum (A, B,
C), C. hyalinus (B, C), Gobionellus boleosoma (B), G. fasciatus (C.), Gobio-
soma (Elacatinus) sp. (A, B, C), G. Tigrigobius dilepis (A, B), G. Tigrigobius
gemmatum (B), G. Tigrigobius pallens (A, B), Gnatholepis thompsoni (A, B,
C), Lophogobius cyprinoides (B), Lythrypnus elasson (A, C), L. hetero-
chromis (A, B), Quisquilius hipoliti (A, B, C). Eleotridae; Erotelis smaragdus
(B). Acanthuridae; Acanthurus bahianus (A, B, C). A. chirurgus (A), A.
coeruleus (A, B, C). Bothidae; Bothus lunatus (C), B. maculiferus (C), B.
ocellatus (A, B, C); Balistidae; Balistes vetula (B), Xanthichthys ringens (C).
Monocanthidae; Cantherines pullus (A), Monocanthus ciliatus (C), M. tuckeri
(A, C), Tetraodontidae; Canthigaster rostrata (A, B, C).

194 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

DIscussION

The Caribbean coast of Central America has been visited by
several expeditions which collected fishes, but until recently no
concentrated effort has been made to study the area ichthyological-
ly, and its fish fauna is poorly known. Shore and trawl collections
have been made by the Bureau of Commercial Fisheries (Bullis
and Thompson, 1965), but few collections have been made in
coral reef areas. Meek and Hildebrand (1923-1928), Fowler (1944),
Caldwell, Ogren, and Giovannoli (1959), Caldwell and Caldwell
(1964) and Caldwell (1963) all contain valuable discussions of
the fishes in the area, but no general work has recently summarized
the information since Fowler (1944).

Considering the small number of collections (38) included
here, all taken in readily accessible areas, it is significant to find at
least 25 new distributional records. Some of these considerably
enlarge the known ranges of the species. The wrasses, one goby
(Quisquilius hipoliti), the scorpaenid, and the opisthognathids were
known to occur widely in Florida, the Bahamas, and the Antillean
regions. Others, such as the clinids in the genera Acanthemble-
maria, Enneanectes, Emblemaria, Malacoctenus, and particularly
Starksia were known only from the Bahamas or restricted areas
nearby.

The discovery of these forms in the British Honduras and Ni-
caragua area indicates a more general distribution of these groups
throughout the Caribbean. Many of these species are small and
are probably transported over uninhabitable areas as larvae. It is
likely that many more coral reef species will be found to be widely
distributed over this region. The only factor restricting the distri-
bution of a given species may well be the lack of a suitable habitat
rather than the lack of means of dispersal.

ACKNOWLEDGMENTS

We thank the following: C. Richard Robins, Institute of Marine
Sciences, University of Miami, for identification of specimens and
review of the manuscript; James E. Bohlke, Academy of Natural
Sciences of Philadelphia, John E. Randall, Bernice P. Bishop Mu-
seum, Honolulu, and John S. Stephens, Jr., Occidental College, Los
Angeles, for identification of specimens; Frederick H. Berry, Bureau

BIRDSONG AND EMERY: New Records of Fishes 195

of Commercial Fisheries, Tropical Atlantic Biological Laboratory,
Miami, for review of the manuscript.

This study was supported in part by NSF contract GB-1450, C.
R. Robins, principal investigator, and NSF contract G-5012, C.
Emiliani principal investigator.

LITERATURE CITED

BOHLKE, JAMES E.., AND C. RicHARD Rosins. 1960. Western Atlantic gobioid
fishes of the genus Lythrypnus, with notes on Quisquilius hipoliti and
Garmannia pallens. Proc. Acad. Nat. Sci. Philadelphia, vol. 112, no. 4,
pp. 13-101, 2 figs,, 3 pls.

BOHLKE, JAMES E., AND Victor G. SPRINGER. 1961. A review of the Atlantic
species of the clinid fish genus Starksia. Proc. Acad. Nat. Sci. Phila-
delphia, vol. 113, no. 3, pp. 29-60, 15 figs.

Butiis, Harvey R., AnD JoHN R. THompson. 1965. Collections by the ex-
ploratory fishing vessels Oregon, Silver Bay, Combat, and Pelican made
during 1956-1960 in the southwestern North Atlantic. Special Scientific
Rep. Fisheries, no. 510, pp. 1-130.

CALDWELL, Davin kK. 1963. Marine shore fishes from near Puerto Limon,
Caribbean Costa Rica. Contrib. in Sci. Los Angeles Co. Mus., vol. 67,
pp. 1-11, 2 figs.

CALDWELL, Davin K., AND MELBA C. CAaLpweLi. 1964. Fishes from the
southern Caribbean collected by Velero III in 1939. Allan Hancock
Atlantic Expedition Rept. vol. 10, pp. 1-61, 2 pls.

CALDWELL, Davin K., Larry H. OGREN, AND LEONARD GIOVANNOLI. 1959.
Systematic and ecological notes on some fishes collected in the vicinity
of Tortuguero, Caribbean Coast of Costa Rica. Rev. Biol. Trop. vol: no.
il, oon 788, OieS. |

CERVIGON, FERNANDO M. 1966. Los peces marinos de Venezuela. Estacién
de Investigaciones Marinas de Margarita, Fundacién La Salle de Cien-
cias Naturales, Monogr. no. 11, pp. 1-951, 385 figs, 1 map, 2 vols.

ESCHMEYER, WILLIAM N. 1965. Western Atlantic scorpionfishes of the genus
Scorpaena, including four new species. Bull. Mar. Sci., vol. 15, no. 1,
pp. 84-164, 12 figs.

Fow Ler, Henry W. 1944. Results of the fifth George Vanderbilt Expedition
(1941) (Bahamas, Caribbean Sea, Panama, Galapagos, Archipelago,
and Mexican Pacific Islands). Monographs Acad. Nat. Sci. Philadel-
phia, no. 6, pp. vi + 1-583, 265 figs., 20 pls.

GILBERT, Carter R. 1965. Starksia y-lineata, a new clinid fish from Grand
Cayman Island, British West Indies. Notulae Naturae. vol. 379, pp. 1-6,
1 fig.

196 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MEEK, SETH E., AND SAMUEL F. HILDEBRAND. 1923-1928. The marine fishes
of Panama. Field Mus. Nat. Hist., Zool. Ser., vol. 15, pts. 1-3.

MILLIMAN, JoHN D., Conran V. W. MAHNKEN, DONALD R. Moore, AND JOHN
A. JonEs. In press. The ecology and productivity of two southwestern
Caribbean atolls, Courtown Cays and Albuquerque Cays.

RANDALL, JOHN FE. 1966. The West-Indian blennid fishes of the genus
Hypleurochilus, with the description of a new species. Proc. Biol. Soc.
Washington vol. 79, pp. 57-71, 1 fig.

RANDALL, JOHN E., AND JAMES E. BOHLKE. 1965. Review of the Atlantic
labrid fishes of the genus Halichoeres. Proc. Acad. Nat. Sci. Philadel-
phia, vol. 117. no. 7, pp. 235-259, 11 figs.

Ropins, C. RicHARD, AND JAMES FE. BOHLKE. 1964. Two new Bahama gobiid
fishes of the genera Lythrypnus and Garmannia. Notulae Naturae 364:
1-6, 2 figs.

ROSENBLATT, RicHAarD H. 1960. The Atlantic species of the blennioid fish
genus Enneanectes. Proc. Acad. Nat. Sci. Philadelphia, vol. 112, no. 1,
pp. 1-23, 4 figs.

SPRINGER, Vicror G. 1959a. Systematics and zoogeography of the clinid
fishes of the subtribe Labrisomini Hubbs. Univ. Texas, Inst. Mar. Sci.
Publ., vol. 5, pp. 417-492, 4 figs, 7 pls.

—. 1959b. A new species of Labrisomus from the Caribbean Sea, with
notes on other fishes of the subtribe Labrisomini. Copeia, 1959, no. 4,
pp. 289-292, 1 fig.

STEPHENS, JOHN S., Jr. 1963. A revised classification of the blennioid fishes

of the American family Chaenopsidae. Univ. California, Publ. Zool.,
VOlOS sivas. Goppp:.lelates alors:

Institute of Marine Sciences, University of Miami, Rickenbacker
Causeway, Miami, Florida 33149. Contribution No. 956.

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968)

Cuban Lizards of the Genus Chamaeleolis
OrLANDO H. GARRIDO AND ALBERT SCHWARTZ

THE endemic Cuban lizards of the genus Chamaeleolis Du-
méril and Bibron are among the most bizarre of the anoline
lizards in the West Indies. Reaching snout-vent lengths somewhat
in excess of 170 mm, Chamaeleolis is structurally characterized by
its prominent bony head casque (a feature which it shares with
Anolis equestris Merrem) and its peculiar body scalation, consist-
ing of large, flat, irregularly shaped and sized dorsal and lateral
scales. The behavior of Chamaeleolis is equally peculiar; in
contrast to the agility of the similarly sized A. equestris, Chamaele-
olis is deliberate and slow, in the fashion of some of the lower
primates and Old World chamaeleontids. Wilson (1957), com-
menting on an individual in captivity, noted that the lizard “could
be left on a laboratory table without much danger of wandering
far away; it would often remain in the same spot for hours or even
days without changing its position.” Its lack of aggressiveness was
demonstrated by Wilson’s placing a live male A. equestris and a
cardboard model resembling a male Chamaeleolis near it without
eliciting any overt response. The lizard might best be described
as extremely lethargic and phlegmatic; the few specimens ob-
served in the field by the junior author confirm the sluggishness of
Chamaeleolis, and Ruibal (1964, p. 482) noted that specimens
secured by him likewise did not attempt to escape.

Possibly owing at least in part to its slowness and consequent
inconspicuousness, Chamaeleolis has always been rare in herpeto-
logical collections made in Cuba. Stejneger (1917, pp. 267-268 )
listed the specimens in the United States National Museum; Barbour
and Ramsden (1919, p. 129) noted its widespread occurrence in
Cuba but regarded it as rare. Ruibal (1964, p. 480) listed
the genus as occurring in all the provinces of Cuba. The rarity
of Chamaeleolis in collections has formerly precluded any analysis
of variation in the genus, and it has been to our great good
fortune that we have been able to bring together a very large
percentage of specimens in both Cuban and American collections
(which, interestingly, nicely supplement one another as far as
geographical coverage of Cuba is concerned). We have examined
specimens in the American Museum of Natural History (AMNH),

198 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

Museum of Comparative Zoology (MCZ), Museum of Zoology,
University of Michigan (UMMZ), and United States National
Museum (USNM )—a total of 30 lizards; for the opportunity to
study these specimens we are indebted to Charles M. Bogert and
George W. Foley, Ernest E. Williams, Charles F. Walker, Doris
M. Cochran, and James A. Peters. A similar courtesy has been
extended us by the curators of several Cuban collections: we have
examined material from the Museo Poey at the Universidad de la
Habana (MP), including those formerly housed at the Universidad
de Oriente (UO), and the collections of Miguel L. Jaume (MJ)
and Mario S. Buide (MSB). We are grateful to Carlos Storch,
Dario Guitart, and Vitelio Vitier, as well as to Srs. Jaume and
Buide, for allowing us to examine the specimens in their care.
Additionally, Héctor Sagué D., director of the Instituto de Biologia,
Academia de Ciencias, has greatly facilitated our work; speci-
mens in the collection of the Instituto de Biologia are designated
by IB. There are 38 Chamaeleolis in Cuban collections, bringing
the number of specimens examined by us to a total of 68. Two
specimens have been placed in the Albert Schwartz Field Series
(ASFS). No previous workers have examined conjointly such a
number of these rare lizards.

We wish also to thank Edmond V. Malnate both for his infor-
mation on the specimens of Chamaeleolis in the Academy of
Natural Sciences of Philadelphia (ANSP) and for his assistance
with literature. Similar information has been generously pro-
vided by Jean Guibé on original material in the Muséum National
d'Histoire Naturelle (MNHN) in Paris. Margaret S. Shaw at
the American Museum and Thomas W. Schoener at Harvard Uni-
versity have been extremely helpful in the matter of literature.
The junior author was able to carry out his collecting in Cuba
between 1957 and 1960 with the aid of two National Science
Foundation grants (G-3865 and G-6252) which are hereby grate-
fully acknowledged. The illustrations are the work of David C.
Weber:

The present paper stems from observations made by the senior
author while comparing material collected by himself and by
Charles T. Ramsden (housed today in the Universidad de Oriente )
with specimens from more western Cuban localities. Even casual
study convinced him that Chamaeleolis, rather than being mono-

GARRIDO AND SCHWARTZ: Cuban Lizards 199

typic as had been uninimously held, was composed of two distinct
species, which differed strikingly in several characters. This in-
formation was relayed to the junior author who in turn borrowed
the Chamaeleolis in American collections and quickly confirmed
the existence of two species. The differences between them are
remarkable and very distinctive; surely only the lack of abundant
comparative material has prevented previous workers (with the
notable exception of Cope) from reaching the same conclusion.
The junior author had unwittingly collected specimens of both
species (four of one, one of the other) and was unaware, like
others, that differences observed between them were anything more
than individual variation. Such is not the case, and the two
species are readily diagnosible on the basis of many characters
of scutellation and apparently by some phases of their respective
color repertories and patterns.

Four trivial names have been applied to these distinctive
lizards. The earliest name is chamaeleonides Duméril and Bi-
bron, 1837, followed by fernandina Cocteau, 1838 or 1839 (we
follow, in dating names in de la Sagra’s Historia fisica, politica y
natural de la isla de Cuba, the rationale outlined by Smith and
Grant, 1958, for the earliest date for the Spanish edition and
attribute the name solely to Cocteau rather than to Cocteau and
Bibron in accordance with Smith and Grant’s review), cocteawi
Fitzinger, 1843, and porcus Cope, 1864. The generic name, al-
though formally proposed and diagnosed by Cocteau and Bibron,
had been previously used by Duméril and Bibron (1837, pp. 168,
171) in reference to Cocteau’s work; this previous usage seems
adequate, in accordance with Articles 11 and 12 of the Interna-
tional Code, to attribute the generic name Chamaeleolis to Du-
méril and Bibron, as has been done by Romer (1956, p. 535).
The apparent anomaly of Duméril and Bibron referring to an
(apparently) as yet unpublished name may be resolved by con-
sidering the fact that Bibron worked with Cocteau on the reptile
section of de la Sagra and was thus familiar in advance with any
nomenclatural changes to be made in the latter work. It is per-
haps fortunate that the generic name Chamaeleolis can be at-
tributed to Duméril and Bibron, since in the Spanish edition of
de la Sagra, the name is spelled variously—Chamoelolis (pp. 72
and 73, in a discussion of Cuban anoline lizards), Chamaeleolis

200 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

(p. 90, where the generic name is formally proposed), and
Chamaeolis (p. 90, where the trivial name fernandina is proposed).
We would also like to point out that many authors have spelled
the trivial name “chamaeleontides” and have attributed this
variant to Duméril and Bibron. There is no justification for this
spelling since Duméril and Bibron spelled the specific epithet
without the t, and we have followed their original orthography.

At least one of the four proposed trivial names (porcus) is
readily identifiable from the original description. Mr. Malnate
has checked the holotype for us and found that it agrees excellently
with the diagnosis proposed by Cope (1864, p. 168). The three
older names (chamaeleonides, fernandina, cocteaui) are less clear-
ly identifiable on the basis of their original descriptions, cocteaui
being especially so. However, Dr. Guibé (in litt., 22 November
1966) advised us that these three names are based upon the same
specimen (MNHN 1004), so that there is no question of either
fernandina or cocteaui being anything other than direct synonyms
of the prior chamaeleonides.

There is some oblique evidence that fernandina Cocteau may
have priority over chamaeleonides Duméril and Bibron. These
latter authors (1837, p. 168, 171) referred to Chamaeleolis Fern-
andina Cocteau and commented that “Notre Anolis caméleonide
est pour M. Th. Cocteau le type d'un genre particulier auquel il
assigne pour principal caractére d’ avoir lécaillure ventrale granu-
leuse. Le nom par lequel il le désigne est celui de Chamaeleolis.”
Such a statement clearly implies that Duméril and Bibron were
aware of Cocteau’s (proposed) generic name as noted before.
Dumeril and Bibron (p. 168) mentioned the name as appearing
in table 12 of the French edition of the de la Sagra work, but
made no further comment on pagination of the Cocteau descrip-
tion in the French edition. Whether their reference was due to
foreknowledge of Cocteau’s work (since Cocteau and Bibron
worked together on the reptile section of de la Sagra) or to their
having seen it in print is equivocal. Since Smith and Grant (1958)
have gone to considerable effort to establish dates and authorships
for published names in de la Sagra’s Historia, and since their re-
search indicates that Duméril and Bibron antedate de la Sagra by
a year, it seems pointless to pursue this line of reasoning further.
It is possible that fernandina Cocteau antedates chamaeleonides

GARRIDO AND SCHWARTZ: Cuban Lizards 201

Dumeril and Bibron, but rather than assume this as fact, we have
adhered to the dating of these two works by Smith and Grant and
continue to use chamaeleonides as the prior name.

Although the two species differ from one another in several
scale characters, the most easily recognized feature is the differ-
ence between the median gular scales. Both sexes of Chamaeleo-
lis have a large, wrinkled, and minutely scaled dewlap. Be-
ginning at the chin and extending posteriorly to about the anterior
distensible portion of the dewlap proper there is a pair of enlarged
paramedian rows of scales. In one species (Fig. 1, left), these
scales are short, squat, conical or more or less pyramidal, and set
next to each other in an uninterrupted series except for occasional
smaller scales. In the other species (Fig. 1, right), the para-
median gulars are elongate, flexible, barbel-like, and separated

Fig. 1. Diagrammatic representation of paramedian gular scales (left) in
Chamaeleolis chamaeleonides, based upon USNM 27497 and (right) in Cham-
aeleolis porcus, based upon USNM 29837.

202 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

from each other by several to many much smaller tiny granular
or irregularly shaped scales. These two types of paramedian gular
scale arrangements are correlated with a constellation of other
features of scutellation which show no intermediates. We have no
doubt whatsoever that we are dealing with two distinct species
which appear to be broadly sympatric over much of Cuba (with
the interesting and possibly significant exception of much of
Oriente Province). The distributions of the two species are puz-
zling, and in no one place have both been taken precisely to-
gether (although we attach little importance to this fact since
the total number of specimens is small for such a large island
as Cuba and the maximum number of either species from a single
locality is 10—Bayate, Guantanamo, Oriente Province). Detailed
discussions of the distributions of the species must be preceded by
clarification of the nomenclature.

SYSTEMATIC ACCOUNT

Chamaeleolis chamaeleonides Duméril and Bibron

Anolis chamaeleonides Duméril and Bibron. 1837. Erp. gén., vol. 4, p. 168.

Chamaeolis [sic] fernandina Cocteau. 1838 or 1839, in de la Sagra, His-
toria fisica, politica, y natural de la isla de Cuba, vol. 4, Reptiles y
Peces, p. 90.

Pseudochamaeleon cocteaui Fitzinger, 1843, Syst. Rep., p. 63.

Type locality. Cuba; here restricted to the vicinity of La
Habana, Habana Province, Cuba.

Distribution. Cuba, from Pinar del Rio Province in the west
as far east as southwestern Oriente Province in the east; unknown
east of a line drawn vertically between Gibara in the north and
Pico Turquino in the south (Fig. 2).

Definition. A species of Chamaeleolis characterized by a com-
bination of 1) paramedian gular scales small, truncate, short,
conical to pyramidal (Fig. 1), 2) supralabials separated from
circumorbital ring by one (occasionally two) row of scales, 3)
dorsal scales subcircular to irregularly shaped with many smaller
interstitial scales, 4) ventral scales small to tiny, 42 to 83 in
distance equal to length from tip of snout to anterior bony border
of orbit, 5) zone of transition between large lateral scales and
tiny ventral scales relatively gradual, 6) scales on lower jaw below

‘snauod "5 ‘sapomo ‘sapuoajapuyy9 *D “So[suUeIT,
‘syoajapuDy Dy JO satoods OX} FO WOLNLA}SIp Surmoys ‘eqny jo def °G “ST

204 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

angle of jaw heavily rugose, and 7) posterior expanded portion
of casque with relatively large scales.

Discussion. Throughout its broad range, C. chamaeleonides
is constant in the diagnostic characters noted above. The shape
of the paramedian gulars is variable, but the variation is not
correlated with either sex or geography and seems merely to be
individual. The paramedian gulars have their bases appressed to
one another and are very rarely separated by occasional inter-
mediate scales similar to the small adjacent gulars. The para-
median gulars are never long and filamentous or barbel-like as
they are in the other species. The second most easily determinable
character of C. chamaeleonides is the heavy rugosity of the scales
below the angle of the jaw on the lower mandible. Although
most lateral head scales are somewhat rugose, those in this particu-
lar area are especially so, and the two species can easily be
separated merely by running the finger over the scales below the
angle of the jaw. Two small specimens (snout-vent lenghts 46
and 54 mm) lack these asperities, but all lizards in excess of 90 mm
snout-vent length show them clearly and not appreciably less well
developed than large adults.

The dorsal scales on either side of the dorsal midline do not
differ in size, shape or arrangement from those of the sides; the
more dorsal rows have the scales subcircular or irregular in out-
line, and scattered between them are many tiny interstitial scales.
The lateral scales are like the dorsals in configuration and in
having surrounding granules. The zone of transition between the
lateral scales and the ventrals is more gradual than the condition
in the second species. The ventrals are small and somewhat
variable in size between individuals and are about the same size
as the unspecialized lateral gular scales.

The largest male (USNM 51891), a truly giant and bulky
lizard, has a snout-vent length of 177 mm; the largest female
(MCZ 38419) has an almost equal snout-vent length of 172 mm.
The sexes are easily distinguished externally by the presence
of a pair of conspicuously enlarged postanal scales in the males;
both males and females have dewlaps. The smallest specimen
examined (MCZ 13328, a female, snout-vent length 46 mm) has
the umbilicus still unclosed.

Through the courtesy of Dr. Guibé and the cooperation of Dr.

GARRIDO AND SCHWARTZ: Cuban Lizards 205

Williams we have examined the holotype of Anolis chamaeleonides
Dumeéril and Bibron (MNHN 1004); it is an adult female with a
snout-vent length of 160 mm, short and squat paramedian gulars,
and 61 ventrals in the snout-eye distance. The specimen departs
slightly from our concept of C. chamaeleonides in that the supra-
labials are not quite completely separated from the circumorbital
ring by a complete row of scales on the left side. The anomaly
is doubtless due to the fact that the scales involved are irregular
in configuration in the holotype, probably because of an old injury.

The use of the snout-eye distance as a standard for counting
dorsal scales in Anolis equestris (Schwartz, 1964) has been em-
ployed in counting ventral scales in the present study. The
comments made by Schwartz (1964, p. 408) apply equally well to
making counts of ventral scales in Chamaeleolis. A reasonably
accurate count can be made on specimens which have been pre-
served in an extended position, but material which has been
curled in a jar when preserved inhibits making counts which are
as accurate as we would like. Nevertheless, the rather gross
differences in numbers of ventrals between the two species (de-
spite a small amount of overlap in counts) make us sure that
there is indeed a very distinct difference in number of ventrals
in the snout-eye distance. Mere inspection of specimens of both
species reassures the observer that there is a strong basic difference,
which we have attempted to quantify by using a “standard dis-
tance.”

A criticism of the snout-eye distance is of course the difference
in relative and actual snout length between juveniles and adults.
Small C. chamaeleonides are not mere miniature adults; the casque
is barely indicated and the snout is short and not attenuate as it is
in subadult and full grown lizards. However, the ventral counts
on the two small juveniles in our series (46 and 63 scales) fall
within the parameters established from adult lizards (42 ventrals
at the lower extreme on two females with snout-vent lengths of
157 and 158 mm, 83 ventrals on a male with snout-vent length of
177 mm), so that there is no skewing of data from the inclusion
of these two tiny lizards with their much larger relatives.

Three specimens taken by the junior author and Ronald F.
Klinikowski in 1959 and 1960 at Los Paredones, Camagiiey Prov-
ince, in the Sierra de Cubitas, were encountered at night. A

206 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

small male was sleeping on a limb of a coffee tree about 5 feet
(1.5 meters) above the ground. A large male and a small male
were collected near to one another on a tree limb (the larger
lizard) about 12 feet (3.7 meters) above the ground and on a
pendant vine about 6 feet (1.8 meters) above the ground. In both
these latter instances, the lizards were sleeping on substrata which
about equalled the diameters of the lizards themselves. In no
case was there any attempt at concealment, and the Chamaeleolis
were fully exposed. The specimen from near San Vicente, Pinar
del Rio Province, was collected by a native from an exposed fence
post 3 feet (0.9 meters) above the ground.

Field notes on the living male (AMNH 78073) C. chamaeleo-
nides from San Vicente were as follows: dorsum light gray, venter
pure white with black to gray spots; dewlap white with four
purplish (Pl. 10, I 4; Maerz and Paul, 1950) blotches. Tail with
seven purplish to violet bands and tail tip; legs banded with
purple, toes black. A few scattered purple scales on side of body.
Tongue inky blue posteriorly and anteriorly, with a yellowish
transverse area between the base and blue-black tip. A male from
Los Paredones (AMNH 83625) was dorsally gray with black
reticulations; the legs were faintly greenish and the dewlap very
pale peach at its edge and grayer basally.

An adult female (IB 1149) from Monte Alto near Juragua,
Las Villas Province, had the body grayish and brown with black
spots, vermiculations, and dots. There was a coal-black spot,
larger than about four or five of the smaller dorsal spots combined,
above the axilla. The casque was completely spotted with black
dots, and more widely separated black spots occurred on the face
and sides. The fused eyelids were beige with six or seven brown
rays extending from them. The iris was black with a yellowish
ring. The venter was whitish with leopard-like spots on the sides
at the junction of lateral scales with the smaller ventrals, under
the hindlimbs, in the region of the vent, on the throat, and on
the anterior portion of the dewlap. The spots gave the appearance
of the result from splashings of dirty water, having a color between
clear brown and yellowish. The dewlap was basally brownish vio-
let and distally pale pink. After death, the grayish spots and
reticulations on the head and body became black, so much so
that the head became completely black.

GARRIDO AND SCHWARTZ: Cuban Lizards 207

An adult female (IB 1086) from Cinco Tiras near Juragua,
Las Villas Province, was grayish to beige in life, with a series of
black reticulations over the whole body, most noticeably on the
sides of the venter and along the sides of the dorsal crest. The
limbs were lightly greenish over gray with three poorly visible
stripes on the forelimbs. The casque was reticulated or spotted
with blackish dots and spots. The dorsal reticulations continued
(in brown) on the sides of the face behind the orbits and along
the upper labials. The eyelids were grayish with about six
grayish radiating lines. Behind the head and above the shoulder
there was a blackish spot, narrower below and broader above.
There were no transverse bars on the tail or on the back, although
on the former there were remnants of some darker spots on the
gray ground. The ventral color was sandy white with dirty yellow
spots as far as the throat. The dewlap on either side of the
paramedian scales was spotted with circular leopard-like spots,
some brownish and others yellowish. The ground color of the
dewlap was dull whitish pink, crossed by about six diffuse whitish
bars; the central portions of these bands were marked with grayish
green, and the base of the dewlap was yellowish. The underparts
of the limbs and tail were spotted with a dirty brownish.

A third specimen (a male, ASFS V11171) from the same
region resembled the above descriptions in several features of body
color and pattern. The dewlap, after the lizard had been pre-
served for a week, was recorded as having a yellow ground
crossed by three peach-colored bands.

From the above notes it is obvious that C. chamaeleonides
has a great versatility in its color and pattern repertory. Our
experience is limited, and we are unaware which of the above
data may be individual or which may be due to regional (sub-
specific?) differences in pattern and color. Relatively recently
and carefully preserved C. chamaeleonides regularly show some
sort of throat spotting, either light spots on a dark ground, or dark
spots on a light ground (usually the latter); this is a feature
which we did not observe in any living or preserved specimens
of the second species, and we conclude that the latter lacks this
style of pattern in its repertory. Much additional material will
be needed to demonstrate whether there are indeed subspecies of
C. chamaeleonides based upon pattern and color. As far as our

208 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

present data are concerned, there are suggestions that there may be
such geographically correlated differences in pattern, but the spe-
cies remains uniform in scutellation throughout its range.

C. chamaeleonides occurs from Pinar del Rio Province to west-
ern Oriente Province. It alone has been collected in the proy-
inces of Habana, Matanzas, and Camagiiey; although the second
species is known from one specimen from Pinar del Rio Province,
C. chamaeleonides is the dominant form in western Cuba and is
presumably so in central Cuba at least as far east as Camagtiey
Province. There are specimens from sea-level (La Mulata, Pinar
del Rio Province; Punta de Maya, Matanzas Province) to ele-
vations of about 1000 feet (305 meters) in the Sierra de los
Organos in Pinar del Rio Province and about the same height at
Los Paredones in the Sierra de Cubitas, Camagiiey Province.

Ruibal (1964, p. 482) noted that “C. chamaeleonides” was
restricted to the shaded portions of broadleaf forest; the five
specimens of Chamaeleolis collected by him are all C. chamaeleo-
nides so his remarks can be applied without hesitation to the
habitat of this species. Our experiences confirm the occurrence
of C. chamaeleonides in forested areas, although it does on oc-
casion resort to more open situations (i.e., the individual collected
on an exposed fencepost at San Vicente).

Stejneger (1917, p. 268) reported C. chamaeleonides from two
further localities in Pinar del Rio Province—Cabafias and Cabo
de San Antonio. The specimens confirming these records are no
longer in the United States National Museum, but there is of
course no reason to doubt them. The Cabo de San Antonio
locality is exceptionally interesting, since the area immediately
about the cape is one of xeric coastal woods. Perhaps in extreme
western Cuba, C. chamaeleonides survives in such a habitat in
sharp contrast to the more mesic forests which it usually occupies
elsewhere. There is no assurance that the species at Cabo de
San Antonio is chamaeleonides, but we expect it there in geo-
graphical grounds. It should also be pointed out that the figure
(Stejneger, 1917, fig. 35) of the lateral view of the head of a
specimen (USNM 27502) from San Diego de los Banos, Pinar
del Rio Province, shows the supralabials in contact with the
circumorbital ring—a character which is not typical of C. chamaele-
onides. The drawing is incorrect, and the two rows of scales in

GARRIDO AND SCHWARTZ: Cuban Lizards 209

this specimen are indeed separated by a complete row of small
scales.

Specimens examined. Cuba, Pinar del Rio Prov., Potrerito, Sumidero, 1
(MP); Sierra de Pica Pica, Sumidero, 1 (JB 1084); Banos San Vicente, 1
(MCZ 38419); 2 km N San Vicente, 1 (AMNH 78073); San Diego de los
Banos, 3 (AMNH 58901, USNM 27502-03); La Mulata, 1 (USNM 51891);
Sierra de Anafe, 1 (MJ); Habana Prov., Caimito de Guayabal, 1 (MCZ
13328); Santiago de las Vegas, 2 (MCZ 7895, USNM 27497); Arroyo Ber-
mejo, | (MSB lla); Matanzas Prov., Punta de Maya, 1 (MSB IIb); Las
Villas Prov., kilometer 13 from Santo Tomas, Ciénaga de Zapata, 1 (IB 1085);
Aguada de Pasajeros, 1 (MCZ 8948); Cinco Tiras, near Juragua, 1 (IB 1086);
Monte Alto, 2 km SE Juragudé, 2 (ASFS V11171, IB 1149); Camagiiey Prov.,
Finca San Pablo, ca. 15 km SW Camagiiey, 1 (MCZ 57933); Los Paredones,
Sierra de Cubitas, 4 (AMNH 83625, AMNH 96555-56, MCZ 59329); near
Los Paredones, 1 (MCZ 59330); Oriente Prov., near Buey Arriba, SW of
Bayamo, 1 (MCZ 59331); no locality data other than Cuba, 2 (MNHN 1004,
AMNH 46101). There are three specimens (ANSP 8132, ANSP 11879,
ANSP 11974) with locality data merely “Cuba” which we have not examined
but which Edmond V. Malnate has checked for us and found to represent
C. chamaeleonides.

Chamaeleolis porcus Cope

Chamaeleolis porcus Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia, p.
168.

Type locality: Cuba; here restricted to the vicinity of the city
of Guantanamo, Oriente Province, Cuba.

Distribution: Primarily the eastern two thirds of Oriente Prov-
ince, east of a line drawn vertically between Gibara in the north
and Pico Turquino in the south, but outlying populations occur in
the Sierra de Trinidad in Las Villas Province and in the Sierra del
Rosario in Pinar del Rio Province (Fig. 3).

Definition. A species of Chamaeleolis characterized by a com-
bination of 1) paramedian gular scales long, flexible, barbel-like
(Fig. 1), 2) supralabials in (usually broad) contact with circum-
orbital ring, 3) scales in about four dorsalmost rows longer than
broad (elongate oval in shape), with few or no tiny interstitial
scales or granules, fairly sharply set off from balance of dorsal and
lateral scales which likewise have very few or no _ interstitial
scales between them, 4) ventral scales fairly large, 22 to 48 in
distance equal to length from tip of snout to anterior bony border
of orbit, 5) zone of transition between large lateral scales and
smaller ventral scales relatively abrupt, 6) scales on lower jaw

210 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

below angle of jaw smooth or only very slightly irregular, and 7)
posterior expanded portion of casque with relatively smaller scales.

Discussion. C. porcus occurs, as far as known, in three
widely separated areas. The region of its broadest known dis-
tribution lies in the eastern two thirds of Oriente Province, where
it is the only species known; in Oriente, marginal western stations
for its occurrence are Playa de Guarda la Vaca in the north and
near Los Negros in the south. Thanks to the interest of Charles
T. Ramsden, who resided in the city of Guantanamo and whose
coffee estates were located about that city and between Guan-
tanamo and Santiago de Cuba, there is a splendid series of C.
porcus from this region in Oriente, Material from elsewhere on
the island is sparse; there are two specimens (MCZ 53598, USNM
156786 ) from the Sierra de Trinidad in Las Villas Province and a
single specimen (MP) from Rangel in the Sierra del Rosario in
Pinar del Rio Province. The two Sierra de Trinidad lizards are
the only representatives of the genus from that mountain range,
but both C. chamaeleonides and C. porcus are known from the
Sierra del Rosario, although the former is apparently more com-
mon there than the latter. ©. porcus presents fanmextremely
disjunct distributional pattern, with a main center in eastern
Oriente.

The long, flexible, barbel-like paramedian gular scales at once
separate C. porcus from C. chamaeleonides. No specimen ex-
amined shows any intermediacy in this character, despite, for
instance, the juxtaposition of the two species in the Sierra del
Rosario (Rangel and San Diego de los Banos) or in the Sierra
Maestra (Los Negros and Buey Arriba). The individual elon-
gate paramedian gulars in C. porcus usually do not have their
bases approximated but rather have them separated by an inter-
posed barbel-like scale in the preserved lizard seems to be placed
atop a small raised mount, the slopes of which are covered with
tiny granules. The precise shape of the elongate scales is variable,
some being almost spicule-like and others with wider bases, with
various degrees of intermediacy between these extremes. The
scales are never short and squat as they are in C. chamaeleonides.
There is no sexual difference in development of paramedian gu-
lars, and little ontogenetic change, since juvenile C. porcus (40
mm in snout-vent length) are just as readily identified on this

GARRIDO AND SCHWARTZ: Cuban Lizards Dallil

basis as are much larger specimens. There is also little evidence
of geographic variation in the development of the paramedian
gular scales; the single subadult female from Rangel (snout-vent
length 111 mm) has extremely long paramedian gulars with wide
bases. The two Sierra de Trinidad lizards (males, with snout-
vent lengths of 116 and 160 mm) do not differ in the condition
of the gulars from Oriente specimens. Additional material from
the Sierra del Rosario may show that this western population is
racially distinct from the more eastern populations; the same
may also be true of the Las Villas lizards, although the two
specimens available do not differ appreciably from their Oriente
relatives.

The largest male C. porcus has a snout-vent length of 162 mm,
and the largest female 171 mm. Both species reach about the
same size. On the chance that there might be a difference in
casque development between C. chamaeleonides and C. porcus,
measurements of casque length (from tip of snout to posterior
margin of casque) and width (across widest portion of casque)
were taken on all specimens of both species. Although there is an
impression of greater casque development in C. chamaeleonides,
measurements indicate that there are no differences between the
species (or between the sexes of each species) in this character.
The holotype of C. porcus (ANSP 8133) is a female with a
snout-vent length of 149 mm.

The size of the ventral scales in C. porcus is considerably
larger than those of C. chamaeleonides. The ranges of the counts
in the two species (22 - 48 in porcus, 42 - 83 in chamaeleonides )
overlap slightly, but the overlap is without particular significance
due to the total number of scales involved in each species and to
the possible minor inaccuracies of the counts, as previously noted.
The contact between the supralabials and the circumorbital ring
is a constant feature in C. porcus. Usually the contact is broad,
but in a few specimens a few longitudinally elongate scales are
interposed between the supralabials and the ring at one or both
ends; however, they never completely separate the two scale
features.

The most extensive recently collected series of C. porcus is
that taken by the senior author at La Florida, Oriente Province.
These five specimens showed a duality of coloration; they were

212 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

able to assume a pale phase (brownish gray or greenish gray)
or to become black. All specimens were capable of becoming
black, but in the pale phase two were regularly brownish gray
and the remaining three were regularly greenish gray. Such differ-
ences are not due to the sex of the lizards involved, since there
is a single male and four females in the series. The greenish
lizards had the dewlap orangish yellow basally and medially, and
in the region where the dewlap approached the throat the former
was a bluish gray tone. Four well defined dark brownish bands
crossed the dewlap. The face and sides of the head had irregular
brown markings on a bluish or clear gray ground. The temple
anterior to the ear opening was uniform clear gray. Those lizards
which were brownish gray in the pale phase were an ashy brown.
There was a grayish white supraaxillary patch on the shoulder.
The color of the dewlap was generally the same in these brownish
lizards as in the greenish gray individuals, although the yellow
was brighter. The ventral color, especially near the limbs, was
yellowish, becoming brownish. The tongue was completely black
except for the sides which were grayish. The single juvenile from
this same locality (IB 1093; snout-vent length 65 mm) was colored
identically to the brownish gray adults.

Field notes on an adult female (AHMN 83626) from near
Felicidad, Oriente Province, taken by the junior author, state only
that the dewlap was dull purple; the lizard was collected on the
base of a Roystonea in a very wet coffee plantation surrounded
by hardwood forest.

Although the number of C. porcus available to us is much
greater than C. chamaeleonides, none of the former species shows
any indications of having been dotted or blotched in the fashion of
the latter. It seems likely that, despite a probably wide variety
of colors and patterns in C. porcus, this species never assumes a
spotted or blotched phase. Inspection of specimens of both C.
chamaeleonides and C. porcus does not reveal any constant fea-
tures of pattern which may confidently be used to distinguish
them; considering the known versalitity of C. chamaeleonides in
this respect, it is doubtful that such differences would be determ-
inable from preserved specimens.

The character of the dorsal and lateral scales which we use
to distinguish C. porcus and C. chamaeleonides is difficult to state

GARRIDO AND SCHWARTZ: Cuban Lizards 213

except in broad terms. The more dorsal scale rows in C. porcus
are composed of scales which are elongate and ovid (rather than
subcircular or squarish as in C. chamaeleonides) and either lack
or have only a very few scattered interstitial scales. Below
these dorsal rows (more laterally), the scales become subcircular
and more like those of C. chamaeleonides but continue to have
very few or no interstitial scales. The transition zone between the
enlarged laterals and the much smaller ventrals is, relative to the
condition in C. chamaeleonides, abrupt in C. porcus; although we
are convinced that the difference is real it is difficult to explain
easily and impossible to quantify. It should be noted that we have
not employed, in diagnosing the two species, the number of dorsals
in any set distance on the back or sides. Attempts at making
such counts end in failure, since the variable and intergrading
sizes of the dorsal scales prevents taking any meaningful count.

There may also be differences between the two species in size
and arrangement of head scales; we have partially diagnosed the
species by the relative size of the scales on the expanded portion
of the casque but have not emphasized this presumed difference.
The problem with cephalic squamation is that in any but small
juveniles, the individual head scales become extremely rough; the
boundaries of the individual scales are impossible to determine
because of their rugosity and because of the additionally obscur-
ing effect of cephalic dark pigment. The casque assumes a very
pebbled and dark appearance in many lizards, and the individual
scale boundaries cannot be determined. Our comments on size
of the casque scales depend upon examination of the few juveniles
of the two species where the cephalic scales are not rugose and
their boundaries are clearly determinabie. However, the juvenile
sample of both species is much too small to justify making any
certain or quantitative statement regarding this feature; data
are simply not determinable from subadult or adult Chamaeleolis,
unless they are bleached from long immersion in preservative
with resultant loss of pigment.

The altitudinal distribution of C. porcus ranges from sea-level
(Playa de Guarda la Vaca) to elevations of 2000 feet (610 meters )
in the Oriente mountains (El Yunque de Baracoa; Santa Maria de
Loreto). Most examples are from lower elevations, but this may
indicate only accessibility of habitat to collectors. The fact that

214 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

many of the UO specimens are the result of Ramsden’s interest
in these bizarre lizards with probable advice to workers on his
cafetales to secure whatever specimens were seen is reflected in
the locality data and relatively low to moderate elevations for
the Ramsden material.

The series of C. porcus from La Florida offers some evidence
of habitat and behavior of this species. La Florida lies very near
or in the Sierra de Purial. The region is mountainous and the
area where the lizards were collected is dark and moist; the
forest is so dense that hardly any sunlight penetrates the canopy.
Several of the lizards were in rose-apple (Eugenia jambos L.)
woods and others were encountered on other trees. Associated
anolines included Anolis equestris Merrem, A. allogus Barbour
and Ramsden, and A. angusticeps Hallowell. The animals were
very difficult to locate and with the exception of one which was
surprised climbing on a tree trunk, other individuals tried to
escape in the manner of A. equestris by running up the trunk.
One C. porcus was encountered low on the trunk of a rose-apple;
when the lizard became aware of the approach of the collector,
it scurried to the opposite side of the tree and with great speed
ascended the trunk. The tree was felled, but the lizard was not
found. Another C. porcus ran up the trunk and, upon reaching
the top, hid itself in the terminal leaves of a branch. When
further pursued, this lizard leaped to the ground where it was
captured. The senior author was guided to this precise locality
by Sr. Rubio Suarez of Baracoa (to whom we are most grateful
for assistance). Sr. Suarez stated that La Florida was the only
locality in the Baracoa area where the “chipojo prieto” could be
encountered.

From the above observations, it is apparent that C. porcus
is at least capable of quick and decisive action. If the two
species differed in their escape and activity patterns, these might
as well be used to characterize them. The lethargic specimen
cited by Wilson (1957) from Mina Carlota, Las Villas Province,
is one of the three C. porcus from non-Oriente localities, however.
Thus, C. porcus under some circumstances may be slow and de-
liberate and in others may react in the rapid pattern of Anolis
equestris. There is still no evidence that C. chamaeleonides
is also capable of such speedy activity.

GARRIDO AND SCHWARTZ: Cuban Lizards DIS)

Since natural history data on Chamaeleolis are so very limited,
annotated labels made by Ramsden on two specimens of C. porcus
and observations by the senior author are especially interesting.
Of an adult female from Santa Maria de Loreto, with a snout-vent
length of 170 mm, Ramsden recorded that an egg had been
deposited on 4 September 1948 while the lizard was held in
captivity. Two females from La Florida, collected by the senior
author on 18 September 1965, contained well developed eggs; one
egg measured 23 mm in length and was shelled and ready for
deposition. In color and shape the egg was like that of A. eques-
tris. A subadult male with snout-vent length of 110 mm was
taken on a coffee bush in June 1949. This lizard died in captivity
in September 1949, after having consistently refused a diet of
fruit and roaches. Since Ramsden had found that Anolis and
Leiocephalus readily ate fruit and roaches, he offered these items
to Chamaeleolis, but compietely without success.

Specimens examined. Cuba, Pinar del Rio Prov., Rangel, 1 (MP); Las
Villas Prov., Mina Carlota, Sierra de Trinidad, 1 (MCZ 53598); Topes de
Collantes, 1 (USNM 156786); Oriente Prov., Playa de Guarda la Vaca, 1
(UO); Guarda la Vaca (not mapped), 1 (MCZ 47895); Isla Saetia, 1
(UMMZ 98014); Baracoa, 1 (USNM 29837); near Baracoa, 1 (MCZ 11205);
La Florida, near Baracoa, 6 (IB 1088-89, IB 1091-93, ASFS V11172); El
Yunque de Baracoa, 1000-1800 feet, 1 (MCZ 42507); costa sur, Baracoa,
Punta Caleta, 1 (AMNH 17719); mountains north of Imias, 1 (MCZ 42503);
8 mi. NE Felicidad, 1 (AMNH 83626); Finca “La Rosita” de Borrero, Rio Frio,
El Cobre, 4 (UO); Hermanos Borrero (not mapped), 1 (UO); El Campanario,
Guaso, Guantanamo, 1 (UO); Los Hondones, Guantanamo, 1 (UO); Bayate,
Guantanamo, 10 (UO); Rio Seco wood, San Carlos Estate, Guantanamo, 1
(UO); La Maya, 1 (MCZ 8501); Finca “Tsabelita” de Ramsden, La Maya, 1

(UO); Santa Maria de Loreto, Loma los Ciegos, Ti Arriba, 2000 feet, 2 (UO);
near Los Negros, Jiguani, 1 (MCZ 8459).

DISCUSSION

The distribution of the two species of Chamaeleolis is puz-
zling. The geographic evidence indicates that C. porcus is the
species endemic to the massifs of extreme eastern Oriente Prov-
ince (including, apparently, the Sierra Maestra, whence there is
only the one record from Los Negros). The locus of origin of C.
chamaeleonides is not so clear, since this species is virtually
island-wide except for the eastern portion of Oriente. The abun-
dance of records of C. chamaeleonides in the montane areas of
Pinar del Rio Province in western Cuba suggests that this species

216 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

has evolved there and has spread thence throughout most of Cuba,
to and including the Sierra Maestra where the two species appar-
ently at least approach one another closely. The abundance of rec-
ords for the Sierra de los Organos and Sierra de Rosario in Pinar del
Rio may be due only to the attraction that these spectacularly
scenic ranges offer for the collector. .Absence of records for C.
chamaeleonides throughout most of Matanzas, Las Villas, and
Camagiiey provinces does not imply that the species is absent
there, but rather that much (but by no means all) of the lowland
areas of these provinces has been denuded of its original forest
cover for cultivation. Recently collected specimens from the Cién-
aga de Zapata and near the city of Camagiiey serve to demon-
strate that the lizard is capable of surviving in more remote lowland
areas (the former locality) or in woods in regions which are
otherwise savanna or whence the forest has been removed (the
latter ).

The geographic picture is made more complex, however, by
the occurrence of C. porcus in two outlying situations (the Sierra
del Rosario in Pinar del Rio; the Sierra de Trinidad in Las Villas )
which lie within the known distribution of C. chamaeleonides.
In the Sierra de Trinidad, C. porcus is the only known species
(there are but two specimens of Chamaeleolis known from this
range), but C. porcus appears to be greatly outnumbered by
C. chamaeleonides in the Sierra del Rosario. If C. porcus
is assumed to be strictly montane in its extra-Oriente distribution,
its absence in the Sierra de Cubitas in Camagiiey is puzzling;
that range is apparently inhabited only by C. chamaeleonides.
The Sierra de Cubitas, however, is lower in elevation than both
the Sierra de los Organos-Sierra del Rosario massif on one hand
and the Sierra de Trinidad on the other.

Chamaeleolis is unknown from the Isla de Pinos, but we
imagine that it will ultimately be taken there. The Isla de Pinos
has been so recently separated from the Cuban mainland that it
seems very likely that Chamaeleolis (presumably C. chamaeleo-
nides) reached the Isla de Pinos but remains as yet uncollected.
Such densely wooded areas as the Paso de Piedras and the slopes
of the Sierra de Casas and Sierra de Caballos would seem appro-
priate for these lizards. It is of interest to note that the equally
large Anolis equestris occurs on the Isla de Pinos but has been

GARRIDO AND SCHWARTZ: Cuban Lizards SALT

rather rarely collected there. [Since the present manuscript
was completed, a pair of C. chamaeleonides was secured by Gil-
berto Silva and Jorge de la Cruz on 23 March 1967 at Santa Isabel,
a wooded hill on the north side of the Ciénaga de Lanier.
Santa Isabel is virtually surrounded by the waters of the Ciénaga,
since a tongue of the Ciénaga borders it on the west. The sur-
rounding flat lands are characterized by stands of the palm Coc-
cothrinax miraguano Beccari. |

The genus Chamaeleolis belongs to the Island Alpha group of
anoline lizards (Etheridge, in litt.), but it appears not to be
closely related to other Alpha anoles on the islands in the western
Caribbean (Cuba and Hispaniola). The degree of difference be-
tween Chamaeleolis and most other western Caribbean anolines
(all of which are placed in Anolis with the exception of the
remarkably divergent Chamaelinorops on Hispaniola) suggests
that the genus has had a long and independent history from the
balance of the Alpha group. Chamaeleolis may represent a local
endemic Alpha development on Cuba, a relict member of a group
of such distinctive anolines which has become extinct elsewhere,
or a remnant of a previous invasion of this line of anoline evolution
into Cuba. The usually slow and deliberate movements of Cham-
aeleolis suggest that its dispersal under its own locomotor powers
might be extremely slow. For C. chamaeleonides to have reached
much of Cuba also intimates that long periods of time have
elapsed for such intra-island dispersal to have taken place.

We suggest that the two species of Chamaeleolis are the
result of a very early invasion of Cuba by an Alpha anoline
stock which has differentiated in situ there into two (eastern and
western?) species which have diverged, in several scale and color
repertory characters, to the species level. Both species have
moved out from their regions of origin throughout most of the
island; of the two perhaps C. porcus was primarily lowland and
C. chamaeleonides primarily highland, although this may not have
been the case at all. It is difficult to visualize situations wherein
such usually phlegmatic and deliberate lizards would come into
direct or even indirect competition either with other members of
their own species or those of the other species, once the two have
achieved overlapping ranges. Therefore, it is possible that there
has been a gradually intermixing of the distributions of the two

218 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

species with no dichotomy of habitat (both are forest dwellers )
or elevation (both, in the portions of their ranges where each is
the sole species, occur in the lowlands and at least to intermediate
elevations in the highlands).

The apparently relict western and central distribution of C.
porcus today may well be an artifact of the seeming rarity of
these lizards. The speed with which specimens of Chamaeleolis
become available for study is matched by the customary slowness
of the lizards themselves. Possibly, both C. porcus and C. cham-
aeleonides coexist over much of central and western Cuba, but
the former species remains uncollected there. However, the fact
that C. porcus is known from two isolated mountain massifs in
western and central Cuba suggests that, during the Cenozoic
inundations and emergences of Cuba and its later (pre-Miocene)
status as an archipelago, lowland populations of Chamaeleolis
have been exterminated, but that prior to this time both species
were widespread on the island. Thus, lowland populations (of
both species) were removed and both species were restricted to
upland localities during periods of inundation. Such outliers of
C. porcus on the Sierra del Rosario and Sierra de Trinidad, far
removed from the “parent” population in the massifs of eastern
Oriente, may well be remnants of long previous island-wide dis-
persions of C. porcus. If such an interpretation approaches fact,
then the Pinar del Rio mountains may well have been the place
of origin of C. chamaeleonides and the eastern Oriente ranges that
of C. porcus. The absence of C. porcus on the Sierra de Cubitas
or the more southern Camagiiey Sierra de Nasjasa may mean that
at various times these lower lying ranges have been more com-
pletely submerged than the higher and more persistent areas in
extreme eastern, central, and western Cuba. Once these lower
ranges became permanent, they were recolonized by C. cham-
aeleonides.

In summary, we suggest that the disjunct distribution of C.
porcus represents an old remnant distribution whose details have
been profoundly affected by the Cenozoic history of Cuba. The
more continuous distribution of C. chamaeleonides on the other
hand would seem to be more recent—a Miocene or later dispersal
from one or more refugia in eastern Cuba. This interpretation
requires that C. chamaeleonides is the more vagile and that it has

GARRIDO AND SCHWARTZ: Cuban Lizards 219

in a relatively short time re-expanded its range from the western
to the eastern portion of the island. C. porcus, on the other
hand, remains restricted primarily to its old range in the moun-
tainous section of eastern Oriente, with two relict outlying popu-
lations in higher mountain ranges in central and western Cuba.

RESUMEN

Se creia que el endémico género cubano Chamaeleolis (Iguanidae) con-
tenia una sola especie, la C. chamaeleonides Cocteau. Pero en realidad existen
dos especies, la segunda siendo C. porcus Cope. Las diferencias entre las dos
especies son: 1) escamas gulares paramediales pequenas, cortadas, cénicas y
piramidales en la chamaeleonides; largas y flexibles en la porcus; 2) escamas
supralabiales separadas del anillo de escamas circumorbitales por una hilera de
escamas en la chamaeleonides, aquéllas en contacto en la porcus; 3) en la
chamaeleonides las escamas dorsales son subcirculares con muchas escamas
interstitiales, en la porcus las cuatro filas de escamas mas dorsales son mas
largas que anchas y con pocas o ninguna escama interstitial, y el resto de las
escamas dorsales carecen de escamas interstitiales; 4) en la chamaeleonides
la distancia del hocico al borde anterior de la érbita corresponde a un numero
entre 42 y 83 de escamas ventrales, siendo estas pequenas o diminutas; en la
porcus la mencionada distancia contiene de 22 a 48 escamas, lo que indica
que las escamas ventrales son mas grandes; 5) las escamas debajo del Angulo
de la boca son muy Aasperas en la chamaeleonides, poco asperas o lisas en la
porcus.

C. chamaeleonides se encuentra por casi toda la isla con excepcidn del
este de la provincia de Oriente (al este de la linea Gibara-Pico Turquino).
C. porcus reside principalmente al este de la linea Gibara-Pico Turquino, pero
se la encuentra tambien en la Sierra de Trinidad en Las Villas (dos ejem-
plares ) y en la Sierra del Rosario en Pinar del Rio (un ejemplar ).

La historia de la geologia cubana explica dichas distribuciones. Se propone
que la porcus fue la especie endémica de las sierras orientales y la chamaeleon-
ides la de las sierras pinarenses. La presencia de la porcus en dos zonas lejos de
la zona principal de la especie en Oriente sugiere que antiguamente ambos
especies de distribuyeran por todas partes de las isla. Esta antiguas extensiones
geograficas se modificaron profundamente en el periodo cenozdico cuando Cuba
estaba submergida y las poblaciones de las dos especies fueron extirpadas en
los llanos, dejando poblaciones relictas de la porcus en las sierras mas occi-
dentales. Cuando Cuba obtuvo su configuracién actual, C. chamaeleonides
volvia a extender su distribucién por toda la isla mientras que C. porcus esta
restringida a los residuos de una distribucidn anteriormente mas amplia.

LITERATURE CITED

Barsour, THOMAS, AND CHARLES T. RAMSDEN. 1919. The herpetology of
Cuba. Mem. Mus. Comp. Zool., vol. 24, no. 2, pp. 71-213.

CocrEau, J. T. (AND G. Brsron, in part). 1838 or 1839. Reptiles y peces
in de la Sagra, Historia fisica, politica y natural de la isla de Cuba.
Paris, vol. 4, pp. 1-255.

Corr, E. D. 1864. Contributions to the herpetology of tropical America.
Proc. Acad. Nat. Sci. Philadelphia, pp. 166-181.

220 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

DuMERIL, A. M. C., AND G. Birsron. 1837. Erpétologie générale. Paris, vol.
AY joyon lel

Maerz, A., AND REA Pau. 1950. A dictionary of color. New York, McGraw-
Hill Book Co., pp. v-vii, 1-23, 137-208, 56 pls.

RoMER, ALFRED S. 1956. Osteology of reptiles. Chicago, Univ. Chicago
Press, pp. viii-xxi, 1-772, 248 figs.

RurpaL, Ropotro. 1964. An annotated checklist and key to the anoline
lizards of Cuba. Bull. Mus. Comp. Zool., vol. 130, no. 8, pp. 475-520,
18 figs.

ScHwARTZ, ALBERT. 1964. Anolis equestris in Oriente Province, Cuba. Bull.
Mus. Comp. Zool., vol. 131, no. 12, pp. 405-428, 7 figs.

SmirH, Hosparr M., AND CHAPMAN GRANT. 1958. The proper names for
some Cuban snakes: an analysis of dates of publication of Ramon de
la Sagra’s Historia Natural de Cuba, and of Fitzinger’s Systema Rep-
tilium. Herpetologica, vol. 14, no. 4, pp. 215-222.

STEJNEGER, LEONHARD. 1917. Cuban amphibians and reptiles collected for
the United States National Museum from 1899 to 1902. Proc. U. S.
Natl. Mus., vol. 53, pp. 259-291, 128 figs.

Wison, Epwarp O. 1957. Behavior of the Cuban lizard Chamaeleolis
chamaeleontides (Duméril and Bibron) in captivity. Copeia, no. 2,
DIAS

Instituto de Biologia, Academia de Ciencias, Capitolio Nacional,
La Habana, Cuba; Miami-Dade Junior College, Miami, Florida,
SOLOTOL SAY

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968 )

Natural Factors Affecting Deer Movement

RicHARD F’, HARLOW AND WILLIAM F. OLIVER, JR.

Tyson (1952) was the first to correlate the number of deer in
a given area with number of tracks. The validity of this census
method is dependent on sampling intensity and knowledge of deer
movement. Considerable variability per count has been reported
(Tyson, 1959; Downing, Moore, and Kight, 1965) but the reasons
for this variability, outside of sampling error, have not been investi-

gated.

In this study, data were gathered and analyzed to evaluate the
possible effect of vegetation types, moon phases, and weather on
deer movement. The track counts were made on eight randomly
selected plots, each approximately 1 mile long, along graded roads
within Crandall Pasture, an area of approximately 15,000 acres
maintained near Fernandina Beach, Florida, by the Southeast
Timber Division of Rayonier Incorporated. Counts were made in
August of each year from 1962 through 1966. Four counts were
made in 1963, and three counts were made in each of the other
years.

The eight track-count plots were randomly selected and _ per-
manently marked. In the afternoon preceding the day of the
count, a drag was used to obliterate all existing tracks. The track
counts were then conducted the next morning between 6 and 8
AM, and only those tracks that actually crossed the 1-mile plot
roads were tallied. Local weather data were obtained from Ime-
son Airport, which is 20 airline miles from the census area, for
each day that tracks were counted. Moon phases were extracted
from the World Almanac and Book of Facts. Percentages of the
types of vegetation were recorded for each mile of road.

DISCUSSION

Deer crossings in relation. to date of census, approximate mile-
long plots, and moon phases were subjected to an analysis of vari-
ance. Results of this analysis are presented in Table 1.

The total number of deer crossings counted along the eight
l-mile plots did not differ significantly between moon phases nor
between years during the 5-year census period. The most signi-

222 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 1
Analysis of variance of deer crossings

Source df SS MS F

Moon phase 3 322 107.33 0.49
Years 4 220 55.00 0.67
Plots 7 24307 3472.43 26nliGea
Moon phase x years 12 2645 220.42 2..69**
Moon phase x plots Hil 1876 89.33 1.09
Years x plots 28 3717 ISPS 162s
Moon phase x years x plots 44 3610 82.04
Total 119 36697

Note: df, degrees freedom; SS, sum of squares; MS mean square; F, ratio of
the mean squares (calculated value).

* Significant :

**Highly significant

ficant factor noted in the analysis was that track counts were con-
sistently high in some plots and low in others over the entire 5
census years. This tendency suggests that deer may have a rela-
tively consistent nonrandom distribution or movement which may
be due to some unmeasured habitat characteristic. The tendency
for track counts to remain stable within a plot (Table 2) suggests
that the track count method might give a good indication of vari-
ation in deer populations, but the high variability in day-to-day
movement of deer necessitates that biologists considering this cen-
sus method recognize its limitations as a means of estimating deer
numbers. Downing et al. (1965) found that 10 or more counts
involving some 30 deer crossings per count would be required to
detect a 20 per cent difference in deer movement (and probably
deer populations ) 95 per cent of the time. It is therefore advisable
to make several counts of a large number of plots to minimize this
variability.

The effect of moon phase-year interaction on deer crossings
was highly significant. The reason for this significant interaction
cannot be explained biologically, but it appears to be due to the
1963 effect of moon phase, as suggested by the single year analyses
shown in Table 3.

A significant interaction occurred between years by plots. Al-
though some plots had consistently more tracks than others, dif-
ferences were noted from year to year. There is no biological

HARLOW AND OLIVER: Deer Movements 225

TABLE 2
Deer track census data arranged chronologically by year and moon phase
PLOTS
Date and

Moon Phase il 2, 3 4 5 6 HM 8
29 Aug. 62 NM 4 8 10 36 16 43 4 6
*Histimated FQ 2 1} 12 34 Lg 38 8 6
15 Aug. 62 FM 0 9 14 44 28 48 5 0
22 Aug. 62 LQ 2 if let om 23 AQ, 5 4
21 Aug. 63 NM 0 23 8 29 5 20 5 12
28 Aug. 63 FQ 2 ol 18 30 21 49 {i 16
4 Sept. 63 FM 8 12 17 57 21 (3 Zi 6
14 Aug. 63 LQ 4 5 3 18 3 23 8 5
“Estimated NM eae | Ce ie Seley Mn ee
12 Aug. 64 FQ il 3 5 36 C 20 eg 3
19 Aug. 64 FM 1 9 4 28 9 30 18 3
26 Aug. 64 LQ 0 13 7 48 i, 51 13 6
27 Aug. 65 NM 2 18 14 74 9 25 12 9
*Estimated FQ 3 13 11 43 10 26 1M if
13 Aug. 65 FM 9 5 5 31 5 20 15 3
20 Aug. 65 LO 3 Seo G3) OL” 20" It 10
16 Aug. 66 NM 5 11 1 DAT 7 19 38 5
23 Aug. 66 FO A Oak aa Be 6 6
30 Aug. 66 FM 1 li IL 38 3 65 oD) 5
*Fistimated LQ 2 Wal 11 35 10 35 18 6

*Estimated—An estimation of missing plot data derived from previously col-
lected data.

explanation apparent, although it is possible that the population
may have undergone some shift.

The effect of moon phase-plot interaction was not significant.

Table 4 lists the percentages of the five types of vegetative as-
sociations, including open water, for each mile of road census, as
well as the total number of tracks counted for each mile during
the 5-year study period. The primary purpose of this table is to
show the habitat configuration of the area under investigation. The
data were insufficient for computing valid statistical relationships.

The data in Table 5 indicate that the uniform weather condi-
tions which prevailed during the track count periods did not have
a noticeable effect on deer movement as evidenced by the ex-

22.4 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 3

Analyses of variance of deer crossings for years 1962 through 1966

Source

1962
Plots
Moon phases

Error

Total

1963
Plots
Moon phases

Error

Total

1964

Plots
Moon phases
Error

Total

1965
Plots
Moon phases

Error

Total

1966
Plots
Moon phases

Error

Total

df

14

23

14

23

SS

6022.50
28.09
133.25

6183.84

ATA 2
1633.97
2102.28

8447.97
4080.63
191.50

590.50

4862.63

5941.63
313.00
935.00

7189.63

2969.83
457.75
1656.42

5084.00

MS

860.36
14.04
9.52

673.10
544.65
100.11

583.00
95.75
42.18

848.80
156.50
66.78

424.26
228.87
118.31

Guim
5.44**

tO 2as
2.2°7

PATOL
2.34

BuO
1.93

Note: df, degrees freedom; SS, sum of squares; MS, mean square; F, ratio of
the mean squares (calculated value).

**Highly significant

HARLOW AND OLIVER: Deer Movements

225

Types of vegetation adjacent to census strips and deer tracks counted per mile

TABLE 4

Vegetation

types IL 2 3 4
Flat-
woods, % ol 39 52 14
Pine-oak
uplands, % 15 AQ 16 NS
Swamps, % 33 I 29 2,
Hammocks, % 0 0 Di 2h
Salt water
marsh, % 0 0 2 iB)
Open
water, % 0 0 0 11
Total
tracks, 38 185 156 565
number

PLOTS

5

6 u
50 25
9 0
40 19
1 31

0 24

0 1
502 198

Note: For brevity percentage of vegetation has been rounded off.

TABLE 5
Climatic conditions and number of deer crossings tallied during the 5-year
census period

Average Amount of Average

Date Temp. (°F) wind
speed.

Avg. Min. (mph )
15 Aug. 62 84 74 5.8
29, Aug. 62 85 74 4.1
29 Aug. 62 84 72 4.9
14 Aug. 63 83 72 6.7
21 Aug. 63 79 70 5.9
28 Aug. 63 81 74 6.1
12 Aug. 64 84 76 1235
19 Aug. 64 78 72 6.2
26 Aug. 64 81 13 7.9
13 Aug. 65 84 Me 9.1
20 Aug. 65 84 74 71
27 Aug. 65 84 73 8.5
16 Aug. 66 81 72 6.9
23 Aug. 66 86 74 7.0
30 Aug. 66 81 2 10.2
T— 0:05 —0.03 —0.02

r =—correlation coefficient

possible
sunshine

87
35
95
50
AT

relative

93

Total
number
tracks

humidity over the
(Per cent) (Per cent)

72
68
66
76
80
81
78
86
74

miles

148
135
127

69
102
174

92
102
145

93
137
163
123
112
191

226 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

tremely weak relationships. Downing et al. (1965) also found no
correlation between weather and tracks within an enclosure in
Georgia.

CONCLUSIONS

This study produced evidence that white-tailed deer of the
Coastal Plain are nonrandomly distributed but have a consistent
pattern of movement. The uniform weather conditions which pre-
vailed during the track count apparently had no effect on deer
movement. Deer activity apparently was not associated with any
particular moon phase.

LITERATURE CITED

Downline, R. L., W. H. Moore, AND JoE KicHt. 1965. Comparison of deer
census techniques applied to a known population in a Georgia enclosure.
Paper presented 19th Ann. Conf. Southeast. Assoc. Game and Fish
Comm., Tulsa, Oklahoma, 13 pp. (mimeo).

Tyson, E. L. 1952. Estimating deer populations from tracks. A preliminary
report. Presented at 6th Ann. Conf. Southeast. Assoc. of Game and
Fish Comm., Savannah, Georgia, 15 pp. (mimeo).

——. 1959. A deer drive vs. track census. Trans. Twenty-fourth N.
Amer. Wildlife Conf., pp. 457-464.

Southeastern Forest Experiment Station, U. S. Department of
Agriculture, P. O. Box 2570, Asheville, North Carolina 28802.

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968 )

Round-Tailed Muskrat in West-Central Florida

Joun R. Pau

DEspPITE several intensive recent investigations (Hall and Cock-
rum, 1953; Schwartz, 1953; Burt, 1954 and Birkenholz, 1963) into
the life history and distribution of the round-tailed muskrat, Neo-
fiber alleni, records of the species from west-central Florida are
lacking. Schwartz (1953) indicated that the area from Taylor and
Suwannee counties, Florida, south to Hillsborough County did not
appear to be presently occupied by round-tailed muskrats although
Simpson (1929) recovered Pleistocene remains of N. alleni from
localities within this area. Schwartz (1953) concludes that the
present range of Neofiber (peninsular Florida and extreme south-
east Georgia, except for the area mentioned above) appears to be
a remnant of one previously more extensive. This paper reports on
records of Neofiber from two new localities and serves to fill some
of the distribution gaps.

One record of Neofiber was provided by C. E. Winegarner,
who on 29 January 1967, obtained the remains of a diamond-back
rattlesnake, within which was found a virtually intact Neofiber
skull. The snake carcass was decomposed with only skin and
skeletal elements remaining and lying in a field one mile south of
Lutz, Hillsborough Co., Florida. This locality is approximately
150 yards from a pond which could have supported round-tailed
muskrats.

A population of N. alleni persists in an isolated marsh in Hills-
borough River State Park (HRSP), an area approximately 15 miles
north of Tampa, near the border of Hillsborough and Pasco coun-
ties. Nine specimens of Neofiber were taken in steel traps from
this pond in June and July 1966; included were four adult males,
three adult females, and two juvenile females. All specimens were
deposited in the collections of the Zoology Department, University
of South Florida, Tampa.

This marsh is approximately three acres in size with the dom-
inant vegetation consisting of a mixture of pickerelweed (Ponte-
deria lanceolata) and maidencane (Panicum hemitomon). The
pond is dependent on rainfall for its water supply and fluctuates
in depth seasonally. In June and July of 1966 the water was 18-30

228 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

inches in depth. Normal winter and spring water levels are 4-12
inches although during the spring months of 1967 standing water
occurred only in several alligator holes. Even at the lowest water
levels alligators continued to reside in this marsh and may be
potential predators on the rats. During the driest periods the rats
burrowed into the banks of several small islands in the marsh and
no evidence of grass houses or feeding platforms could be found.

The population existing in the marsh appears to be relatively
small, probably less than 25 individuals and considerably fewer dur-
ing the dry months. Reproductive activity is apparently coordinated
with the onset of the rainfall in the early summer. All specimens
collected prior to July 20 were adults. Two young rats taken on
20 and 22 July 1966, weighed 118 and 133 grams. Growth data
of Birkenholz (1963) for known-age animals would indicate the
juveniles in the present study were five to six weeks old. This
would place the date of parturition around 15 June. None of the
adult females taken were pregnant or lactating.

The population of Neofiber in HRSP is ecologically isolated;
the marsh is totally surrounded by heavily wooded areas and is
located 2-3 miles from the nearest suitable habitat. Superficial ex-
amination of the ponds in the surrounding areas has shown no
evidence of Neofiber. The cranial measurements (Table 1) of the

TABLE 1
Body and cranial measurements of adult Neofiber alleni

Present Study Schwartz (1953)
mean alleni nigrescens
(range ) mean mean
Specimens measured ih 25 14
Condylobasal Length 43.6 (41.9—44.9) 46.2 46.8
Zygomatic Breadth 27.5 (26.4—28.5) 27.9 XS) IL
Nasal Length 12.8 (12.4—13.0) 13.6 13.8
Breadth of Nasals 5.0 ( 4.6— 5.3) Do 5.4
Upper Molar Row 11.4 (11.0—12.0) ILs 12.0
Interorbital Constriction 5.1 ( 5.0— 5.2) Sal 5.1
Length of Palatine Slits 8.1 ( 7.7— 8.5) Us) 8.1
Body Length O20 (lia2=—213)) 203.0 203.0

Hind Foot 41.0 (40—44) 43.1 43.5

PauL: Round-tailed Muskrat 229

HRSP specimens are smaller in most dimensions than those given
by Schwartz (1953) for Neofiber from adjacent areas. Schwartz
(1953) labeled the populations immediately east of this area as
intergrades between N. a. alleni and N. a. nigrescens. The dorsal
pelage color of the HRSP specimens is somewhat variable, but
tends toward a dark Fuscous Black (Ridgway, 1912). This color,
along with the smaller size, indicates a probable closer relationship
of the HRSP specimens with the more southern populations of
N. a. nigrescens.

LITERATURE CITED

BirRKENHOLZ, D. C. 1963. A study of the life history and ecology of the round-
tailed muskrat (Neofiber alleni True) in north-central Florida. Ecol.
Mon., vol. 33, pp. 187-213.

Burt, W. H. 1954. The subspecies category in mammals. Syst. Zool., vol. 3,
pp. 99-104.

Hai, E. R., ANp E. L. Cockrum. 1953. A synopsis of the North American
microtine rodents. Univ. Kansas Publ., Mus. Nat. Hist., vol. 5, pp. 373-
A98.

Ripeway, R. 1912. Color standards and color nomenclature. Washington,
D. C., published by the author.

ScHwartz, A. 1953. A systematic study of the water rat (Neofiber alleni).
Occ. Pap. Mus. Zool. Univ. Michigan, no. 547, 27 pp.

Smpson, G. G. 1929. The extinct land mammals of Florida. 20th Ann.
Rept. Florida State Geol. Surv., pp. 229-80.
Illinois State Museum, Springfield, Illinois 62706

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968 )

Mating Behavior of Peromyscus polionotus

MicHAEL H. SmiITH

STUDIES on interspecific behavioral discrimination have involved
almost all of the species of Peromyscus in the United States (Blair
and Howard, 1944; Clark, 1952; Blair, 1953 and 1954; Harris, 1954;
Bradshaw, 1965; Tamsitt, 1961b; McCarley, 1964; Moore, 1965;
Smith, 1965 and 1966). Unfortunately a comprehensive knowledge
of this multidimensional phenomenon does not necessarily result
from the inclusion of many species. Most of the existing data con-
cern only one aspect of behavioral discrimination, the tendency to
form homo- or heterospecific social aggregates under crowded lab-
oratory conditions. The interpretation of these data usually depends
upon the assumption that there is a direct relationship between the
tendencies to associate and to breed with another animal. Differ-
ences in the sequence of behavior culminating in mating could
prove this assumption to be in error.

Tamsitt (196la) described mating behavior in detail for P. com-
manche, P. nasutus, and P. truei of the P. truei species group. The
purpose of this paper is to present similar data for P. polionotus
of the P. maniculatus species group and to compare these results
with those of Tamsitt.

METHODS

During 1962 and 1963, 64 pairs of mice captured in the field
were used in the laboratory for breeding stock. Occasionally, one
escaped or died and had to be replaced; additional pairs were added
later to the colony. Four Florida subspecies were represented: P.
polionotus subgriseus from Ocala National Forest, P. p. phasma
from Anastasia Island, Saint Johns County, P. p. leucocephalus
from Santa Rosa Island, Okaloosa County, and P. p. rhoadsi from
Archbold’s Biological Station, Highlands County.

The mice were kept in cages similar to those described by
Layne (1958); the sides were made of one-quarter inch hardware
cloth and the wood tops were removable. Each cage was placed
on a tray covered with sawdust. The ambient temperature in the
laboratory was usually 24 + 2° C; occasionally it varied as much
as 5°C. Relative humidity was measured with a sling psychrometer

SMiTH: Mating of Beach Mouse 23k

at different times during the day and night; it varied from 54 to 84
per cent. Overhead fluorescent lights were automatically turned on
at 0630 and off at 2030; no outside light entered the room. Purina
laboratory chow and water were supplied ad lib.

Mating was regularly accompanied by sound and occurred in
the evening while the lights were still on in the colony room. De-
tailed notes were made on 16 different pairs of P. p. subgriseus
for a total of 25 matings. Additional scattered observations were
made on the other three subspecies. One pair was observed at a
time. All of the matings occurred during the post-partum heat.
The cages were checked in the morning for new-born litters, and
if present, observations were begun that night 30 minutes before
the overhead lights went off. A reflector lamp with a 100-watt red
light located eight to ten feet from the cage was used after the
room became dark.

Sometimes the mice appeared to have no interest in breeding.
In a majority of these cases, however, copulation occurred within
a few minutes after the mice were given a ball of cotton about 3
cm in diameter. Thirteen of the 25 matings occurred with the
previous litter present and 12 without it. Four additional matings
were observed in a wooden cage (50 cm high, 45 cm wide, 75 cm
long) with a glass front and one-quarter inch hardware cloth on the
back. The large cage was used because Tamsitt (196la) suspected
that certain aspects of the breeding behavior might have been
caused by the small size of his cages.

RESULTS

Mating of the old-field mouse varied from pair to pair and with
the number of copulations that took place on the same evening
(Fig. 1; Table 1). The sequence usually associated with the first
few copulations will be presented first and the variation from this
sequence will follow.

The female initiated mating by positioning herself in front of
the male. Her ears were erect, the tail was low, and the body was
held high. The white underparts of the side of her body next to
the male were displayed, and the eye on the same side was slightly
closed. She moved forward displaying one side of her body, and
just before making contact, lowered her body and turned her head
upward and to the side, thus exposing her white throat. The male

232, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

MALE FEMALE

rn ef TAKES INITIATIVE |
4

WASHES FEMALE

erent

RUNS IN FRONT OF MALE

Sy ee “—““SF UPRIGHT POSITION fers
connec --f ESCAPE FROM FEMALE +; i SSE STOPS RUNNING
[DiScONTINUES MATING] : :
[MOUNTS FEMALE i concmecmecmccncce? DEFENSE POSTURE Tee und

[_ WASHES ates ABDOMEN 1 2 ||

Jones bi vseasss nes sennuenes nares’

[FORWARD THRUST & INTROMISSION 1 ARCHES BACK

FI OFSUA UN'S OPES areteeeteeeeereee SAE, FORWARD MOVEMENT [o—=——~
i

[LICKS GENITALIA LICKS GENITALIA

:
A E
i : i

j i

Bae oes aaTiAL WASH 3:

ae AG COMPLETE WASH

Fi

>
i i
F i

i
[___INGESTION

| grin —)

Traeseeenevsessanersseeevons:

: 4
eg EGEST 1ON

ansenneans ones =

i
RESTING

Fig. 1. Sequence of behavior patterns during mating. Solid lines indicate
the usual sequence of behavior leading up to and following the first few copu-
lations. Broken lines indicate alternate sequences used at various times during
mating.

had his body relatively high, ears erect, and tail low. The male
also closed his eye on the side of his body closest to the partner
just before contact was made.

Mutual grooming and then smelling of the genital region oc-
cured. The partners occasionally walked in a tight circle in naso-
anal contact. Grooming was usually confined to the sides and back
of the body of the other animal; occasionally the neck and the
sides and dorsal part of the head were included. After the female
stopped smelling the male’s genital region, she started running in a

SMITH: Mating of Beach Mouse 233

tight circle in front of the male with her head, tail, and body close
to the floor. In the large cage, the female ran back and forth on
the floor in front of the male. If the male did not chase the female,
she repositioned herself in front of him and repeated the entire
sequence.

The naso-anal contact became so vigorous in some cases that
the female was pushed forward. In other instances, the male re-
peatedly moved its head upward after placing it under the female’s
anal region. This movement elevated the posterior half of the fe-
male’s body 3 to 5 cm exposing her white belly. The male fre-
quently tried to mount the female if she slowed down slightly.
Successful mounting occurred only when the female came to a
complete stop. At this moment she extended her body forward,
angled the tail laterally, arched her back, and thus raised the ex-
posed perineum. The male mounted by extending his body over
hers and then grasped her body in the region of the diaphram
with his front legs. Intromission consisted of a single forward
thrust of such force that the female was normally pushed forward.
The female was never heard squeaking during intromission. After
the forward thrust, the male relaxed its grip of the female. The
male always lifted one leg as he drove the penis forward. Occa-
sionally, the other leg was also lifted at the end of the forward
thrust, and thus, the male rode on the back of the female. In these
cases, the male slowly fell off the female onto his side or back. If
one foot remained on the floor, the male dismounted as the female
moved forward. The male then sat upright on the hind legs and
tail, bent its head toward the floor, pulled sheath of the penis back
with the forepaws, and took the withdrawn penis into the mouth
and washed it. At the same time, the female licked her genitalia
from a similar position. Both sexes went through a complete
“wash”, which includes the entire body and tail, following the
cleaning of the genital area. The “wash” of the female was of
shorter duration than that of the male. She went back to the nest,
which contained the newborn young, and then ate or drank before
initiating the next mating. The male was frequently still cleaning
his fur when the female repositioned herself in front of him. Mice
always copulated many times during the course of an evening
(Table 1). No conspicuous marking behavior, such as urination or
dragging the genital region on the floor, was observed.

234 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

TABLE 1

Mating data for laboratory-reared Peromyscus polionotus subgriseus

Number of pairs observed 16
Successful matings 25
Mean number of mounts per mating 49.8
Mean number of thrusts per mating 45.6
Intromissions per total mounts 91.6%
Mean number of washing of penis by male per mating UProll
Mean observation time in minutes per mating 152.6

Copulations were not randomly distributed over time but oc-
cured in groups (or series) with a greater amount of time between
groups than between successive copulations. Between each series
of copulations the animals rested and showed no sexual behavior
for at least three minutes. The male frequently lay facing the nest
from the farthest possible point in the cage, keeping his body and
tail flat on the floor, ears erect, and limbs extended in a position
similar to that used by heat-stressed animals. The female usually
stayed in the nest with the young mice. As the length of the period
of mating increased, both mice spent more time drinking, eating,
urinating, and defecating during the rest periods. The number of
series of copulations per mating period ranged from six to twelve
and averaged 9.2. The mean number of copulations per series
decreased and the length of the rest period between each series
increased as the number of copulations completed that night in-
creased. The mean number of copulations for the first series was
6.8 and for the last series was 2.6. The average length of the rest
period between the first and second series was 3.8 minutes, and it
was 15.3 minutes before the last series.

Mating behavior on any one evening gradually changed as the
total number of completed copulations increased. The male re-
sponded to changes in the female’s behavior more quickly than
before. Grooming and smelling of the genital region were frequent-
ly deleted. The male often started chasing the female without be-
ing approached by her. The washing of the penis occurred less
frequently, and the partial wash of the head and ears were more
common than the complete performance. The female cleaned her
genitalia less frequently.

SmiTH: Mating of Beach Mouse 235

In almost all of the later matings the male took the initiative.
He would approach the female head on while she was in the nest.
The female would frequently assume an upright posture, and the
male would attempt to push the female over on her back and wash
her white belly. The female uttered frequent, high-pitched sounds
during the initial advances of the male and occasionally while she
was on her back. When the female righted herself, she would run
out of the nest with the male chasing after her and the normal
sequence of events occurred.

Sometimes, when the female resisted the male’s efforts, the
male picked up a new-born animal and carried it out of the nest.
The female immediately retrieved the young animal and after de-
positing it in the nest, the female would run from the nest with
the male following. The male frequently attempted to mount the
female while she was retrieving the young animal but was never
successful. Half of the males picked up young animals and moved
them out of the nest during their mating behavior. The other males
concentrated their efforts on the females. New-born animals were
displaced several times from their nest by one male the first time
it was observed and not at all during the second period of observa-
tion. The general scheme of this male’s mating behavior was sim-
ilar on both occasions except in this one regard.

Females gradually became more aggressive and drove their
mate and previous litter from the nest when they were still present.
The female frequently lunged at the male and tried to bite him.
Eventually, the male left the female alone in the nest and mating
ceased. In some cases, the female left her previous litter in the
nest although she drove her mate out. The period during which
the previous litter and/or the adult male were kept out of the nest
varied from several hours to several days.

The young animals from the previous litter were allowed only
restricted movements during the mating of their parents. Initially,
they spent most of their time in the nest but were later found eat-
ing and drinking near the water bottle. As the female approached,
they would run back to the nest. Except in one case, the young
animals played no direct part in the mating behavior. One juvenile
male entered into the chasing of the female. The juvenile male
never attempted to mount his mother or make any kind of physical
contact with her. He ran behind his father and frequently tried

236 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

to mount him. The adult male payed little attention to the activity
of the juvenile, but the female frequently lunged at him and
eventually chased him back into the nest with the other young
mice.

On four occasions, mating was observed when there was more
than one adult male in the cage. Usually only one male bred with
the female. The others either remained in one corner of. the cage
or they attempted to mount the male that chased the female. Once
two males alternately tried to mount a female, but one repeatedly
failed because the female never stopped and arched her back when
he tried to mount.

The only consistent difference observed in the mating behavior
of the several subspecies was the lack of squeaking by females of
P. p. phasma.

Discussion

As pointed out by Smith (1966), the normal social unit in this
species is an adult male and female with or without young. Fe-
males are normally dominant over their mates and play a major
role in the process of pair formation and maintenance of the pair
bond. No other species of Peromyscus is known to be as social as
P. polionotus. A proper interpretation of the behavior of the old-
field mouse and its comparison with that of other species in this
genus can only be made when these unique characteristics of the
species are kept in mind.

Mating was the result of a series of social interactions in which
the female at first played a submissive role and then later a domi-
nant one. The change was gradual and was accompanied by an
increase in the amount of initiative shown by the male. Squeaking
by the female only occurred in ambivalent situations characterized
by both avoidance and approach tendencies. Once the female
started to run in front of the male or attacked him, she would stop
squeaking. Display of the white underparts of the female’s body
was also a part of her submissive behavior. Reversal of the normal
social hierarchy seemed to be a necessary prerequisite for the
completion of mating. The re-establishment of the female’s domi-
nance over the male was correlated with the cessation of mating.

The six phases of the mating pattern described by Tamsitt
(196la) for the P. truei species group were also evident in the
behavior of P. polionotus. They were “(1) initiation of courtship

SmitTH: Mating of Beach Mouse 237

by the female, (2) circling of the female before the male, (3) pos-
turing by the female before the male, (4) mounting by the males,
(5) thrust-intromission by the male, and (6) dismounting by the
male.” Of these arbitrary divisions only circling in front of the
male by the female was considered to be a laboratory artifact
caused by the small cage size. The essential part of the circling
was probably the movement away from the male, or fleeing; this
being interpreted as submissive behavior.

The overall similarity of the mating patterns was probably due
to the close phylogenetic relationship of the mice. Both of the
species groups belonged to the subgenus Peromyscus. Despite the
similarity, a detailed comparison of the results also revealed some
consistent differences.

Only the females squeaked, but they did so at different times.
Those in the P. truei group usually squeaked once during intro-
mission; P. polionotus females were quiet at this time but vocalized
during their interactions with the males prior to mounting. How-
ever, this behavior was not an essential part of mating since the
P. p phasma females always remained silent and the females of the
P. truei group occasionally did likewise. These sounds may be
used for species recognition in P. polionotus but were more likely
a sign of submission by the female. The silence of the P. p. phasma
females may be related to the open habitat on the beach dunes
where they are found, but P. p. leucocephalus females vocalized
occasionally, although not as frequently as some of the other sub-
species. The latter subspecies was also collected on the sand dunes.
Silence may be selectively advantageous in open habitats, because
certain predators can use sounds to locate their prey.

Males of all of the subspecies of P. polionotus were observed
carrying newborn young out of the nest. This behavior was not
mentioned for P. truei by Tamsitt, but he did not observe mating
during the post-partum heat so there were no young animals
present. Handling of the young by the male, however, was not an
essential part of mating. Some of the males never exhibited this
behavior, while others did so only during one of the several ob-
served mating periods.

The nest acts as a focal point around which the mice center
their mating behavior. This may be due to a conflict between mat-
ing behavior and maternal behavior in the female, or it indicates

238 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

that mating normally takes place within the burrow in the field. A
third alternative is that it is related to the small size of the cages.
The last does not seem likely since the mice used their nest in the
big cage just as often as in the small ones. By mating underground
the mice would not be as susceptible to predation. The old-field
mice would be perfectly safe if they left the entrance to their bur-
row sealed as they do during the day (Smith, 1966).

The last and perhaps most important difference is the lack of
conspicuous marking behavior by the P. polionotus females. In the
P. truei group females dragged the perineal region on the floor
propelling themselves by movements of the forelimbs. Nothing
even remotely similar to this was observed in P. polionotus. Tam-
sitt interpreted the females behavior as functioning in depositing
olfactory stimuli for the male. After being stimulated the male
would make contact with the female and a series of epimeletic
activities would ensue, their function being to condition the mice
to each other's presence. This type of conditioning was not nec-
essary in P. polionotus because of the prior formation of the pair
bond (Smith, 1966). The various characteristics of the mice that
contribute to the formation of the pair bond in P. polionotus were
probably more important than the minor variations encountered in
their mating behavior as compared with that of the P. truei group.
Of course these same characteristics can be used for discrimina-
tion during mating, but if they were they did not seem to be em-
phasized to the same degree in the different species of mice.
Recognition was already established before mating started in P.
polionotus.

SUMMARY

Mating is the result of a series of social interactions in which the
female at first plays a submissive role and then later a dominant one.
Six phases are recognized in the mating behavior of P. polionotus:
(1) initiation of courtship by the female, (2) running away by the
female, (3) posturing by the female before the male, (4) mounting
by the male, (5) thrust-intromission by the male, and (6) dismount-
ing by the male. These are essentially the same as those described
by Tamsitt (1961la) for the P. truei group. There are differences in
the mating sequence of the two types of mice, but only one of these
seems to be essential to the completion of mating. This is the mark-
ing behavior of the females of the P. truei group, an action which

SMitH: Mating of Beach Mouse 239

was not observed in P. polionotus. One of the functions of this be-
havior is probably species recognition, and since recognition is
established prior to mating in P. polionotus, marking would be
superfluous in this species.

ACKNOWLEDGMENTS

I would like to thank Dr. E. G. F. Sauer for his patience and
guidance as chairman of my Ph.D. committee. Drs. R. Beyers, D.
Coleman, and F’. Golley critically read the final draft of the paper,
and my wife, Irma Smith, assisted me in various ways. A grant-in-
aid from Sigma Xi and support from the Department of Zoology at
the University of Florida helped finance the study. Preparation of
the manuscript was aided by AEC Grant AT(38-1)-310 and NSF
Grant GB5140. To all of the above I express my sincere apprecia-
tion.

LITERATURE CITED

Buair, W. F. 1953. Experimental evidence of species discrimination in the
sympatric species, Peromyscus truei and P. nasutus. Amer. Natur., vol.
87, pp. 103-105.

1954. Tests for discrimination between four subspecies of deer mice
(Peromyscus maniculatus). Texas Jour. Sci., vol. 6, pp. 201-210.

AND W. E. Howarp. 1944. Experimental evidence of sexual isolation
between three forms of mice of the cenospecies Peromyscus maniculatus.
Contrib. Lab. Vert. Biol. Univ. Michigan, vol. 26, pp. 1-19.

BrapsHaw, W. N. 1965. Species discrimination in the Peromyscus leucopus
group of mice. Texas Jour. Science, vol. 17, pp. 278-293.

Cuark, W. K. 1952. Isolating mechanisms, competition and geographic vari-
ation of the Peromyscus boylei group in Oklahoma and Texas. Unpubl.
Ph.D. dissertation, Univ. Texas, 102 pp.

Harris, VAN T. 1954. Experimental evidence of reproductive isolation be-
tween two subspecies of Peromyscus maniculatus. Contrib. Lab. Vert.
Biol. Univ. Michigan, vol. 70, pp. 1-13.

Layne, J. N. 1958. A simple and inexpensive small animal cage for laboratory
use. Turtox News, vol. 36, pp. 208-209.

McCartey, H. 1964. Ethological isolation in the cenospecies Peromyscus
leucopus. Evolution, vol. 18, pp. 331-332.

Moore, R. E. 1965. Olfactory discrimination as an isolating mechanism be-
tween Peromyscus maniculatus and Peromyscus polionotus. Amer. Mid].
Natur., vol. 73, pp. 85 -100.

240 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES

SmitH, M. H. 1965. Behavioral discrimination shown by allopatric and sym-
patric males of Peromyscus eremicus and Peromyscus californicus be-
tween females of the same two species. Evolution, vol. 19, pp. 430-435.

1966. The evolutionary significance of certain behavioral, physiologi-
cal, and morphological adaptations of the old-field mouse, Peromyscus
polionotus. Unpubl. Ph.D. dissertation, Univ. Florida 187 pp.

TamsitT, J. R. 196la. Mating behavior of the Peromyscus truei species group
of white-footed mice. Amer. Mid]. Natur., vol. 65, pp. 501-507.

1961b. Tests for social discrimination between three species of the
Peromyscus truei species group of white-footed mice. Evolution, vol.

15, pp. 555-563.

Institute of Ecology and Department of Zoology, University of
Georgia (mailing address: Savannah River Ecology Laboratory,
c/o U. S. Atomic Energy Commission, SROO, Box A, Aiken, South
Carolina 29801).

Quart. Jour. Florida Acad. Sci. 30(3) 1967 (1968)

FLORIDA ACADEMY OF SCIENCES

INSTITUTIONAL MEMBERS FOR 1967

Archbold Expeditions

Barry College

Central Florida Junior College
Florida Atlantic University
Florida Institute of Technology
Florida Presbyterian College
Florida Southern College

Florida State University
Jacksonville University
Marymount College

Miami-Dade Junior College
Mound Park Hospital Foundation
Nova University of Advanced Technology
Polk Junior College

Rollins College

St. Leo College

Stetson University

University of Florida

University of Florida
Communications Sciences Laboratory

University of Miami
University of South Florida

University of Tampa

FLORIDA ACADEMY OF SCIENCES
Founded 1936

OFFICERS FOR 1967

President: Jackson P. SICKELS
Department of Physical Sciences, University of Miami
Coral Gables, Florida

President Elect: CLARENCE C CLARK
Department of Physical Sciences, University of South Florida
Tampa, Florida

Secretary: Joun D,. McCrone
Department of Zoology, University of Florida
Gainesville, Florida

Treasurer: JAMES B. FLEEK
Department of Chemistry, Jacksonville University
Jacksonville, Florida

Editor: Pierce BRODKORB
Department of Zoology, University of Florida
Gainesville, Florida

Membership applications, subscriptions, renewals, changes
of address, and orders for back numbers should
be addressed to the Treasurer

Correspondence regarding exchanges
should be addressed to

Gift and Exchange Section, University of Florida Libraries
Gainesville, Florida

Bea aan CC RI Al eed eermeren nti hers aes ai
JLSNI NVINOSHLINS S31YVYUAIT_ LIBRARIES SMITHSONIAN INSTITUTI
= w = ts: 2) = ie
< = = ah <x Ae
ha w : a ow DD XN“
Ye Nie oO 4 Se O a
Lp = = = i=
= > = } > =
wu Ps Y * woe w” ‘
RIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYV
cole > n = aes ” i
a aM a 182) a
ee = oe “ tee
< % <
Gm wud) = us) SS =
=) faye : E =
ol : a)
{LSNI_NVINOSHLINS S3IY¥Vu¥g!I
te : x zZ c
w = we = a
me) = 0 = 4 oe
> = > ra (ics
Ron E a ae =
D z oD Zz ne
RIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IuY
wn ne w = re
= Sie = Wy = =
= z= \. oi Ay, = a
oC) 2 OE FAA EE
~S 2 a Ss \ ay YG fh es 3
Ye ee Ae : :
ILSNI_ NVINOSHLIWS SAIYVeGIT LIBRARIES SMITHSONIAN
= z = = Yiu, %
c < a < Yi fy 7 o
— om = o Of Ue? 7: =
= mM — oO. =
fe) a Oo ase oO
= os Pa ae A
RIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS -
z ats = a pa =
fe) iB Ss) oo. a ee
= 2 5 2 NR 5
Ey = - calle =
- a ~ e =
ao = uy =
ILSNI S SASIYVYE!IT LIBRARIES
Zz ” z a “ =
ies = ps = <
: = ;
lp 3 7 8 QS F a)
Y ce rene: e O r
Wp eS = a z = <N
3 S ae 5 7
aie cae
RIES SMITHSONIAN INSTITUTION NOILALILSN! NVINOSHLINS Sa 1yV ue
ws Z ut a > So wl Kann
oc. ce a ie a
< } 2. < =A < [9
re a = a 5 oc \h
mM = aa — m 7;
uk a oO aE Oo ae ;
ai mz ats Zz FY ess |
ILSNI_NVINOSHLINS S34 iy¥Vugd bday LIBRARI ES_ SMITHSON AN
ee | | ar ee oO aap ~~ a oO Sy, = A

ee Ae - pa ey eee aR OS om
mn 5 a ESO EG
a Naa | z nee in 2
NVINOSHLINS a

SAIUVYGIT LIBRARIES SMITHSONIAN INSTITU

S

RIES SMITHSONIAN INSTITUTION

re '
NVINOSHLIWS
SMITHSONIAN
NVINOSHLIW
SMITHSONIAN

NOILNLILSNI NVINOSHLIWS S3I1uVY

EX
ss

SMITHSONIAN INSTITUT

S

ARIE

LSNI_NVINOSHLINS
ii “

LIBRARIES
LIBRARIES

LIBR

SAlYVUdIT LIBRARIES

S|

pSMITHSONIAN INSTITUTION NOILALILSNI_ NVINOSHLIWS

INSTITUTION NOILNLILSNI

saiuvugiy

INSTITUTION NOILNLILSNI

SSiuYVdd!)
S saidvug

*

‘>

NVINOSHLINS

SNI_NVINOSHLIWS

. “
ory

SAIYVYEIT LIBRARIES SMITHSONIAN INSTITUT

SMITHSONIAN
NVINOSHLIWS
SMITHSONIAN
NVINOSHLIW

MITHSONIAN

= ” = =
aA ow + mn od
pol “4 Re je
ha Se I yp nt
SoS). 2S O si a.
iemeey = —t we
RI ees BO ORIEN INSTITUTION NOLLALILSNI NVINOSHLIWS Saluvy
vo. . Pe es Ss
POM = ©,3) (9) 2 8S |
5 Gy = i : a
2° | Z fs ue
_SN NVINOSHIIWS Saluvudly LIBRARIES) SMITHSONIAN INSTITUT
3
Ba
UY
oO
os
>
=

NVINOSHLIW
SMITHSONIAN
SMITHSONIAN

1ES SMITHSONIAN NOILNALILSNI NVINOSHLINS SS3IYVY!
i US a BP eos yl
al aM u (ep) ah
bivee en ee ui oc
ti a. 4 od <
cm 3S S 5 ae
oe) = —t z Pe ne

Ok) Cae TE TIMI. OOo ADA eed ,ErRmRAMei ewe CRBITLIOCRITARL TRIO Rr

SMITHSONIAN INSTITUTION LIBRARIES

IAAT

9088 01354 1693

Pea ; ty, ; ‘ '
‘ 2 Fi fk aL
; E '
5 F
74 pire * 2 ‘ ‘
<: ; Be: < H Bye ‘
i ‘ : ‘
1 r " ‘ ; :
‘ Art ;
’ 44 ‘ :
4 woaek L P Z ew.
4 Ut oe
he wee ae é
' tee er a sek ak Oe ‘ Hi i
wy i Fi ' "4
be ‘ Foy outs yi $1 1 : wag :
i Yat ORES ie eRe ae Tar aa 4 ¥
aera { : 1
t : ‘ ‘ : t be r 4
i é vias Fi 1 i if :
ea | pe) nip ‘ 3 . oe ‘ ‘
. yo ‘ : t H he