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Group Title: Bulletin of the Florida State Museum, Biological Sciences
Title: A Systematic study of two species complexes of the genus Fundulus (Pisces Cyprinodontidae)
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Permanent Link: http://ufdc.ufl.edu/UF00095803/00001
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Title: A Systematic study of two species complexes of the genus Fundulus (Pisces Cyprinodontidae)
Physical Description: 64 p. : ill. ; 23 cm.
Language: English
Creator: Relyea, Kenneth
Florida State Museum
Donor: unknown ( endowment )
Publisher: Florida State Museum
Place of Publication: Gainesville Fla
Gainesville Fla
Publication Date: 1983
Copyright Date: 1983
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Subject: Fishes -- Classification   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography : p. 46-50.
Bibliography: Bulletin of the Florida State Museum, Biological Sciences ; volume 29, number 1
Statement of Responsibility: Kenneth Relyea.
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Bibliographic ID: UF00095803
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
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Resource Identifier: oclc - 09765579

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Full Text









of the
FLORIDA STATE MUSEUM
Biological Sciences
VOLUME 29 1983 NUMBER 1






A SYSTEMATIC STUDY OF TWO SPECIES COMPLEXES
OF THE GENUS FUNDULUS
(PISCES: CYPRINODONTIDAE)

KENNETH RELYEA


UNIVERSITY OF FLORIDA


GAINESVILLE









Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL
SCIENCES, are published at irregular intervals. Volumes contain about 300 pages and are not
necessarily completed in any one calendar year.













OLIVER L. AUSTIN, JR., Editor
RHODA J. BRYANT, Managing Editor

Consultants for this issue:

GEORGE H. BURGESS
STEVEN P. CHRISTMAN
CARTER R. GILBERT
ROBERT R. MILLER
DONN E. ROSEN













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Copyright @ by the Florida State Museum of the University of Florida

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Publication dates: 22 April 1983


Price: $3.30























TABLE OF CONTENTS


INTRODUCTION ................................................. 3

ACKNOWLEDGEMENTS ............... ..... .. ..... ..... ........................ 3

M ETHODS ................ .......................................... .............. 3

KEY TO SPECIES ...................... ............................................ 4

SYSTEMATIC ACCOUNTS ................. ........................................... 5

Fundulus heteroclitus (Linnaeus) .......... ....... ........................ 5

Fundulus bermudae (Giinther) ................. .........................11

Fundulus rhizophorae (Goode) ...................... ................... 12

Fundulus grandis grandis (Baird and Girard) ..................................15

Fundulus grandis saguanus (Rivas) ..............................................20

Fundulus grandissimus (Hubbs) .............................................23

The Fundulus majalis Species Complex ................ .....................27

Fundulus majalis (Walbaum)-Fundulus similis (Baird and Girard) ..............27

Fundulus persimilis ......................... .............................. 37

COMPARISON OF THE SPECIES COMPLEXES ................. ...........................40

ZOOGEOGRAPHY ..................................................................41

LITERATURE CITED ................... ............................................. 46











A SYSTEMATIC STUDY OF TWO SPECIES COMPLEXES
OF THE GENUS FUNDULUS
(PISCES: CYPRINODONTIDAE)
KENNETH RELYEA1
ABSTRAcr: Two Fundulus species complexes, the Fundulus heteroclitus-F. grandis and F.
majalis species complexes, have nearly identical overall geographic ranges (Canada to north-
eastern Mexico and New England to northeastern Mexico, respectively; both disjunctly in
Yucatan). Fundulus heteroclitus (Canada to northeastern Florida) and F. grandis (northeast-
ern Florida to Mexico) are valid species distinguished most readily from one another by the
total number of mandibular pores (8 and 10, respectively) and the long anal sheath of female
F. heteroclitus. Fundulus maialis and F. similis are conspecific (F. majalis has nomenclatural
priority) and intergrade in northeastern Florida in the same region in which F. heteroclitus and
F. grandis are sympatric. Overlapping populations in other species groups of fishes, notably
Sphoeroides, Chasmodes, and Menidia, occur in the same northeastern Florida area. Both
Fundulus population complexes have disjunct Gulf and Atlantic populations, differentiated
allopatric populations in the Florida Keys (and Cuba for F. grandis), and allopatric related
species in Yucatan (F. grandissimus and F. persimilis). Presumably F. grandis and southerly F.
majalis ("similis-type") evolved in the Gulf of Mexico in isolation from Atlantic coast ancestors
during some Pleistocene interglacial period, and have recontacted Atlantic populations in
northeastern Florida since the Wisconsin glaciation. Yucatan populations of Fundulus, Flori-
dichthys, Menidia, and Cyprinodon and Florida Keys populations of Fundulus, Menidia,
Syngnathus, and possibly Lucania and Cyprinodon are considered to be glacial relicts.

RESUMEN: Dos complejos de species del g6nero Fundulus: Fundulusheteroclitus-F. grandis y
F. majalis tienen una distribuci6n geografica general casi id6ntica (desde Canada al noreste de
Mexico y desde Nueva Inglaterra al noreste de Mexico, respectivamente, siendo ambas
disyuntivas en Yucatan). Fundulus heteroclitus (desde Canada al noreste de Florida) y F.
grandis (desde el noreste de Florida a Mexico) son species vilidas distinguibles facilmente
entire si por el numero total de poros mandibulares (ocho y diez, respectivamente) y la larga
membrana anal en la hembra de la especie F. heteroclitus.
Fundulus majalis y F. similis son coespecificos (F. maialis tiene prioridad en la nomencla-
tura) y se alternan al noreste de Florida, en la misma region en la cual F. heteroclitus y F.
grandis son simpatricas. Poblaciones de otros grupos de peces con distribuci6n superpuesta
como es el caso de Sphoeroides, Chasmodes y Menidia, se presentan en esa misma area del
noreste de Florida.
Ambas poblaciones del complejo Fundulus tienen distribuci6n disyuntiva en el Golfo de
Mexico y en la costa Atlantica, asi como poblaciones alopitricas en los Cayos de Florida (y
Cuba en el caso de F. grandis) y species alopatricas relacionadas en el Yucatan (F. grandissi-
mus y F. persimilis).
Presumiblemente, F. grandis y F. majalis (tipo similis) se originaron en el Golfo de M6xico,
aisladas de sus ancestros de la costa Atlantica durante alguin period interglacial del Pleis-
toceno y posteriormente recontactaron a las poblaciones del noreste de Florida a partir de la
glaciaci6n del Wisconsin. Las poblaciones deFundulus, Floridichthys, Menidia y Cyprinodon
en Yucatin, asi como las poblaciones de Fundulus, Menidia Syngnathus y probablemente
Lucania y Cyprinodon en los Cayos de Florida, son consideradas como relictos vivientes del
period glacial.
'The author is a member of the Department of Zoology, University of Kuwait, Kuwait.

RELYEA, K. 1983. A Systematic Study of Two Species Complexes of the Genus Fundulus
(Pisces: Cyprinodontidae). Bull. Florida State Mus., Biol. Sci. 29(1): 1-64.







RELYEA: GENUS FUNDULUS


INTRODUCTION
The North American killifishes of the genus Fundulus are relatively well
known taxonomically. Two general reviews of the genus have appeared
(Miller 1955, Brown 1957), and Rosen (1973) included it in his general
discussion of the brackish water species of Cyprinodontidae. All studies at
the species level have dealt with freshwater forms (Fundulus rathbuni
Brown 1955; F. catenatus and F. stellifer Thomerson 1969; F. cingulatus
and F. chrysotus Brown 1956a; F. olivaceus and F. notatus Brown 1956b,
Braasch and Smith 1965, Thomerson 1966; F. notti and F. lineolatus Brown
1958, Rivas 1966, Wiley 1977; F. blairae Wiley and Hall 1975; F. waccamen-
sis Hubbs and Raney 1946; F. diaphanus Shapiro 1947). The last species
often enters brackish water. Other freshwater species would be worth-
while candidates for taxonomic analysis.
No systematic study has been concerned with brackish water forms, of
which 10 species are currently recognized in the Atlantic Ocean and Gulf of
Mexico. This study deals with two brackish water species complexes, the
Fundulus heteroclitus-F. grandis and F. majalis-F. similis population com-
plexes, which have remarkably similar overall distributions.
The F. heteroclitus-F. grandis complex ranges continuously (or nearly
so) from Labrador (Kendall 1909) to northeastern Mexico (Miller 1955).
Allopatric populations occur on Bermuda (F. bermudae and F. rhizopho-
rae) and the Yucatan Peninsula of Mexico (F. grandissimus) (Hubbs 1936).
The F. majalis-F. similis complex ranges from New Hampshire (Jackson
1953) to northeastern Mexico (Miller 1955). The similarity of the distribu-
tion of these two population complexes allows comparison of geographic
variation between them, and provides a basis for zoogeographical interpre-
tations.
Although Brown (1957:73) placed members of both species complexes
in the subgenus Fundulus, it is not certain that the two groups are closely
related (Miller 1955:7-9).
ACKNOWLEDGEMENTS
I extend especial gratitude to Dr. Royal D. Suttkus (Tulane University) and Dr. Ralph W.
Yerger (Florida State University) for their patience and guidance. In addition to the above, the
following people provided specimens for examination and/or advice: James E. Bohlke,
Carter R. Gilbert, Robert R. Miller, Edward C. Raney, Luis R. Rivas, C. Richard Robins,
Walter R. Courtenay, Jr., Robert Kanazawa, Martin Moe, Robert Christensen, Richard M.
Blaney, Glenn Clemmer, Alfred Smalley, E. O. Wiley, Harold Dundee, Michael Dahlberg,
Gerald Gunning, and Donn E. Rosen. I am most grateful to these people. Ahmed Khalifa,
Kuwait University, prepared the line drawings.
I am also indebted to my wife Gail, and to former Jacksonville University students Harry
Sahlman, Julian Humphries, Bruce Sutton, and Charles Duggins for assistance in the field.
METHODS
Counts and measurements used follow the methods proposed by Hubbs and Lagler
(1958), with exceptions as noted below. The lateral scale count was made by counting








BULLETIN FLORIDA STATE MUSEUM


anteriorly from the caudal base to the scale at, or just posterior to, the dorso-posterior rim of
the operculum, in the manner proposed by Brown (1957:70). The last two dorsal and anal rays
were counted as separate rays. Caudal ray counts refer to branched rays only; no ontogenetic
changes in this character occur in these species complexes. Vertebral counts include the
hypural plate. Body depth was measured at the level of the pelvic fin base. Gill-raker counts
were made on the first (anterior most) arch.
Morphometric data, expressed as thousandths of standard length (hereafter referred to as
"SL") were maintained separately for the sexes, whereas meristic data for both sexes were
combined. For body proportions, 10 males and 10 females were measured for each species
from a given geographic area with the following exceptions: Fundulus grandissimus (Yuca-
tan), 3 specimens of each sex; F. majalis from the Savannah River to Matanzas Inlet, and from
Mosquito Lagoon southward, 15 specimens of each sex. Most morphometric data were not
significant and are not included in this paper; significant items are discussed. Morphometric
features examined were: lengths of head, snout, orbit, caudal peduncle, predorsal area, and
anal and dorsal fin bases; height of pectoral and pelvic fins; interorbital width; caudal
peduncle and body depth; and length of the anal sheath (ovipositor, females only).
Cephalic sensory pore and pit organ arrangements were also examined using Gosline's
(1949) study as a basis.
The statistical procedure used was the t-test (N > 30, two-tailed, p = .01) (see Sokal and
Rohlf 1969:220-221). The test is designed to compare population sample means by using the
formula:
X1 XS
t-=
S S,2 + S22
N2 N,
where X, and X2 are the means to be compared, S is the standard deviation, and N the number
of individuals of each sample to be compared. The null hypothesis in all cases is: The means
being compared are not different. A value of t of 2.58 or less would mean acceptance of the
null hypothesis with a 1% chance of error. Values of t greater than 2.58 suggest rejection of the
null nypothesis (i.e. the means can be considered significantly different at the level of
p = .01). These numbers do not mean that a population can automatically be considered a
subspecies or species. This is still a matter of judgment. The fragmentation of each species into
populations is arbitrary, but necessary for comparisons, and must be taken into account in
making taxonomic decisions.
Institutions from which specimens were examined are: Academy of Natural Sciences of
Philadelphia (ANSP), United States National Museum of Natural History (USNM), Univer-
sity of Michigan Museum of Zoology (UMMZ), Cornell University (CU), Tulane University
(TU), Florida State University (FSU)', University of Florida (UF), University of Miami
(UMIM), University of Miami Institute of Marine Science2, (UMML), and Jacksonville
University (JU).
KEY TO THE SPECIES OF THE Fundulus heteroclitus-Fundulus grandis
AND Fundulus majalis SPECIES COMPLEXES

1 a. Preorbital region scaled....................................2
b. Preorbital region unsealed .............................3
2 a. Dorsal fin rays 12-16, rarely 11 (only 12 of 1780
specimens examined had 11); New Hampshire to
northeastern Mexico.................................. Fundulus majalis
b. Dorsal fin rays 9-11, usually 10; known only from
Yucatan, Mexico ...................................Fundulus persimilis
These collections have been transferred to the Ichthyology Division, Florida State Museum, University of Florida, and are
now catalogued as UF-FSU.
'Now the Rosenstiel School of Marine and Atmospheric Science.


Vol. 29, No. 1








RELYEA: GENUS FUNDULUS


3 a. 8 mandibular pores.....................................4
b. More than 8 mandibular pores ..........................6
4 a. Anal rays 11-14, usually 12-13; gill rakers 12-15,
usually 12-13; predorsal scales 13-16, usually
14-15; caudal peduncle circumferential scales
18-20; known only from Bermuda ....................Fundulus bermudae
b. Anal rays 9-12, usually 10-11; gill rakers 8-17,
usually 10-13; predorsal scales 11-21, usually 12-18;
caudal peduncle circumferential scales 16-20 ............ 5
5 a. Caudal peduncle circumferential scales 19-20;
predorsal scales 13-21, usually 14-18; gill rakers
8-17, usually 10-13; Canada to northeastern Florida.......Fundulus heteroclitus
b. Caudal peduncle circumferential scales 16-17;
predorsal scales 11-17; usually 12-14; gill rakers
9-14, usually 10-11; known only from Bermuda ..........Fundulus rhizophorae
6 a. 10 mandibular pores; 13-19 predorsal
scales; northeastern Florida to north-
eastern Mexico; Cuba ................................Fundulus grandis
b. 12 mandibular pores; 21-25 predorsal
scales; known only from Yucatan Peninsula,
Mexico ..................................... ........Fundulus grandissimus

SYSTEMATIC ACCOUNTS
THE FUNDULUS HETEROCLITUS-FUNDULUS GRANDIS
SPECIES COMPLEX
A. Fundulus heteroclitus AND BERMUDA POPULATIONS
Fundulus heteroclitus (LINNAEUS)
FIGURE 1

Cobitis heteroclita Linnaeus 1766:500 (original description of F. heteroclitus, Charleston,
South Carolina; type specimen now in Linnaean Society of London collection); Goode and
Bean 1885:204-205 (type examined); specimen skinned and glued to herbarium sheet; skin
122 mm TL; counts: dorsal rays 12, anal rays 10, lateral scales 33); Jordan and Evermann
1896:641 (synonymy).
Cobitis killifish Walbaum 1792:12 (original description, Long Island, New York); Jordan and
Evermann 1896:641 (synonymy).
Poecilia macrolepidota Walbaum 1792:11 (original description, Long Island, New York);
Jordan and Evermann 1896:641 (synonymy).
Poecilia fasciata Bloch and Schneider 1801:453 (original description); Jordan and Evermann
1896:641 (synonymy).
Poecilia coenicola Bloch and Schneider 1801:452 (original description, Carolinas); Jordan and
Evermann 1896:641 (synonymy).
Hydrargyra swampina Lacepede 1803:378 (original description, South Carolina); Jordan and
Evermann 1896:641 (synonymy); Bailey and Wiley 1976:477-840 (evidence for retention of
H. swampina as a synonym of Fundulus heteroclitus).
Fundulus mudfish Lacepede 1803:37 (Carolinas); Jordan and Evermann 1896:641 (synonymy).
Esox pisciculus Mitchill 1815:440 (original description, New York); Jordan and Evermann
1896:641 (synonymy).
Esox pisculentis Mitchill 1815:441 (original description, New York); Jordan and Evermann
1896:641 (synonymy).








BULLETIN FLORIDA STATE MUSEUM


FIGURE 1. Fundulus heteroclitus (JU 775), St. Johns County, Florida. Upper: male, 60 mm
SL; Lower: female 64 mm SL.
Hydrargira nigrofasciata LeSeur 1817:133-134 (original description, Newport, Rhode Island);
Jordan and Evermann 1896:641 (synonymy).
Fundulus nigrofasciata Storer 1837:295 (original description, Massachusetts); Jordan and
Evermann 1896:641 (synonymy).
Fundulus pisculentis Storer 1837:294 (original description, Massachusetts); Jordan and Ever-
mann 1896:641 (synonymy).
Fundulus viridescens DeKay 1842:217 (original description, New York); Jordan and Ever-
mann 1896:641 (synonymy).
Fundulus zebra DeKay 1842:218 (original description, New York); Jordan and Evermann
1896:641 (synonymy).
Fundulus fonticola (?) Cuvier and Valenciennes 1846:198 (original description, Puerto Rico);
Jordan 1887:526 (examination of type specimen); Jordan and Evermann 1896:643 (des-
cription, Puerto Rico); Hubbs 1926:7 (F. fonticola = F. antillarum?); Jordan, Evermann,
and Clark 1930:175 (West Indies); Hildebrand 1935:49 (existence of F. fonticola doubtful);
Myers 1938:358-360 (existence doubtful); Rivas 1948:216-217 (existence doubtful); Miller
1955:3 (status of F. fonticola unclear).
Fundulus nisorius Cope 1870:490 (original description, Gaboon, West Africa); Fowler
1916:417-418 (species redescribed; allied to Fundulus heteroclitus); Hubbs 1926:6 (synon-
ymy; West African type locality presumably in error); Miller 1955:3 (synonymy).
Fundulus fasciatus Uhler and Lugger 1876:126 (original description, Maryland); Jordan and
Evermann 1896:641 (synonymy).
Fundulus vinctus Jordan and Gilbert 1882:354-356 (original description, Cape San Lucas,
California); Hubbs 1931:2-3 (synonymy; type locality presumably in error).
Fundulus antillarum Fowler 1916:418-420 (original description, St. Martins, West Indies);
Myers, 1938:358-360 (type locality F. antillarum doubtful); Rivas 1948:216-217 (synon-
ymy); Miller 1955:3 (synonymy; ANSP types examined; type locality presumably in error).
Fundulus heteroclitus grandis Bean 1891:92 (Chesapeake Bay).


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


Fundulus heteroclitus badius Garman 1895:96 (original description, Gulf of St. Lawrence,
Canada).
Fundulus heteroclitus heteroclitus Jordan and Evermann 1896:640-641 (original description,
Virginia to Florida); Brown 1957:73 (Gulf of St. Lawrence to Matanzas River, Florida).
Fundulus heteroclitus macrolepidotus Jordan and Evermann 1896:641 (original description,
Maine to Virginia); Breder 1948:83 (Gulf of St. Lawrence to Virginia); Brown 1957:73
(synonymy).

MATERIAL EXAMINED.-(The number of specimens follows catalogue number in paren-
theses.) LABRADOR: USNM 86223(2); NOVA SCOTIA: UMMZ 138840(36), 138834(15),
USNM 35339(14); QUEBEC: CU 18357(34), UMMZ 174659(4); MAINE: TU 43451(20),
43452(4), UMMZ 145494(80); NEW HAMPSHIRE: UMMZ 140525(80); MASSACHUSETTS:
CU 31101(46), 47234(31), TU 19743(10), 23221(9); CONNECTICUT: UMMZ 138841(100);
NEW YORK: UMMZ 103897(80), CU 38503(36); NEW JERSEY: CU 26970(12), 32197(7),
30365(4), UMMZ 109874(100); PENNSYLVANIA: UMMZ 99463(100); DELAWARE: UMMZ
99430(43), 99428(5), 99429(5); MARYLAND: TU uncatalogued (68), UMMZ 159890(22);
VIRGINIA: UMMZ 156714(6), 171290(20); NORTH CAROLINA: ANSP 80837(6), UMMZ
132806(3), 65237(1), 55997(4), 126285(29); SOUTH CAROLINA: ANSP 7509-19(11), UMMZ
139418(100); GEORGIA: ANSP 80833(29), 80834(36), TU 21339(160), 21232(477); FLORIDA:
FSU 6525(82), 10432(8), 8896(19), JU 36, 60, 111, 673, 644, 790, 476, 457, 252, 154, 1463, 709,
876, 838, 794, 806, 796, 775, 786, TU 41592(2), 41583(13), 41588(3), 41843(1), UF 2980(4),
USNM 104956(7).

DIAGNOSIS.-A species of Fundulus most closely related toF. bermudae
and F. rhizophorae, with which it shares the characters of 8 total mandibu-
lar pores and a long anal sheath in females (Table 6). Somewhat less closely
related to F. grandis (with which it is partly sympatric), from which it
differs in having 8 (vs. 10) total mandibular pores and a longer anal sheath
in females (see Table 6). Also distinguished from F. grandis by gill raker
number (8-17), usually 10-13, versus 8-12, mode 10, in sympatric F. gran-
dis), vertebral number (30-36, usually 32-34, vs. 33-37, usually 35-36), and
length of the anal sheath of females (59-233, population means are 87-200
expressed as thousandths of standard length; vs. 52-80, mean 69.0, in
sympatric F. grandis). Fundulus heteroclitus is distinguished from F. ber-
mudae by anal rays (9-12, usually 11, vs. 11-14, usually 12-13) and predorsal
scales (13-21, usually 15-16, vs. 12-16, usually 14); and from F. rhizophorae
by predorsal scales (13-21, usually 15-16, vs. 11-17, usually 12-14), caudal
peduncle circumferential scales (17-22, usually 19-20, vs. 15-18, usually
16-17), and pectoral fin rays (16-21, usually 17-19, vs. 15-18, usually 16-17).
Maximum body size in about 120 mm SL.
DEscRIPTON.-Meristic data, in addition to that given in the diagnosis
and in Tables 1-5, are as follows: pelvic fin rays 6-6; branchiostegal rays
5 + 5; dorsal fin rays 10-15 (mode = 12); anal fin rays 9-12 (mode = 11);
branched caudal rays 14-21, usually 16-18; caudal peduncle circumferential
scales 17-22, usually 19-20; and pectoral fin rays 16-21 (mode = 18-19).
The preorbital area is naked, and 4 irregular scale rows occur on the
operculum. There is an enlarged interorbital scale surrounded by a rosette







BULLETIN FLORIDA STATE MUSEUM


of 7 smaller scales.
Cephalic sensory pore arrangement has been described by Gosline
(1949). There are 4 preorbital pores. In northern populations a pit organ lies
between preorbital pores 2 and 3, but anterior of a line connecting those 2
pores. This pit organ is absent in many individuals in southern populations.
Of 68 specimens examined, 66 had 14 preopercular pores, 1 had 15, and 1
had 16.
The lower pharyngeal plates are narrow and elongated and have
numerous, small, conical teeth. The upper pharyngeal plates are rounded.
Mature males in life are colored blue to olive dorsally, shading to a
lighter olive ventro-laterally. The ventral region is bright yellow in breed-
ing males (hence the common name yellow bellied cobbler). Small, light
spots occur on the body, especially posteriorly. The median fins are blue
with light spots, and have a yellow or orange margin. Pelvic and pectoral
fins are also yellow or orange. An ocellus occurs in the dorsal fin of males,
but this is obscured by increasing pigmentation in the dorsal fin as the fish
matures (Fig. 1).
In males, 7-10 contact organs appear on the posterior rim of each body
scale, and numerous contact organs are found on the margins of the head
scales. These structures also occur on the rays of the dorsal and anal fins,
and occasionally on the pectoral fins.
Lateral scale number (Table 1) and predorsal scale number (Table 2)
show general north-south dines, with numbers decreasing southward, and
with significant differences in means between populations. These charac-
ters will not separate F. heteroclitus and F. grandis.
The mean number of gill rakers (Table 3) for F. heteroclitus south of the
St. Johns River, Florida (12.36), is statistically different from the mean for
Florida east coast F. grandis (9.95), where the two species are sympatric.
Northern specimens of F. heteroclitus (Chesapeake Bay northward) have
significantly fewer gill rakers (8-13, usually 10-12) than southern specimens
(10-17, usually 11-13).
There is no difference in number of caudal-peduncle circumferential
scales between F. heteroclitus and F. grandis, and little variation among F.
heteroclitus populations. The mode is 20 in all populations.
The pectoral ray mean for F. heteroclitus south of the St. Johns River
(18.50) differs from the mean for Florida east coast F. grandis (17.95). As
with gill rakers, northern and southern populations of F. heteroclitus may
also be indicated here (means < 18.28 vs. means > 18.50, respectively).
Vertebral number (Table 4) generally decreases southward with signif-
icantly different means between populations suggesting strong clinal patt-
erns. The mean number of vertebrae for southern F. heteroclitus (33.27)
differs significantly from the means for Florida east coast F. grandis
(35.27).


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


The total number of mandibular pores (Table 5) completely separates
F. heteroclitus and F. grandis (and F. grandissimus), and is the best single
diagnostic character. The mode for F. heteroclitus is 8 (mean 7.95) and for
F. grandis is 10 (mean 9.94) with no overlap in counts.
MORPHOMETIC VARIATION.-No significant variation in morphometric
features was found. See discussion of morphometric variation for F. gran-
dis for comparisons with that species, and Table 6, which shows intraspe-
cific and interspecific variation for anal sheath length in females.
RANGE.-This species is known from northeastern Florida to southern
Newfoundland (Fig. 2).
DIscussIoN.-Four subspecies of Fundulus heteroclitus have been rec-
ognized (see synonymy). Bean (1891:92) recognized F. h. grandis from
Chesapeake Bay, but my data show no distinctive differences for any
feature for Chesapeake Bay populations. Garman (1895:96) described F. h.
badius from the Gulf of St. Lawrence, Canada, and this was later retained
as a valid form by Jordan et al. (1930:176). Breder (1948:83) included Gulf
of St. Lawrence specimens in F. h. macrolepidotus (Gulf of St. Lawrence to
Virginia), and Brown (1957:73) included Gulf of St. Lawrence specimens in
F. h. heteroclitus. Obviously, these latter authors did not consider F. h.
badius to be valid, and my data support this. Fundulus heteroclitus ranges
continuously from the Gulf of St. Lawrence to Florida, and Canadian
populations differ in no way from New England specimens. These are
included together in my tabulation of meristic features.
Although Brown (1957:73) recognized one subspecies from Canada to
Florida, F. h. heteroclitus, Jordan and Evermann (1896:640-641) had deli-
neated F. h. heteroclitus (Virginia to Florida) and F. h. macrolepidotus
(Maine to Virginia). My data suggest some differentiation of populations
from Cape Lookout northward through Canada from populations to the
south of Cape Lookout (see Remarks). One meristic feature, gill raker
number, does show a significant difference between populations north and
south of Cape Lookout, but a vague dine is indicated. Other meristic
features do not show significant differences, or clinal trends are clearly
evident if differences are apparent. One morphometric character, anal
sheath length of females, is shorter in populations north of Chesapeake
Bay, but this does not coincide with the defined subspecies F. h. heterocli-
tus and F. h. macrolepidotus. In addition, a general increase in anal sheath
length southward is clear from Table 6.
Some meristic variation may be expected on the basis of environmental
temperatures, but characters such as anal sheath length and gill raker
number point to more specific reproductive and feeding differences.
Brummett's (1966:616-620) demonstration that eggs ofF. heteroclitus from
Woods Hole have chorionic fibrils, in contrast to eggs from Beaufort
specimens, points to further differences between northern and southern






BULLETIN FLORIDA STATE MUSEUM


A?- ," Canada-
.Y- .. ,.. -S.,
.-^ ,h -,. ,..

Cape Co

!. ,* .- -. -

Iong island
"y.- r-L- o ng


12,

SChesapeake Bay


A," .- i J Cape Lookout

harleston
Savannah R.

1J vSt. Johns R.





Type Locality


FIGURE 2. Map of the geographic distribution of Fundulus heteroclitus.


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


populations. These meristic, morphometric, and egg morphology differ-
ences could be interpreted as evidence for the validity of the nominal
subspecies F. h. heteroclitus and F. h. macrolepidotus. However the differ-
ences do not coincide along distributional lines, and some variability
should be expected in a wide ranging species such as F. heteroclitus. I
choose not to fragment F. heteroclitus into subspecies, and in doing so
follow Brown (1957).

Fundulus bermudae GUNTHER
FIGURE 3






















FIGURE 3. Fundulus bermudae (USNM 164795), Mangrove Lake, Bermuda. Upper: male 85
mm SL; Lower: female 78 mm SL.
Fundulus bermudae Giinther 1874:370 (original description, Bermuda; precise type locality
unknown; present location of types unknown; counts: dorsal rays 14, anal rays 12, lateral
scales 35); Jordan and Evermann 1896:643-644 (in part: description); Bean 1906:35 (in part:
distribution); Beebe and Tee-Van 1933:57 (in part: description).
Fundulus heteroclitus bermudae Hubbs 1926:8 (synonymy); in part: distribution); Jordan,
Evermann, and Clark 1930:175 (synonymy); in part: distribution); Miller 1955:3,7 (in part:
distribution); Brown 1957:69, 73 (in part: distribution); Collette 1962:438 (Bermuda
endemic).
MATERIAL EXAMINED.-The number of specimens follows catalogue number in paren-
theses. BERMUDA: USNM 164793(105), 164794(127), Mangrove Lake.
DIAGNOSIS.-A species of Fundulus most closely related to F. heterocli-
tus and F. rhizophorae, with which it shares the characters of eight total
mandibular pores and a long anal sheath in females (Table 6). It is sympat-
ric with F. rhizophorae, from which it is distinguished by number of dorsal








BULLETIN FLORIDA STATE MUSEUM


fin rays (11-15, usually 13-14, vs. 10-13, usually 11-12), anal fin rays (11-14,
usually 12-13, vs. 9-12, usually 11), branched caudal fin rays (16-19, usually
17-18, vs. 13-18, usually 15-16), predorsal scales (12-16, usually 14-15, vs.
11-17, usually 12-14), gill rakers (12-15, usually 12-13, vs. 9-14, usually
10-11), caudal peduncle circumferential scales (18-20), vs. 15-18), and
pectoral fin rays (17-20, usually 18-19, vs. 15-18, usually 16-17). Fundulus
bermudae may be distinguished from F. heteroclitus by number of anal fin
rays and predorsal scales; F. heteroclitus has 9-12, usually 11, anal fin rays
and 13-21, usually 15-16, predorsal scales.
DEscRIPTION.-Fundulus bermudae has 31-36 (usually 33-34) vertebrae,
31-36 (usually 33-35) lateral scales, 14 preopercular pores, 5 branchiostegal
rays, and 6 rays in each pelvic fin, in addition to the meristric features given
in the diagnosis. See Table 7 for a summary of meristic features. Head
scalation and sensory pore arrangement are similar to those occurring in F.
heteroclitus.
I have not seen living material of this species. Gunther (1874:370)
described this species as being brownish-olive with faint dark green cross-
bands. There seems to be little difference between this species and F.
rhizophorae and F. heteroclitus in coloration of preserved specimens, with
the exception that F. bermudae has a wide cream-colored band extending
above the lateral line from the operculum to the caudal peduncle. As in the
other species, the median fins are heavily pigmented (presumably blue) in
males, and pale or clear in females. The anal sheath of females is long as in
the other two species, to which F. bermudae is obviously closely related
(see Table 6 for comparison of anal sheath length).
Maximum body size is about 100 mm SL.
RANGE.-This species is known only from Bermuda. Its current preval-
ence and distribution there are undetermined.
DIscussIoN.-Hubbs (1926) considered F. bermudae to be a subspecies
of F. heteroclitus. Because of its isolation and the degree of morphological
divergence from F. heteroclitus, I believe that Ginther's (1874) original
interpretation of F. bermudae as a distinct species should be accepted. The
long anal sheath of females and the eight total mandibular pores clearly ally
F. bermudae with F. rhizophorae, which also occurs on Bermuda, but
which is distinguished by several meristic features (see Diagnosis). The two
Bermuda species probably represent two invasions of Bermuda by heter-
oclitus or pre-heteroclitus populations, or (less likely) one invasion fol-
lowed by separation in isolated lacustrine situations.
Fundulus rhizophorae Goode 1877:298 (original description, Basden Pond, Bermuda; types
USNM 21943, but present whereabouts unknown; counts: dorsal rays 12, anal rays 11,
lateral scales 35).
Fundulus bermudae Jordan and Evermann 1896:643-644 (synonymy; in part: description);
Bean 1906:36 (in part: description); Beebe and Tee-Van 1933:57 (in part: description).


Vol. 29, No. 1








RELYEA: GENUS FUNDULUS


Fundulus heteroclitus bermudae Hubbs 1926:8 (synonymy; in part: distribution); Collette
1962:440 (synonymy).
MATERIAL EXAMINED.-The number of specimens follows catalogue number in paren-
theses. BERMUDA: JU 1550 (150), USNM 175776 (70), 112083 (247), all from St. George I.,
Lover's Lake.
Fundulus rhizophorae GOODE
FIGURE 4






















FIGURE 4. Fundulus rhizophorae (JU 1550), Lover's Lake, Bermuda. Upper: male42 mm SL;
Lower: female 43 mm SL.
DIAGNOSIS.-A species of Fundulus most closely related to F. bermudae
and F. heteroclitus, with which it shares the characters of eight total
mandibular pores and a long anal sheath in females (Table 6). It is distin-
guished from F. bermudae by number of dorsal fin rays (10-13, usually
11-12, vs. 11-15, usually 13-14), anal fin rays (9-12, usually 11, vs. 11-14,
usually 12-13), branched caudal fin rays (13-18, usually 15-16, vs. 16-19,
usually 17-18), predorsal scales (11-17, usually 12-14, vs. 12-16, usually
14-15), gill rakers (9-14, usually 10-11, vs. 12-15, usually 12-13), caudal
peduncle circumferential scales (15-18, usually 16-17, vs. 18-20, usually 19),
and number of pectoral fin rays (15-18, usually 16-17, vs. 17-20, usually
18-19). Fundulus rhizophorae is distinguished from F. heteroclitus by
number of predorsal scales, caudal peduncle circumferential scales, and
pectoral fin rays; F. heteroclitus has 13-21 (usually 14-18) predorsal scales,
17-22 (mode 19-20) caudal peduncle circumferential scales, and 16-21
(usually 17-19) pectoral fin rays.








BULLETIN FLORIDA STATE MUSEUM


DIAGNOSIS.-A species of Fundulus most closely related toF. bermudae
and F. heteroclitus, with which it shares the characters of eight total
mandibular pores and a long anal sheath in females (Table 6). It is distin-
guished from F. bermudae by number of dorsal fin rays (10-13, usually
11-12, vs. 11-15, usually 13-14), anal fin rays (9-12, usually 11, vs. 11-14,
usually 12-13), branched caudal fin rays (13-18, usually 15-16, vs. 16-19,
usually 17-18), predorsal scales (11-17, usually 12-14, vs. 12-16, usually
14-15), gill rakers (9-14, usually 10-11, vs. 12-15, usually 12-13), caudal
peduncle circumferential scales (15-18, usually 16-17, vs. 18-20, usually 19),
and number of pectoral fin rays (15-18, usually 16-17, vs. 17-20, usually
18-19). Fundulus rhizophorae is distinguished from F. heteroclitus by
number of predorsal scales, caudal peduncle circumferential scales, and
pectoral fin rays; F. heteroclitus has 13-21 (usually 14-18) predorsal scales,
17-22 (mode 19-20) caudal peduncle circumferential scales, and 16-21
(usually 17-19) pectoral fin rays.
DEscRIPTION.-Fundulus rhizophorae has 31-34 (usually 32-33) verte-
brae, 14 preopercular pores, 5 branchiostegal rays, and 6 rays in each pelvic
fin, in addition to the characteristics given in the diagnosis. Table 7 sum-
marizes meristic data.
Goode (1877:298) described F. rhizophorae as being a light, tawny
yellow with transverse bands of greenish-brown. I have examined freshly
preserved material, from which I present the following additional informa-
tion: Males resemble F. heteroclitus males. The body is olive to blue in life
with light spots, especially posteriorly. The ventral region is yellow, as in F.
heteroclitus. Median fins are heavily pigmented (blue) with light spots.
Females are drab brown (tawny?) with faint, dark vertical bars, as is true
also of F. heteroclitus.
Males possess 7-10 extremely long contact organs on the rim of each
body scale. Contact organs also occur on the head, dorsal fin, and anal fin.
Maximum size attained is about 65 mm SL. This is somewhat smaller
than the maximum size of F. bermudae and F. heteroclitus.
RANGE.-Fundulus rhizophorae is known only from Bermuda. In addi-
tion to the type locality, the species is known from Lover's Lake on St.
Georges Island (USNM 175776, USNM 112083, JU 1550). The species is
particularly abundant at this latter locality, and seems to be isolated from F.
bermudae. Even though two species of Fundulus occur on Bermuda, they
apparently are not sympatric.
DIscussioN.-Jordan and Evermann (1896) placed F. rhizophorae in the
synonymy of F. bermudae. In my opinion, these two forms are distinct
species, both derived from F. heteroclitus, and I believe that the meristic
data presented here argue for that conclusion. In addition, F. rhizophorae
appears to be smaller than the other two species. The possibility exists that
the degree of difference between F. rhizophorae and F. bermudae could


Vol. 29, No. 1








RELYEA: GENUS FUNDULUS


be interpreted as merely ecophenotypic variation. However, given the
allopatric distribution of both Bermuda forms and F. heteroclitus and the
meristic differences existing among them, specific status for each seems
reasonable.

B. Fundulus grandis grandis AND Fundulus grandis saguanus
Fundulus grandis grandis BAIRD AND GIRARD
FIGURE 5























FIGURE 5. Fundulus grandis grandis (JU 842), Franklin County, Florida. Upper: male 105
mm SL; Lower: female 104 mm SL.
Fundulus grandis Baird and Girard 1853:389 (original description, Indianola, Texas; two male
syntypes, 99-101 mm SL, USNM 15298).
Fundulus grandis grandis Rivas 1948:215-222 (original description of F. g. saguanus and
comparison with F. g. grandis); Brown 1957:73 (identification in key; distribution).
Fundulus pallidus Evermann 1892:84 (original description, Galveston, Texas); Garman
1895:96 (F. pallidus placed in synonymy of F. grandis); Miller 1955:5-7 (synonymy).
Fundulus floridensis Girard 1859:157 (original description, Charlotte Bay, Florida); Garman
1895:96 (F. floridensis placed in synonymy ofF. grandis); Jordan and Evermann 1896:641-
642 (synonymy).
Fundulus heteroclitus Jordan and Gilbert 1883:334 (distribution); Evermann 1892:84 (Texas);
Nichols 1942:125-126 (Gulf coast of Florida).
Fundulus heteroclitus grandis Jordan and Swain 1884:230 (Cedar Keys, Florida); Garman
1895:96 (F. grandis and F. heteroclitus clearly distinguished); Jordan and Evermann,
1896:641-642 (Gulf of Mexico); Jordan, Evermann, and Clark 1930:176 (Gulf of Mexico);
Brown 1957:73 (distributions of F. grandis and F. heteroclitus delineated and species
recognized as distinct).








BULLETIN FLORIDA STATE MUSEUM


MATERIAL EXAMINED.-The number of specimens follows catalogue number in paren-
theses. FLORIDA: Florida State Board of Conservation 1225(6), 1288(9), 903(6); FSU
11210(55), 11474(110), 12413(12); JU 113, 1618, 44, 1730,1499, 1464, 1011, 1024,903, 912, 922,
875,890,891,691,686,780,770,798,842,839,639,530,475,453,367,164,86,82; TU 39581(10),
38091(4), 39587(1), 39616(8), 20477(11), 20477(11), 20388(9), 18183(5), 12539(5), 41206(14),
21261(57), 20366(11), 2273(112), 41387(2), 42448(1), 41069(133), 40562(9), 9742(37), 21385(61),
18421(29), 20959(6), 18355(44), 20424(1), 41069(133), 40562(9), 9742(37), 21385(61), 18421(29),
20959(6), 18355(44), 20424(1), 18220(4), 41566(15), 41077(11), 17815(15), 17846(5), 18389(129),
17915(4), 41584(3); UF 9693(1), 3483(5), 4013(8), 5215(1), 4014(4), 12279(1), 1490(8), 4662(11),
6683(5), 2980(11), 3116(10); UMIM 3152(3); UMML 3264(50); USNM 62668(2), 25310(1).
ALABAMA: TU 8577(9), 14514(2), 15854(2). MISSISSIPPI: JU 235(100); TU 10861(32),
16109(107). LOUISIANA: TU 11492(27), 19048(11), 4344(11), 37219(6), 1394(4), 6425(16),
9253(21), 6546(81), 17338(16), 8271(4). TEXAS: JU 1639,1645,1657,1683,1688; TU 39872(4),
2163(75), 22190(76), 21505(195), 22063(15), 21474(6), 41099(7); USNM 15298(2), Indianola,
1853, (Cotypes). MEXICO, TAMAULIPAS: TU 38959(14), Rio Purificacion, 38.5 km NW of
Victoria, Hwy 85, 25 Dec. 1964; TU uncat. (18), 11 km W of Tampico on Hwy 110, 23 May
1961; TU uncat. (59), Rio Purificacion, Caracol, Hwy 85, 20 May 1961; TU uncat. (8), Arroyo
Chorreras, S of San Jose las Norias, 17 Dec. 1960; TU uncat. (3), Arroyo Chorerras, 19 km S of
San Fernando, Hwy 101, 22 May 1962; UF 25975 (30)

DIAGNOSIS.-A subspecies of Fundulus grandis most closely related to
F. g. saguanus, F. grandissimus, and slightly less closely related to F.
heteroclitus, F. bermudae, and F. rhizophorae. Distinguished from F.
heteroclitus by number of gill rakers (7-13, usually 9-11, vs. 10-15, usually
12-13 in sympatric F. heteroclitus), vertebrae (31-36, usually 34-35, vs.
31-37, usually 33-34), total mandibular pores (10 vs. 8), and the short anal
sheath of females (34-103, as thousandths of standard length, vs. 166-194 in
sympatric F. heteroclitus). Fundulus g. grandis is distinguished from F.
grandissimus by number of lateral scales (30-38, usually 33-36, vs. 36-42,
usually more than 38), predorsal scales (13-19, usually 15-17, vs. 21-25,
usually 23-24), caudal peduncle circumferential scales (17-23, usually 20,
vs. 21-25, usually 24), and total mandibular pores (10 vs. 12). Fundulus
grandis grandis is distinguished from the allopatric F. g. saguanus by
number of lateral scales (30-38, usually 33-36, vs. 30-35, usually 32-33), gill
rakers (7-13, usually 9-11, vs. 7-11, usually 9), vertebrae (31-36, usually
34-35, vs. 31-35, usually 32-33), and pectoral fin rays (15-21, usually 17-19,
vs. 16-19, usually 17).
DEsCRIPTION.-Meristic features, in addition to those in Tables 1-5 and
in the Diagnosis, are pelvic fin rays 6-6, branchiostegal rays 5+5, dorsal fin
rays 10-14 (usually 11-12), anal fin rays 10-15 (usually 11), branched caudal
fin rays 15-21 (usually 16-18). Of 136 specimens examined, 132 had 14
preopercular pores, 3 had 15, and 1 had 16. Four irregularly arranged scale
rows occur on the preoperculum. The preorbital area is naked, and the
preorbital pore series consists of four pores with a pit organ directly
between numbers two and three. An enlarged interorbital scale is sur-
rounded by a rosette of seven smaller scales.


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


Males have 7-10 contact organs on the posterior rim of each scale on the
body, and many small contact organs on each head scale. Contact organs
also occur on the rays of the dorsal and anal fins, and occasionally on the
pectoral fins.
The lower pharyngeal plates are elongate and narrow, and have many
small conical teeth. The upper pharyngeal plates are rounded.
Breeding males are deep blue in color dorsally, shading to a lighter olive
ventrally. The body coloration is interrupted by light, narrow, vertical,
irregular bars. Numerous small, light spots occur on the body, especially
posteriorly. The median fins are blue, with numerous light spots, and have
a yellow or orange margin. The pelvic fins are yellow or orange, and have
many scattered melanophores.
A prominent dark blotch on the operculum and a dark predorsal stripe
that fades with age are present in both sexes. Many scattered melanophores
occur on the anterior and posterior portions of the pectoral fin base of both
sexes.
Females are drab olive dorsally, shading to white ventrally and the fins
are cream colored or yellow. Juveniles are generally olive, but their most
distinctive coloration feature is a bright yellow anal fin. Some juveniles
have darker vertical bars on the lighter olive ground color, but these
disappear at 15-20 mm SL.
Maximum size is about 145 mm SL.
TYPES.-Of the two syntypes, the smaller (99 mm SL) has 12 dorsal rays,
11 anal rays, 19 branched caudal rays, 35 lateral scales, 14 predorsal scales,
10 gill rakers, 19 pectoral rays, and 21 caudal peduncle circumferential
scales. The other syntype (101 mm SL) has 13 dorsal rays, 12 anal rays, 18
branched caudal rays, 35 lateral scales, 9 gill rakers, 17 pectoral rays, and 19
caudal peduncle circumferential scales. The predorsal region of this spec-
imen is damaged. Both specimens have a total of 10 mandibular and 14
preopercular pores.
RANGE.-Fundulus grandis grandis ranges from Laguna de Tamiahua,
Veracruz, Mexico (Miller 1955), eastward along the coast of the Gulf of
Mexico to at least Collier County, Florida (southwest Florida, UMIM 3152
and 3264), and disjunctly on the east coast to Florida from Nassau County
(northeastern Florida, JU 36,44, 111, 113,1618) southward to Lake Worth,
Palm Beach County (UF 1490, 4662). The gap between the east coast and
Gulf of Mexico populations is occupied by F. g. saguanus (extreme south-
ern Florida, Florida Keys, Cuba). Fundulus grandis is sympatric with F.
heteroclitus for about 100 miles of coastline in northeastern Florida (Fig. 6).
MERISTIC VARIATION.-The modal number of dorsal rays is 12 in popula-
tions in the Gulf of Mexico east of the Mississippi River and 11 in popula-
tions west of the Mississippi River and on the east coast of Florida. Most
specimens of F. grandis have either 11 or 12 dorsal rays. Atlantic and Gulf













ral
S Cape Canave al
Tampa Ba ------


Gulf of Mexico R "
/ y oa Raton


ey est

ya Uvero
Progreso RioLogr os
Sisal Fundulus grandis grandis
is Type locality- Eg. grandis
Yucatan E g. saguanus
(Type locality= Playa Uvero) '
o Egrandissimus
(Type locality= Progreso)

FIGURE 6. Map of the geographic distribution of Fundulus grandis and Fundulus grandissimus.







RELYEA: GENUS FUNDULUS


coast means are statistically different (11.41 and 12.04, t=7.87), and signifi-
cantly different means between Gulf of Mexico populations may indicate
strong clinal patterns.
The mode of anal rays is 11 in all populations, and means only show
significant variation in the western Gulf of Mexico. Fundulus heteroclitus
also has 11 anal fin rays.
The number of branched caudal rays is extremely variable, no clear
modes are evident, and any statistical difference in means is slight. Approx-
imately 89% of all specimens examined had either 16, 17, or 18 branched
caudal rays.
The modal number of lateral scales (Table 1) is 35 in Gulf of Mexico
populations and 34 on the east of Florida. The east coast race mean (34.37)
is significantly different from the southwestern Florida population mean
(34.72). No major significant variation occurs in Gulf populations, and no
clinal patterns are evident.
The modal number of predorsal scales (Table 2) is 16 in all populations
except in the western Gulf of Mexico (mode = 15), but no difference is
apparent in the western and eastern Gulf means. The Atlantic coast popula-
tion mean differs from the southwestern Florida mean.
The modal gill raker count west of the Mississippi River is 10 (Table 3),
shifts to 9 in the region from the Mississippi River to Cape San Bias, is 10
again farther east and south in the Gulf, and is 10 on the Florida east coast.
The modal number of caudal peduncle circumferential scales is 20 in all
populations. The mean number increases clinally southward along the
Florida Gulf coast, but statistical differences are marginal for compared
means.
The modal number of pectoral rays is 18 in all populations. There is a
distinct increase in mean number from north to south in the Gulf of Mexico.
The western Gulf mean (18.21) differs statistically from eastern Gulf means
(17.90), and the Florida east coast mean (17.95) differs significantly from
the southwestern Florida mean (18.38).
No significant variation occurs for vertebral number (Table 4), the
mode remaining 35 in all populations. Mean vertebral number differs
significantly between southern F. heteroclitus (33.27) and Florida east
coast F. grandis (35.27).
Total mandibular pores (Table 5) are typically 10 in nearly all
specimens.
MORPHOMETRIC VARIATION.-Although no significant morphometric
data were obtained, one feature is worth noting. The length of the anal
sheath in females (Table 6) permits separation of nearly all populations of
F. grandis and F. heteroclitus (except for F. heteroclitus north of Cape Cod
where F. grandis does not occur). The length of the anal sheath of female F.
g. grandis (Florida east coast) ranges from 52 to 80 (thousandths of stand-
ard length, mean 69.0), whereas the longer anal sheath of female F. heteroc-








BULLETIN FLORIDA STATE MUSEUM


litus ranges from 166 to 194 (mean 178.8). In F. grandis, the sheath encloses
only about one-half the length of the first anal ray, whereas in F. heterocli-
tus the first anal ray is completely enclosed. This character is useful for field
identification.
Fundulus grandis saguanus RIVAS
FIGURE 7





















FIGURE 7. Fundulus grandis saguanus (CU 23975), Monroe County, Florida. Upper: male 92
mm SL; Lower: female 87 mm SL.

Fundulus grandis saguanus Rivas 1948:215-222 (original description, Playa Uvero, Cuba,
holotype, USNM 132419, 75 mm SL; 14 paratypes, USNM 132420, 49-65 mm SL); Miller
1955:6, 8 (Cuba); Brown 1957:69 (Cuba).
Fundulus grandis Kilby and Caldwell 1955:203 (ecology; Snake Bight Canal, Florida); Briggs
1958:265 (in part: distribution); Tabb and Manning 1961:614 (ecology; Florida Bay).
MATERIAL EXAMINED.-The number of specimens follows catalogue number in paren-
theses. CUBA: USNM 132419(1), Holotype; USNM 132420(10), Paratypes. FLORIDA: ANSP
79273(4), 97096(6), 89772(1), 74883(6); CU 587(2), 48360(50), 48374(55), 48340(1), 48354(17),
48294(18), 48335(9), 48330(135), 23975(8), 45408(12); FSU uncatalogued (8); JU 718,869,938,
951, 1029, 1448; TU 37200(2); UMIM 5727(5), 5723(17); UMML 419(2), 4374(2), 319(2),
3306(2), 6585(8), 772(1), 3112(28); UF 8632(1), 5091(43); USNM 131871(2), 126799(2),
124660(2), 116897(2).

DIAGNOSIS.-A subspecies of Fundulus grandis, distinguished from the
typical subspecies by number of lateral scales (30-35, usually 32-33, vs.
30-38, usually 33-36), gill rakers (7-11, usually 9, vs. 7-13, usually 9-11),
vertebrae (31-36, usually 32-33, vs. 31-36, usually 33-35), and pectoral fin
rays (16-19, usually 17, vs. 15-21, usually 17-19). Fundulus g. saguanus has
the 10 mandibular pores and short anal sheath ofF. g. grandis, and may be


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


distinguished from F. heteroclitus on the basis of those characters. The total
of 10 mandibular pores also distinguishes F. g. saguanus from F. grandissi-
mus, which has 12 mandibular pores.
DESCRIPTION.-See Tables 1-6 for summary of meristic characters.
Except for those features noted above, these characters are the same as
those found in the typical subspecies.
Coloration is similar to that ofF. grandis grandis, but overall coloration
is much lighter. The blue color of males is pale, and the ventral area is
white. Median fins are not so heavily pigmented as in F. grandis grandis,
and the light spots on the body are more prominent. This lighter coloration
may be correlated with the clear or milky colored water in which this
subspecies lives.
Melanophores are not scattered along the base of the pectoral fin as in F.
g. grandis, but are concentrated into a distinct blotch around the dorsal
origin of the pectoral fin. Usually no melanophores occur on the rest of the
fin base, but on some individuals a few scattered melanophores may occur
there anteriorly and posteriorly. The pelvic fins of males also lack melano-
phores, or have only a few, which is in sharp contrast to F. g. grandis, in
which the pelvic fins have a heavy concentration of melanophores.
Contact organs are poorly developed on F. g. saguanus males. They are
neither as large nor as numerous as on F. g. grandis, and do not appear at all
on many specimens in breeding condition.
The pharyngeal plate is similar to that of F. g. grandis, but the individual
teeth are somewhat larger and more recurved.
Maximum size is about 150 mm SL.
TYPES.-The holotype (USNM 132419), a female 75 mm in standard
length, was collected 18 km northeast of Sagua La Grande, at Playa Uvero,
Las Villas Province, Cuba. The 14 paratypes (USNM 132420: 9 females
38-66 mm SL, 5 males 49-65 mm SL), bear the same locality data as the
holotype. Four of the paratypes are no longer at the USNM and presuma-
bly are deposited elsewhere. These are the only Cuban specimens known.
RANGE.-This subspecies ranges from Cuba (type material) and the
Dry Tortugas (USNM 116897) northward through the Florida Keys and
Florida Bay (westward to Cape Sable, UMIM 5723) to Boca Raton on the
Florida east coast (JU 951, 2 specimens). It is common at least as far north as
the Miami area. It does not appear to contact or intergrade with popula-
tions of F. g. grandis on either the Gulf or Atlantic coasts (Fig. 6).
MERISTIC VARIATION.-Little variation in meristic features occurs
throughout the range of the subspecies, and Cuban specimens closely
resemble specimens from the Florida Keys and Florida bay. Fundulus g.
saguanus differs from F. g. grandis in a number of meristic features (Tables
1-4; and the Diagnosis). In addition, the mean number of pectoral fin rays
for F. g. saguanus (17.44) differs significantly from populations of the








BULLETIN FLORIDA STATE MUSEUM


typical subspecies from southwestern Florida (mean 18.38) and from the
Florida east coast (mean 17.95). The mode for the latter two populations is
18, and is 17 for F. g. saguanus from the Florida Keys and Cuba. Also of
note is that clinal trends in pectoral fin ray and predorsal scale numbers
seen in F. g. grandis are reversed in F. g. saguanus.
MORPHOMETMIC VARIATION.-Fundulus g. saguanus differs little from F.
g. grandis in morphometric features, and consequently these data are not
presented here. However, Rivas (1948:219), in his description of F. g.
saguanus from Cuba, distinguished this form from F. g. grandis (including
Florida Keys specimens in F. g. grandis) on the basis of several morpho-
metric features. Although the Cuban specimens may differ from the F. g.
grandis in the northern Gulf of Mexico, they do not differ from Keys
specimens. Rivas (1948) reported the range for caudal peduncle length of
the Cuban specimens as 4.1-4.5 (ratio of standard length), and the range for
head length as 2.7-3.0 (also in standard length). My data for specimens from
the Florida Keys give a range of 3.5-4.3 and 2.8-3.4 for these characters,
respectively. Rivas (1948) also reported the caudal peduncle depth as
1.9-2.2 in head length, and the interorbital width as 2.1-2.4 in head length.
My figures for these characters are, respectively, 1.8-2.1 and 2.2-2.4, for
Keys' specimens. Subsequently, on the basis of these morphometric data
and previously discussed meristic data, the Florida Keys' populations are
placed with F. g. saguanus.
DIscussIoN OF Fundulus grandis.-Populations of F. grandis from the
Gulf of Mexico seem to be continuous from Mexico (several TU collec-
tions) to southwestern Florida, over which range little morphological varia-
tion occurs. Slight differences in meristic features can be seen to the east
and west of the Mississippi River, but recognition of subspecies is not
warranted.
The allopatric population of F. grandis on the Atlantic coast is much
more similar to the Gulf populations than to those from extreme south
Florida and Cuba. If there are F. grandis between Boca Raton and Lake
Worth, they are very rare, and gene flow is surely minimal.
The extreme south Florida population, F. g. saguanus (including the
lower Florida east coast, Florida Keys, and Cuba) appears to be allopatric
to both the Gulf and Atlantic populations, and differs markedly from both
in most meristic characters (Tables 1-4). There are no characters that will
distinguish Florida Keys (north to Boca Raton) populations from the nomi-
nal Cuban subspecies F. g. saguanus (Rivas 1948), and they should be
referred to that subspecies. My original inclination (Relyea 1967) to elevate
F. g. saguanus (inclusive of the Florida Keys) to species level may well be
correct. Allopatric populations such as this, however, with a low degree of
differentiation, cannot be easily placed taxonomically. Obviously what is
important is the elucidation of the population as allopatric and somewhat


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RELYEA: GENUS FUNDULUS


distinct. Perhaps other workers using different criteria (i.e. biochemical or
chromosomal) will now be able to progress with the species concept in
allopatric killifish populations, such as in the Florida Keys. In my opinion,
killifish populations are too ecophenotypically plastic to be readily and
simply allowed to species or subspecies on the basis of variable body
proportions or indices derived from combinations of traits. Different pop-
ulations differ in different ways.
Experimental work with the influence of temperature and other envir-
onmental factors on development of morphometric, meristic, and pigmen-
tary features is needed. Studies of zygote viability in intraspecific crosses
would also be of interest (i.e. Gulf of Mexico vs. Atlantic coast populations
of F. grandis).
Fundulus grandis from the Gulf of Mexico and Atlantic coast of Florida
are more like one another than either is to extreme south Florida F. g.
saguanus and should be recognized as disjunct populations of the same
subspecies, F. g. grandis.
C. YUCATAN POPULATIONS, Fundulus grandissimus
FIGURE 8






















FIGURE 8. Fundulus grandissimus (JU 1699), Chiquila, Quintana Roo, Yucatan Peninsula,
Mexico. Male 68 mm SL.
Fundulus grandissimus Hubbs 1936:209-210 (original description, Progreso, Yucatan, Mexico;
Holotype, MCZ 32909, 179 mm SL); Miller 1955:1 (Yucatan); Brown 1957:69 (Yucatan);
Miller 1966:788 (Yucatan).







BULLETIN FLORIDA STATE MUSEUM


Fundulus grandis Evermann and Goldsborough 1902:149 (Yucatan; error in identification);
Barbour and Cole 1906:156 (Yucatan; error in identification).
MATERIAL EXAMINED.-The number of specimens follows catalogue number in paren-
theses. MEXICO, YUCATAN: TU uncatalogued (11), Lagoon 0.5 mi. S. of Progreso, 31 May
1961; USNM 50513(15), Progreso, 1935, Paratypes; FSU 27222(18), Progreso Mexico, Quin-
tana Roo: JU 1679(20), at Chiquila, March 1979.

DIAGNOSIS.-A species of Fundulus most closely related to F. grandis,
but differing from that species in the number of lateral scales (36-42,
usually 38-41, vs. 30-38, usually 33-36), predorsal scales (21-25, usually
23-24, vs. 13-19, usually 15-17), caudal peduncle circumferential scales
(21-25, usually 22-24, vs. mode of 20 in F. grandis), pectoral fin rays (15-18,
usually 16, vs. mode of 18), and total number of mandibular pores (12, vs.
10). It also reaches a larger maximum size, the holotype measuring nearly
30 mm longer than the largest known specimens of F. grandis (179 mm vs.
150 mm SL).
DESCRIPTION.-In addition to the meristic features summarized in
Tables 1-5 and in the Diagnosis, F. grandissimus has 14 preopercular pores,
5+5 branchiostegal rays, 6 rays in each pelvic fin, 10-12 dorsal fin rays, 10-12
anal fin rays, and 15-18 branched caudal fin rays. Cephalic squamation and
sensory pore arrangement are similar to those of F. grandis.
Coloration is similar to that of F. grandis. Males are blue to olive with
light vertical streaks and light spots, especially posteriorly, whereas
females are drab olive. Median fins are blue with light spots.
Maximum size is about 180 mm SL.
Contact organs appear to be poorly developed. Females have a short
anal sheath, a feature that allies this species with F. grandis (see Table 6).
TYPEs.-The holotype (MCZ 32909) is a female, 179 mm SL collected at
Progreso, Yucatan, Mexico. The USNM collection has 15 paratypes
(USNM 50513), consisting of 12 females 108-131 mm SL and 3 males
114-142 mm SL. Hubbs (1936) mentioned 10 more paratypes (3 females,
131-151 mm SL and 7 males 97-125 mm SL).
RANGE.-This species is known from Progreso, Sisal, and Rio Lagartos,
Yucatan, and Chiquila, Quintana Roo, in the Yucatan Peninsula of Mexico
(Fig. 6).
DIscussoN.--The most notable meristic features ofF. grandissimus are
given in the Diagnosis. Especially noteworthy is the total number of man-
dibular pores (12), a feature that readily distinguishes F. grandissimus from
the closely related F. grandis. Body proportions are similar to F. grandis.
Hubbs (1936:159) stated that F. grandissimus apparently resembles F.
grandis from the Florida Keys (interpreted here as F. g. saguanus). My data
show that F. grandissimus and F. g. saguanus represent opposite extremes
of divergence within the F. grandis population complex. Meristic counts
are high for F. grandissimus, with marked, statistically significant differen-


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RELYEA: GENUS FUNDULUS


ces in means when compared to F. grandis for lateral scales, predorsal
scales, caudal peduncle circumferential scales, and total mandibular pores.
Meristic counts are low for F. g. saguanus, which has the number of
mandibular pores (10) typical of F. grandis.

ECOLOGY
Fundulus grandis and F. heteroclitus inhabit bays and tidal marshes,
which usually have a mud or sand-mud substrate. The habitat of F. grandis-
simus is similar, but much less is known of that species. Fundulus grandis
has been recorded from salinities ranging from freshwater (Tabb and
Manning 1961) to 760/oo (Simpson and Gunter 1956). Fundulus heteroclitus
is also extremely euryhaline, having been recorded from freshwater (Hil-
debrand and Schroeder 1928; Massmann, 1954), and is common in brackish
situations.
Chichester (1920) observed spring migrations of F. heteroclitus in the
Raritan River, New Jersey, from the mouth of the river to more brackish
and freshwater areas, where they spend the summer and breed. In colder
weather they leave the river and adjacent marshes and go into the deeper
waters of the bays. If trapped in cold waters, they burrow in mud to a depth
of 6-8 inches. Bigelow and Schroeder (1953) and Hildebrand and Schroeder
(1928) also observed this burrowing behavior. Because of the more south-
erly distribution ofF. grandis, it is doubtful that this species undergoes such
a behavior pattern, and I have not noted F. grandis to burrow even when
alarmed. It is my impression that many individuals of F. heteroclitus
remain active throughout the winter months in northeastern Florida;
whether any individuals burrow there is unknown.
Burger (1939) and Matthews (1938, 1939), working at Woods Hole,
demonstrated that water temperature, not photoperiod, affects gonadal
development in F. heteroclitus, and the same is probably true for F.
grandis. Joseph and Yerger (1956) reported a spawning period of March
through September for F. grandis on the Gulf coast, and Simpson and
Gunter (1956:130-131) collected spawning individuals in Texas in October.
Springer and Woodburn (1960) felt that F. grandis spawned from late fall
through early spring in Tampa Bay, while Kilby (1955:197-198) suggested a
continuous breeding season at Cedar Keys and Bayport. I have collected
spawning fish through November on the northern Gulf coast. Apparently,
southern populations of F. heteroclitus also spawn throughout warm
months of the year (personal observation, Duval County, Florida).
Fundulus heteroclitus in the Gulf of Maine spawn from June to August
(Bigelow and Schroeder 1953:163). In Chesapeake Bay the spawning
period is somewhat longer, April through August (Hildebrand and
Schroeder 1928:139), and becomes still longer southward (Brummett
1966:616-620).







BULLETIN FLORIDA STATE MUSEUM


Newman (1907) gave an account of the spawning behavior of F. heter-
oclitus. The male clasps the female with his dorsal and anal fins, and the
pelvic fins of both individuals are forced against a rock or the substrate.
The clasping posture lasts for about two seconds, during which time the
gametes are released. Eggs are placed against rocks, oysters, debris, or at
the base of Spartina or Juncus. Taylor et al. (1979) demonstrated a lunar
periodicity of oviposition in which eggs are placed at the base of Spartina
on the highest monthly tide and left stranded above the high tide mark until
the next new or full moon. I have watched F. grandis spawning at Lanark
Beach, Franklin County, Florida. The male forces the female into coves,
clumps of debris, or among oysters and assumes a clasping posture similar
to that of F. heteroclitus. For further information on killifish reproduction
see Foster (1967a,b).
The small eggs of F. heteroclitus and F. grandis are demersal and
adherent. Hubbs and Drewery (1959) and Newman (1908) have success-
fully obtained artificial hybrids between F. heteroclitus and F. grandis
females and F. majalis males. Natural hybrids are known between F.
diaphanus and F. heteroclitus (Hubbs, Walker, and Johnson 1943).
Members of this species complex form schools, but not to the extent of
F. majalis. Feeding groups of 12-20 individuals are frequently encountered.
Food consists primarily of molluscs, crustaceans, and small fishes. I have
removed the following fishes from the oral cavity of F. grandis: F. majalis,
F. grandis, Cyprinodon variegatus, Gambusia affinis, and Gobiosoma sp.
Radiographs show fish remains in the gut of many specimens of both F.
grandis and F. heteroclitus. Fundulus grandis in aquaria readily eat Gam-
busia affinis. Fundulus heteroclitus is known to feed on gammarid
amphipods (van Dolah 1978).
Seine hauls that yield Fundulus heteroclitus or F. grandis also often
yield F. majalis. Fundulus majalis is a bottom feeder and does not generally
eat other fishes, whereas F. heteroclitus and F. grandis are middle-level
feeders, or top feeders, and do prey on fishes. The perplexing problem is
that where F. heteroclitus and F. grandis are sympatric, they appear to be
fully syntopic, even to the point of forming mutual schools. I, and others,
have collected both species in the same short haul of a seine. No habitat
difference is detectable between them. As the breeding seasons of both
species are extensive, there seems to be no temporal separation, and yet I
have seen no hybrids. Perhaps sexual recognition is well developed in each
species, and interspecific mating is therefore rare. This implies a situation
similar to that in F. notatus and F. olivaceus, in which the species are quite
similar superficially (to the human eye), but which hybridize very infre-
quently (Thomerson 1966). It is also possible that interspecific matings
occur in nature, but zygotes are inviable.


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RELYEA: GENUS FUNDULUS


REMARKS
The patterns of geographic variation in this species complex are: (1)
possible racial differentiation of Fundulus heteroclitus north and south of
Cape Lookout; (2) isolation of two species, F. bermudae and F. rhizopho-
rae from a F. heteroclitus stock on Bermuda; (3) differentiation of F.
heteroclitus and F. grandis as full species (sympatry in northeastern Flor-
ida; diagnostic features are length of anal sheath of females and number of
mandibular pores); (4) differentiation of an extreme south Florida popula-
tion (and Cuba) which is referred to F. grandis saguanus; (5) possible racial
differentiation of F. grandis in the western Gulf of Mexico and on the
Florida east coast; and (6) differentiation of F. grandissimus in Yucatan.

THE FUNDULUS MAJALIS SPECIES COMPLEX
Fundulus majalis (WALBAUM) AND THE NOMINAL
Fundulus similis BAIRD AND GIRARD
FIGURES 9-12


FIGURE 9. Fundulus mafalis (JU 805), St. Johns County, Florida. Upper: male 61 mm SL;
Lower: female 77 mm SL. This is the typical "majalis" pigmentary pattern.
Cobitis majalis Walbaum 1792:12 (original description; Long Island, New York; types not
examined); Jordan and Evermann 1896:639 (placed in Fundulus).
Esox flavulus Mitchill 1815:439 (original description, New York); Jordan and Evermann
1896:639 (synonymy).
Esox zonatus Mitchill 1815:440 (original description, New York); Jordan and Evermann
1896:639 (synonymy).









BULLETIN FLORIDA STATE MUSEUM


FIGURE 10. Fundulus majalis (JU 995), Brevard County, Florida. Upper: male 87 mm SL;
Lower: female 75 mm SL. This represents the classical "similis" pigmentary pattern.


,e<>^ "~


FIGURE 11. Fundulus maialis (intergrade pattern) (JU 768), Flagler County, Florida. Upper:
male 66 mm SL; Middle: female 71 mm SL; Lower: female 88 mm SL.


Vol. 29, No. 1








RELYEA: GENUS FUNDULUS


FIGURE 12. Fundulus majalis (JU 693), St. Johns County, Florida. Juveniles (from top to
bottom) 47, 51, 53, and 57 mm SL.

Hydrargyra trifasciata Storer 1837:417 (original description, Massachusetts); Jordan and
Evermann 1896:639 (synonymy).
Hydrargyra majalis Cuvier and Valenciennes 1846:207 (South Carolina); Uhler and Lugger
1876:128 (Maryland); Yarrow 1877:214 (North Carolina); Goode 1879:118 (St. Johns R.,
Florida); Jordan and Evermann 1896:639 (synonymy).
Hydrargyra vernalis Cuvier and Valenciennes 1846:206 (original description, South Carolina);
Jordan and Evermann 1896:639 (synonymy).
Hydrargyra similis Baird and Girard 1853:389 (original description, Indianola, Texas; types
not examined); Jordan and Everman 1896:638-639 (synonymy).
Fundulus similis Giinther 1866:323 (removed F. similis from genus Hydrargyra and placed it
in Fundulus).
Fundulus majalis Giinther 1866:322 (removed F. majalis from genus Hydrargyra and placed it
in genus Fundulus); Garman 1895:104-105 (description; distribution); Jordan and Ever-
mann 1896:639 (description; distribution; synonymy); Miller 1955:9 (distribution); Brown
1957:72-73 (identified in key; distribution).
MATERIAL EXAMINED.-The number of specimens follows catalogue number in paren-
theses. NEW HAMPSHIRE: UMMZ 145275(1). MASSACHUSETTS: ANSP 7050-51(2),
7118(10); UMMZ 158794(50). NEW YORK: UMMZ 103799(50), 104029(50). NEW JERSEY:
ANSP 7102-17(16), 88226(2), 30987(1); UMMZ 114372(50), 109847(50). MARYLAND: ANSP
45631(2); TU uncat. (15); UMMZ 97817(40), 66865(50). VIRGINIA: UMMZ 99400(14),
99399(13), 99398(2), 102347(1), 171291(3). NORTH CAROLINA: ANSP 80733(18); TU
16510(11); UMMZ 132805(14); USNM 58975(3), 58977(3), 51854(4). SOUTH CAROLINA:
UMMZ 136059(58). GEORGIA: ANSP 80735(4), 80736(16); TU 21235(101), 21351(1); UMMZ
88584(3). FLORIDA: ANSP 89772(1), 74884(10), 30613-16(4); FSU uncat. (15), uncat. (4),
1579(17), 6518(12), 12418(30), 6526(69), 8892(3), 10385(1), 4639(6); JU 46,74,88,114,708,697,








BULLETIN FLORIDA STATE MUSEUM


733,685,693,679,552,474,497,504,165,147,791,643,787,776,768,805,795,793,860,841,840,
892, 877, 906, 1285, 1337, 1468, 1571, 1617; TU 20474(6), 5902(118), 37444(203), 20596(122),
18165(5), 18207(11), 20385(212), 41204(310), 41591(99), 2274(59), 20337(42), 21260(26),
41063(140), 40571(3), 27786(47), 9745(18), 18306(5), 18413(258), 20952(67), 20578(11),
18357(106), 20425(8), 18223(1), 41080(13), 41572(50), 41563(63), 20303(61), 16703(16),
41582(20), 41847(1), 20992(51), Uncat. (KR-Fla.-34) (13), 36160(2); UF 4011(2), 6611(30),
5227(28), 5201(1), 2987(2), 3303(3), 3117(5), 3298(79), 3137(2); UMIM 2603(12), 5314(2),
3151(2); UMML 3321(1); UMMZ 55989(3), 65242(2), 139341(33), 177472(53), 139390(1),
136587(2), 138388(96), 139369(77); USNM 18052(2), 104955(2), 62636(2). ALABAMA: TU
19837(36), 14519(1). MISSISSIPPI: TU 10873(3), 28139(11), 9858(1). LOUISIANA: TU
19508(19), 5881(85), 23209(26), 23635(259), 18760(1), 19162(30), 15337(148), 8267(26). TEXAS:
JU 1640, 1649, 1654, 1667, 1674, 1687; TU 12864(1), 1831(9), 39874(1), 21952(1), 21551(390),
41101(8). MEXICO: Tamaulipas: UMMZ 157322(3), 35 mi. N of Tampico.

DIAGNOSIs.-Members of the Fundulus majalis-F. similis species com-
plex may be distinguished from all other members of the genus by the
scaled preorbital region. Other diagnostic features include: dorsal rays
11-16, (usually 12-15), anal rays 9-13 (usually 10-12), lateral scales 30-37
(usually 33-35), gill rakers 6-10 (usually 7-8), pectoral rays 16-21 (usually
18-19), mandibular pores 8 (occasionally 7), branchiostegal rays 6, and
vertebrae 31-38 (usually 34-36).
DEscRIPTION.-In addition to the meristic features presented in the
Diagnosis and in Tables 8-12, there are 6 rays in each pelvic fin, 13-20
(usually 15-18) branched caudal fin rays, 13-17 (usually 14-15) predorsal
scales, and 15-22 caudal peduncle circumferential scales (usually 20-21
north of Cape Lookout and 17-19 southward).
There are 4 preopercular and 3 preorbital scale rows. Opercular scales
are irregularly arranged. There is an enlarged interorbital scale surrounded
by a rosette of 7 smaller scales. Contrary to Cooke's (1965) statement, the
interorbital scale arrangement will not distinguish this species from F.
heteroclitus (or F. grandis). Preorbital scalation extends dorsally beyond
the second preorbital pore, except in extreme south Florida populations.
This latter exception seems to be related to the elongation of the head
southward in this species complex (see morphometrics). Curiously, the
area immediately dorsal to the fourth (most ventral) preorbital pore,
approximately the size of one preorbital scale, is naked.
Of 183 specimens examined, 181 had 14 preopercular pores, and 2
specimens had 15 pores. Of 320 specimens examined, 251 had a total of 8
mandibular pores, 13 had 6, 53 had 7, and 3 had 9. There are typically 4
preorbital pores. Cephalic sensory pore arrangement was described in
detail for this species by Gosline (1949), and does not differ markedly from
the arrangement for the F. heteroclitus-F. grandis species complex.
The upper pharyngeal plates are separated into anterior and posterior
regions by a distinct groove and are about as wide as long. The anterior
portion is large (nearly as large as the posterior portion), but in the F.


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


heteroclitus-F. grandis species complex the anterior portion is small. Pha-
ryngeal plate structure is the object of continuing research by the author.
On the anterior portion of the upper plate are two vaguely defined trans-
verse rows of teeth, and 4 or 5 transverse rows on the posterior portion.
Pharyngeal teeth are small, numerous, and conical. The lower pharyngeal
plates are fused to form a broad triangular plate. The overall morphologi-
cal structure of the pharyngeal plates of F. majalis-F. similis resembles that
of F. diaphanus and F. seminolis.
Meristic characters and body markings vary considerably geographi-
cally. The dorsal fin ocellus of males is prominent in northern populations,
but is reduced in intensity or lacking in southern populations. Body mark-
ings of females, which range from vertical bars (F. similis type) to longitud-
inal stripes (F. majalis), are discussed in another section. Juvenile pigmen-
tation is like that of females (see Figs. 9-12).
Males in breeding color are a dark steel blue, especially dorsally. The
ventral region of the head and body are bright yellow anteriorly, shading to
white posteriorly, and an intense dark blotch occurs in the middle of the
operculum. The tip of the snout to the occiput is green. The median fins are
dark, and the caudal fin has a dark margin. The pectoral and pelvic fins are
bright yellow. An orange predorsal stripe extends from the dorsal fin origin
to the interorbital region (this stripe may be absent in extreme south Florida
specimens). Contact organs occur on the head, dorsal, and anal fins and on
body scales. In northern populations, 7 or 8 contact organs appear on the
posterior rim of each body scale and 4-6 in southern populations.
Members of the Fundulus majalis-F. similis population complex are
large species, attaining a maximum standard length of about 130 mm.
RANGE.-Fundulus majalis ranges continuously along the Atlantic coast
from New Hampshire, Great Bay, mouth of Lamprey River (Jackson
1953:192), southward to Lake Worth, Palm Beach County, Florida (JU
1337), and disjunctly in the Gulf of Mexico from Collier County, south-
western Florida (UMIM 3151, FSU 1579) continuously northward and
westward along the coast to near Tampico, Mexico (Miller 1955:9) (Fig.
13). An allopatric population occurs in the Florida Keys. Within this overall
range, the nominal F. majalis occurs, according to Miller (1955), from New
Hampshire southward to the Matanzas River, and the nominal F. similis
from the Matanzas River southward through the Florida Keys and west-
ward in the Gulf of Mexico to Mexico. I find (in several JU collections) the
"majalis" pigmentary features farther south than the Matanzas River, to
Flagler County. The "similis" form occurs southward from there. In all
Tables in this paper, therefore, the designation Flagler County includes
specimens that are "maialis" in appearance.
MERISTIC VARIATION.-Along the Atlantic coast the modal number of
dorsal rays (Table 8) is 14 in northern populations and 13 in southern








BULLETIN FLORIDA STATE MUSEUM


\ I . :r


Johns R.
atanzas- ._V -
L Flagler area \


Type localities:
S /A-i @ Fundulus majalis
a" OFE similis


FIGURE 13. Map of the geographic distribution of Fundulus maialis. Mexican localities in the
Tampico area actually farther south than indicated.

populations, the shift occurring between the Matanzas River and Flagler
County populations (both "majalis" types). The means for these two popu-
lations are statistically different, and the trend is clearly clinal. In the
Florida Keys the mode is 12, and the mean is significantly different from
populations adjacent in the Gulf and on the Atlantic coast. Dorsal rays show
a slight increase from north to south in the Gulf of Mexico. Of note is that
87% of the specimens examined from Cape Lookout to the Matanzas River
had 14 or fewer dorsal rays, while 98% from Chesapeake Bay northward
had 14 or more. A strong north-south dine is indicated, including gradation
from "majalis" to "similis." It is not altogether clear whether low counts in
Florida Keys population represent a southerly terminus of the dine or a
separate distinct population.
The modal number of anal rays is 11 in all populations, and any statisti-
cal differences between means is marginal.
The mean number of branched caudal rays (Table 9) for the lower
Florida east coast population (15.87), and for the Flagler County popula-
tion (16.20) differs significantly from the mean for the Matanzas River


Transas

Tamd~ico


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RELYEA: GENUS FUNDULUS


population (16.99). A clinal pattern is evident. The mode for caudal rays
shifts from 16 west of the Mississippi River to 15 eastward and back to 16 on
the Florida east coast from Mosquito Lagoon southward.
The modal number of lateral scales (Table 10) is 34 in all populations
from Cape Lookout southward on the Atlantic coast and throughout the
Gulf of Mexico, but is 32 in the Florida Keys. Populations from Chesapeake
Bay northward show an increase in the number of lateral scales, but means
are not significantly different from those of southern populations. The
mean for the Florida Keys population is significantly lower than means for
adjacent populations.
Predorsal scales do not differ markedly between populations, except
that the mean for predorsal scales in the lower Florida east coast race
(15.24) differs significantly from the Flagler County population mean
(14.62). The modal number of predorsal scales for the Florida Keys popula-
tion does not distinguish this population from Gulf of Mexico populations
(nor does the mean), but the Florida Keys mean (14.76) is statistically
different from the mean (15.24) for the lower Florida east coast population.
The modal number of gill rakers is 7 in all populations except those west
of the Mississippi River (mode = 8). The mean for this latter population is
also significantly different from populations in the Gulf east of the Missis-
sippi River.
Populations north and south of Cape Lookout are sharply delineated by
the number of caudal peduncle circumferential scales (Table 11). The
Chesapeake Bay mean (19.91) is significantly different from the Cape
Lookout-Charleston Bay mean (18.80), but there seems to be a general
clinal decrease southward along the Atlantic coast. In the Gulf of Mexico
the number of circumferential scales increases west of the Mississippi River
(mean is significantly different). Lower Florida east coast populations
resemble the Matanzas River population more closely than they do Gulf of
Mexico populations, which would either indicate gene flow from F. majalis
to the nominal F. similis or convergence in the Atlantic coast environment.
The mean for Florida Keys populations differs significantly from the mean
for Florida east coast populations, but not from the mean for the southward
Florida population in the Gulf.
The modal number of pectoral fin rays is 19 in all populations, except
for the Florida Keys and lower Florida east coast populations which have
18. The means (18.16 and 18.39, respectively) for these latter populations
are marginally statistically different from the Flagler County population
mean (18.73). Flagler County specimens are close, with regard to this
character, to Matanzas River specimens (mean = 18.79). The Florida
Keys' population mean (18.160 differs significantly from the mean (18.85)
for the southwest Florida population, but not from the Florida lower east
coast mean (18.39).







BULLETIN FLORIDA STATE MUSEUM


The number of vertebrae (Table 12) shows little variation throughout
the population complex, except for the Florida Keys. No statistical differ-
ence exists between the Matanzas River, Flagler County, and lower Florida
east coast populations with this character, but the mean for the Florida
Keys population is markedly different from adjacent population means.
The mode is 36 from Chesapeake Bay northward, 35 southward on the
Atlantic coast, 34 or 35 in Gulf populations and 33 in the Florida Keys.
MORPHOMETRIC VARIATION.-Little noteworthy variation occurs in
body proportions throughout the population complex, but head length
data (Table 13) are presented, as Brown (1957:72-73) suggested that this
character will serve to distinguish between F. majalis and F. similis, which
my data do not support. Southern populations on the Atlantic coast (both
males and females) have longer heads than northern populations, but this
character shows broad overlap in range. Gradation from F. majalis to F.
similis seems evident. Florida Keys and Gulf of Mexico specimens (espe-
cially in southwest Florida) have the longest heads. Preliminary data not
presented here indicate that elongation of pharyngeal plates accompanies
this southward increase in head length.
BODY MARKINCS.-The body markings of males (Figures 9-10; also see
excellent illustration in Garman 1895) do not vary greatly geographically.
Males typically have a well defined dorsal fin ocellus, which is formed by a
concentration of pigment on the last four dorsal rays and surrounded by a
cream-colored area. This mark is well defined southward along the Atlan-
tic coast to the Matanzas River. Farther south along the east coast of Florida
and in the Keys and Gulf of Mexico, the intensity of the ocellus decreases.
Miller (1955) did not note the ocellus in Fundulus persimilis from Yucatan.
In addition, the ocellus is usually lacking in the western Gulf of Mexico
population (from which F. persimilis was derived?).
The body markings of females show considerable geographic varia-
tion. Northern females (Fig. 9; also see Garman 1895) have two or three
longitudinal stripes, which begin just behind the pectoral fin and extend
well onto the caudal peduncle. One to four vertical bars appear on the
caudal peduncle. The stripes all lie on the ventral half of the sides. Juveniles
have from 7 to 14 vertical bars. The barred juvenile pattern breaks up by
the flattening of the dorsal region of each bar to form a "T." This process
begins anteriorly and proceeds posteriorly, leaving the last one or two bars
on the caudal peduncle unaffected (Newman 1907). The next step in the
process is the formation of a short longitudinal slash about midway on each
vertical bar. Occasionally a third slash appears farther ventrally (Fig. 12).
As this process continues, the longitudinal slashes coalesce to form
stripes, and the bars are completely disrupted. In northern populations this
process is usually completed by the time an individual attains 20 mm
standard length, whereas in southern populations the process is not com-


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RELYEA: GENUS FUNDULUS


plete until about 40-50 mm SL.
On the lower east coast of Florida, in the Gulf of Mexico, and in the
Florida Keys, the vertical bars are retained throughout the life of the
individual (Fig. 10). Adult female specimens from Flagler County, Florida
(between Matanzas River and Mosquito Lagoon), have a pattern in which
several bars are broken up into spots (Fig. 11). This north-south trend
enhances the idea that F. majalis and F. similis are conspecific.
Several females from the Chandeleur Islands in Mississippi Sound have
a distinct tendency toward the development of longitudinal stripes. In this
same collection (TU 8267) are several males with a well-defined ocellus in
the dorsal fin, showing that these body markings may appear in various
localities. In addition, I have examined adult F. majalis (up to about 70 mm
SL) from New York and South Carolina that have not completely deve-
loped the typical stripes. Juveniles from South Carolina, Georgia, Florida,
and throughout the Gulf of Mexico are indistinguishable, whereas northern
F. majalis juveniles generally show the adult female striped pattern early.
This probably accounts for the erroneous records of F. similis from South
Carolina by True (1883:255) and Jordan and Gilbert (1883:585). I believe
that it may also account for the record ofF. similis from the Matanzas River
by Miller (1955).
Water temperature may be a factor involved in the ontogenetic pig-
mentary changes in this species and should be more thoroughly investi-
gated. Specimens hatched at different times of the year may develop the
characteristic adult pattern at different rates. The single presumed F.
similis from the Matanzas River (UMMZ 139390; a 73 mm female) was
collected on 19 August 1936. Another female (74 mm SL; JU 88) collected in
September near St. Augustine has "similis-like" bars, but two of the bars on
one side are flattened dorsally to form a short stripe. These fish would be
about one year old. By contrast, numerous 30-40 mm juveniles that I
collected in the Matanzas River in August, which are presumably from a
spring spawning of the same year, had nearly completed the pigmentary
changes. Specimens about 30-40 mm SL collected in the same area in
March (presumably from a fall spawning) show only the very beginnings
of pigmentary changes and perhaps would not attain the adult pattern until
the next fall. Such specimens collected in the spring or early summer would
still have vertical bars and would attain adult markings at a later age and
greater size than individuals from a spring spawn. The two "similis-like"
specimens from the Matanzas River have, in my opinion, retained the
juvenile barred pattern, and can be allocated to F. majalis.
Females of the southern races typically have from 8 to 11 dark vertical
bars on each side, but the range is from 7 to 14, the character being very
variable (Fig. 10). The last vertical bar on the caudal peduncle may be
reduced to one, or often two spots, but this feature is changeable on any one







BULLETIN FLORIDA STATE MUSEUM


fish in a matter of minutes (personal aquarium observation). The same is
probably true of the number of vertical bars, although there appears to be
some genetic fixation of this character. One specimen from Fort Walton,
Florida, had only five vertical bars on each side on the posterior half of the
body when taken alive. During several months in an aquarium it never
developed more than five bars on each side.
The vertical bars found on adult females from southern populations
appear to be a retained juvenile character. Presumably the populations in
the Gulf of Mexico and on the lower Florida east coast are derived from a
population that differentiated in isolation from aF. majalis stock during the
Pleistocene (see Zoogeography). One manifestation of this isolation was
apparently the retention of juvenile body markings.
From the previous accounts of geographic variation of meristic and
morphometric characters and body markings, it is evident that Fundulus
majalis and F. similis are conspecific. No character affords specific separa-
tion, and the supposed sympatry of the two forms, based on a single
"similis-like" specimen collected in the Matanzas River, is untenable.
Differences between Atlantic and Gulf of Mexico populations of Fun-
dulus majalis are obvious, and the lower Florida east coast populations vary
with respect to meristic characters in the direction of F. majalis from the
Matanzas River, and yet body markings of the former clearly resemble
those of Gulf of Mexico populations.
The Flagler County specimens are certainly transitional between
Matanzas River and Mosquito Lagoon specimens. Some taxonomists
would consider the Flagler County material to represent intergrades
between two subspecies, but the only characters that would afford slight
separation of such subspecies are dorsal rays and body markings. The clinal
nature of these characters has been discussed previously. Admittedly there
is a definite transition from one form to another along the east coast of
Florida, but this is merely sharp clinal gradation. I find no compelling
evidence for the awarding of subspecific designations, although other
investigators might.
Griffith (1974) has concurred with this position, and Chen's (1971)
comparative chromosome analysis demonstrated no differences between
Fundulus majalis and F. similis. These forms should be considered
conspecific.
Florida Keys populations are allopatric (or appear to be), much like
Fundulus g. saguanus. In some respects these Keys specimens resemble
Atlantic coast populations, and in others Gulf populations. In other respects
(i.e. vertebral number), they differ markedly from other populations. Any
possible gene flow between the populations in the Florida Keys and adja-
cent populations remains to be demonstrated. In some ways (lateral scales)
the Keys' population seems to be the southern end of a dine. My original


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RELYEA: GENUS FUNDULUS


inclination (Relyea 1967) was to describe the Florida Keys population as a
distinct species equivalent to F. g. saguanus. This may yet prove to be the
correct interpretation, but I think that the characters exemplified in the
Keys may be expected in southernmost populations. Yucatan populations
of both species complexes being considered do not show the low meristic
counts expected of southernmost populations (some meristic features
increase from north to south in F. grandis).
This all may be evidence of, as I suspect, a lack of close genetic affinity
of Florida Keys and Yucatan populations, or it could reflect different
developmental environments. One might also be suspicious of the influ-
ence of genetic drift in an isolated small population such as in Yucatan.
Fundulus majalis (and F. g. saguanus) exists throughout the Florida Keys,
however, and genetic drift should probably not play a major part in
determining the observed frequencies of the characters studied; natural
selection and environmental influences, perhaps temperature, on devel-
opment must be assumed to be more significant. The structure of popula-
tions in the Florida Keys and Yucatan is different. In addition, the possibil-
ity of recent or continuing gene flow to the Florida Keys population from
peninsular Florida populations must be considered possible; such gene
flow to Yucatan populations would probably not have been possible for a
long time. Yucatan and Florida Keys populations should not be expected to
vary in the same way from other populations. Perhaps it is most significant
that Yucatan F. grandissimus and F. persimilis differ from other popula-
tions in similar ways, and so do Keys populations of F. grandis and F.
majalis, and yet Keys and Yucatan populations represent different extremes
of morphology in their respective species complexes.
It should also be noted here that the divergence of populations west of
the Mississippi River is not sufficient, in my opinion, to warrant recognition
of eastern and western Gulf of Mexico subspecies.
If subspecies are to be recognized in this population complex, then Gulf
of Mexico, Florida Keys, and lower Florida east coast (Mosquito Lagoon
southward) populations should be considered F. majalis similis; popula-
tions from the Matanzas River northward should be F. m. majalis; and the
Flagler County material should be intergrades.

Fundulus persimilis MILLER
Fundulus persimilis Miller 1955:13-25 (original description, Rio Lagartos, Yucatan, Mexico;
holotype UMMZ 163094, male 75 mm SL; paratypes UMMZ 163095 [30 spec.], UMMZ
162303 [2 spec.]; Brown 1957:69 (Yucatan).

DIAGNOSIS.-A species of Fundulus most closely related to F. majalis
("similis-type"), from which it may be distinguished by the smaller dorsal
and anal fins, the numbers of dorsal fin rays (9-11, usually 10), anal-fin rays







BULLETIN FLORIDA STATE MUSEUM


(9-10, usually 10), vertebrae (35-37, usually 36), and lateral scales (34-37,
usually 35-36) (see Miller 1955:13, Tables I, II, and III).
RANGE.-This species is known only from the Yucatan Peninsula of
Mexico.
DIscussIoN.-Fundulus persimilis has fewer dorsal and anal fin rays and
more vertebrae and lateral scales than F. majalis. These features and the
marked disjunction in its distribution relative to F. majalis leave little doubt
as to its specific validity. It is interesting to note that F. persimilis does not
resemble F. majalis from the Florida Keys, and as with the F. heteroclitus-
F. grandis species complex, the Yucatan population and Florida Keys
population seem to represent opposite morphological extremes.

ECOLOGY
Members of the Fundulus majalis population complex are euryhaline
and inhabit estauries, salt marshes and lagoons. Specimens have been
collected from a salinity range of 3.20/oo (Springer and Woodburn 1960) to
76.10/oo(Simpson and Gunter 1956). Although F. majalis congregates
around the mouths of creeks (Hildebrand and Schroeder 1928; personal
observation), it apparently does not swim upstream much beyond tidal
level. Gunter (1956) included this species in his list of euryhaline fishes, but
it is less euryhaline than either F. heteroclitus or F. grandis (see Griffith 1974
for extensive data).
There is a definite preference for a soft-mud, or sand-mud, substrate,
although F. majalis is extremely common along sandy shore lines (Springer
and Woodburn 1960:27; Hildebrand and Schroeder 1928:141; Bigelow and
Schroder 1953:165). My field observations indicate that the limiting factors
of dispersal are the degree of surf action in a given area and the availability
of shallow water zones. On the east coast of Florida the species is encoun-
tered in tidal marshes and estuaries off the major inlets. At Destin Inlet, in
the northern Gulf, the species is encountered in the inlet along shallow,
sandy shore margins, but not near the mouth of the inlet where the wave
action is intense. In Franklin County, Florida, F. majalis is common along
the sand-mud shorelines of the area, where wave action is minimal because
of the several offshore barrier islands and Alligator Peninsula to the east.
When strong south winds prevail, often in conjunction with an incoming
tide, the fishes retire to protected coves and creek outlets. On Dog Island
(Franklin County), the fishes are abundant on the bay side where surf
action is slight, but they are absent on the Gulf side. The same is true on
Alligator Peninsula. I have seen specimens in tide pools and low surf near
the mouth of the St. Johns River, Duval County, Florida.
Inlets on the east coast of Florida are often quite deep, and present only
a narrow shallow water zone or none at all. Fundulus majalis is rarely
encountered at depths of more than 2 feet, and consequently this species is


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RELYEA: GENUS FUNDULUS


restricted to limited areas on the Atlantic coast of Florida. This results in
small isolated populations wherever suitable conditions exist, severely
limits dispersal, and may, in part, explain the marked transitions from one
population to the next on Florida's Atlantic coast.
The Gulf coast presents nearly continuous shallow water zones, with
little wave action and extensive tidal marshes. Consequently, the species is
continuously distributed, shows less variation, and is abundant in the Gulf.
Numerous mangrove islands and mud flats provide the necessary habitat in
the Florida Keys.
Fundulus majalis forms larger schools (up to about 20 individuals) than
most species in the genus, with the exception ofF. seminolis. Most ecologi-
cal studies including these forms have noted school formation.
Joseph and Yerger (1956) and Kilby (1955) stated that this species (as F.
similis) spawns from spring through fall. Burger (1939) and Matthews
(1938, 1939) showed that water temperature, not photoperiod, stimulates
gonadal development in F. heteroclitus. Harrington (1959) demonstrated
the same for F. confluentus, and this probably prevails in F. majalis. The
length of the spawning period consequently decreases northward. Fundu-
lus majalis spawns in Chesapeake Bay from April to September (Hilde-
brand and Schroeder 1928) and in the Gulf of Maine in late spring and
summer (Bigelow and Schroeder 1953).
Spawning occurs in shallow cup-like depressions in shallow water
(Simpson and Gunter 1956; Martin and Finucane 1969). The large demersal
eggs are forced into the sand or mud during oviposition. The failure of
artificial hybrids (between female F. majalis and male F. heteroclitus and
F. grandis) to survive has been attributed to the large size of the egg (2 mm
diameter), and to the inability of the hybrid embryo to absorb all of the
yolk (Hubbs and Drewery 1959).
Newman (1909) stated that F. majalis spawns in small groups of several
males and females which my field observations support. In general pro-
tected waters with some debris or silt on the bottom are preferred,
although spawning also takes place along shallow, sandy shorelines. The
spawners may eat their own eggs. This is apparently related to general
feeding behavior, as schools ofF. majalis constantly poke into the substrate
for food. Individuals will also burrow into the substrate when alarmed.
Fundulus seminolis exhibits a similar spawning behavior.
Small crustaceans and molluscs are primary dietary items. Numerous
X-ray radiographs show that most individuals ingest great quantities of
sand while feeding. Hildebrand and Schroeder (1928:141) and Bigelow and
Schroeder (1953:163) stated that small fish are eaten. I have found no fish
remains in the alimentary tracts of the many specimens I have examined,
nor have I any evidence that F. majalis eats other fishes. Specimens that I
have kept in aquaria refused to eat small Gambusia affinis.







BULLETIN FLORIDA STATE MUSEUM


COMPARISON OF THE SPECIES COMPLEXES
The similar distribution of the Fundulus heteroclitus-F. grandis and F.
majalis species complexes affords a unique opportunity of comparing
geographic variation and population differentiation between them. A
point of interest is the fact that in both species complexes a distinct species
occurs on the Yucatan Peninsula of Mexico, separated by a distance of
several hundred miles of coastline from related Gulf of Mexico forms.
The pattern of geographic variation within F. grandis and southern
populations of F. majalis are strikingly similar: (1) racial differentiation east
and west of the Mississippi River; (2) isolation on the east coast of Florida;
and (3) allopatry in the Florida Keys. A major notable difference is that F.
grandis had dispersed to Cuba and the Tortugas, whereas F. majalis has
not.
Fundulus grandis does not hybridize with its northern cogener, F.
heteroclitus, while a transition between races of F. majalis is apparent. The
area of sympatry of F. grandis and F. heteroclitus is about the same as the
area of transition between races of F. majalis.
Fundulus majalis and F. heteroclitus both show racial differences north
and south of Cape Lookout, North Carolina. Fundulus heteroclitus has
dispersed farther northward (to Canada) on the Atlantic coast than F.
majalis, and has populated Bermuda (perhaps several times).
The Fundulus heteroclitus-F. grandis complex has been more success-
ful in reaching more isolated, or harsher, regions than F. majalis. This
probably reflects both dispersal ability and ability to exploit new regions
once they are attained. Williams (1960) showed that F. heteroclitus does not
move very much from a given place, even across tidal creeks with moder-
ate tidal currents. Nevertheless, this species has dispersed to Bermuda, and
F. grandis to Cuba and the Tortugas.
Racial differentiation in F. majalis north and south of Cape Lookout
would be the direct influence of temperature, with more northern popula-
tions exhibiting higher counts for meristic features. This is not obvious in F.
heteroclitus, as the northern race tends to have lower meristic features.
Gabriel (1944) demonstrated that F. heteroclitus reared in colder waters
tended to have more vertebrae than those raised in warmer waters. Fahy
(1972) came to the same conclusion for F. majalis. Gabriel (1944) also
demonstrated genetic control of vertebral number, but other characters
should not necessarily respond to water temperature during developmen-
tal stages in the same way, or at all, as do vertebrae, and for pectoral rays,
gill rakers, and caudal peduncle circumferential scales, the northern race
has fewer than the southern race. The length of the anal sheath of females
also increases southward, but how this could be influenced by water
temperature is unclear. Moreover along the Gulf coast these same charac-
ters increase from north to south in F. grandis.


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RELYEA: GENUS FUNDULUS


Racial differentiation in F. heteroclitus and F. majalis is surely a combi-
nation of genetic and environmental influence. Populations of these forms
are continuous along the Atlantic coast, and there is no valid basis for
recognizing the geographic races nomenclaturally.
Racial differentiation in F. majalis and F. grandis in the Gulf of Mexico
is not marked. The strong outflow of freshwater from the Mississippi River
probably serves to keep eastern and western Gulf races at least partially
separated, showing a parallel to William's (1960) work on a magnified
scale.
Florida east coast populations are geographically isolated from Gulf
populations, and gene flow from Atlantic to Gulf is nonexistent. This results
in disjunct populations of F. grandis and F. majalis, with an intervening,
closely related form in the Florida Keys. That the differences between
Atlantic and Gulf races are in part genetic is plausible. As pointed out
previously, lower Florida east coast F.majalis show the genetic influence of
the Matanzas River population, and thus differ from Gulf populations in
that respect.

ZOOGEOGRAPHY
The present distributions ofF. heteroclitus and F. grandis reflect popu-
lation movements (in a long term sense) during and since the Wisconsin
glaciation. Walters and Robins (1961) pointed out that Pleistocene sea
levels may have been as much as 450 feet lower than present sea levels, and
Emiliani (1958) demonstrated that Wisconsin glacial period sea tempera-
tures in Florida may have been as much as 6 C lower than present
temperatures. Such lowered temperatures would have forced the temper-
ate fauna southward into south Florida and the Caribbean. For a discussion
of this Trans-Florida faunal group see Walters and Robins (1961). A fine
discussion of Gulf-Atlantic disjunct distribution can also be found in Dahl-
berg (1970). Hedgepeth (1953) discussed disjunct invertebrate distribu-
tions. The reader is also referred to Miller (1966) and Myers (1966) for
discussions of Caribbean distribution.
As water temperatures have warmed since the last glaciation, popula-
tions have moved northward. Separation into temperate Atlantic and Gulf
of Mexico populations (disjunct Carolinian fauna) was inevitable for tem-
perate forms incapable of existing in tropical south Florida environmental
conditions.
Fundulus grandis would appear to be a Gulf derivative of an F. hetero-
clitus population (or a common precursor of both) isolated during some
earlier glacial-interglacial sequence. The mandibular pore number of 8 in
Fundulus heteroclitus is shared with most other Fundulus. The 10 mandibu-
lar pores of F. grandis is presumably a derived state. The current disjunct F.
g. grandis populations in the Gulf and Atlantic (Keys' populations [= F. g.







BULLETIN FLORIDA STATE MUSEUM


saguanus] considered later) reflect the most recent northward movement
on each side of the Florida peninsula of southerly populations. This disjunc-
tion could be a repetition of the mechanism that earlier isolated F. grandis
and F. heteroclitus. The situation with F. majalis is similar, except that
reproductive isolation had not been achieved between the early Gulf and
Atlantic disjuncts, so that intergradation occurs along the northeastern
Florida coast in the region of contact between Gulf and Atlantic forms.
In this regard, the distributions of several fishes are worth noting:
1) Menidia beryllina, M. peninsula, and M. menidia present a strikingly
similar pattern of distribution to the Fundulus considered here. Menidia
beryllina and M. peninsula appear to have disjunct Gulf and Atlantic
populations (same gaps in southeastern Florida and southwestern Florida
as for Fundulus, although the exact distribution in south Florida and the
northern Florida Keys is as yet unclear). Isolated populations also occur in
Yucatan (M. colei and Menidia species) and the Florida Keys (M. concho-
rum; considered later). Menidia menidia is an Atlantic coast form that
ranges from Canada to northeastern Florida (range nearly identical to F.
heteroclitus). Menidia beryllina occurs throughout the Gulf of Mexico and
ranges northward to Massachusetts on the Atlantic coast. Menidia beryllina
(or M. peninsula) and M. menidia may hybridize in the Matanzas River in
northeastern Florida (Gosline 1948). The reader is referred to Gosline
(1948), Robbins (1969), Johnson (1975) and Duggins (1980) for further
discussion of Menidia systematics.
2) The zone of overlap between Chasmodes bosquianus (disjunct Gulf
and Atlantic populations) and C. saburrae (intervening peninsular Florida
form) is near Marineland in northeastern Florida, the area of contact
between Gulf and Atlantic populations of Menidia and Fundulus discussed
above (Springer 1959; Williams 1983).
3) The region of sympatry of the temperate Atlantic puffer Sphoeroides
maculatus and the southern puffer, S. nephelus (essentially a Gulf form) is
in the same northeastern Florida area noted above (Shipp and Yerger
1969).
4) Floridichthys carpio ranges from Cape San Bias, near Port St. Joe,
Florida (Kaill 1967), southward along Florida's Gulf coast through the
Florida Keys and northward on the Atlantic coast to Lake Worth, Palm
Beach County, Florida (JU 1338). It then occurs disjunctly(?) on the Atlan-
tic coast from the waters around Fort Pierce northward to the southern end
on Mosquito Lagoon, Volusia County, Florida, just to the north of the Cape
Canaveral area (JU 1298) (the same northeast Florida area noted above).
This presently known distribution suggests separation of the two popula-
tions in South Florida. Two subspecies of F. carpio are recognized from
Yucatan (Hubbs 1936); but Duggins (1980) challenged that view.
Another related factor worth noting is that Floridichthys carpio is not
common north of the Cedar Key area of Florida's Gulf Coast (about 290 N


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RELYEA: GENUS FUNDULUS


Lat.). In addition, the disjunct northeastern Florida population is concen-
trated near the same latitude. This also marks the zone of transition
between previously mentioned species and is approximately the northern
distributional limit of mangroves. Parr (1933) emphasized a winter thermal
barrier to the dispersal of marine organisms along the Atlantic coast, and
pointed out the abrupt change of average winter temperatures of about 60"
F north of Cape Canaveral to 70 F to the south (due to the Gulf Stream).
This marked temperature change between the southern and temperate
parts of Florida (Carolinian Province) is not evident on the Gulf coast, and,
with the exception of Floridichthys carpio, is not reflected by fish
distributions.
The distributional limits of killifishes in the western Gulf of Mexico are
also near the northern limits of mangrove growth (northern limit is about
20N Lat., Tamaulipas, Mexico; Thorne 1954). Currently accepted south-
ern limits of some cyprinodontids in the western Gulf area: Fundulus
grandis, Laguna de Tamiahua, Veracruz, Mexico (Miller 1955); F. majalis,
near Tampico, Mexico, (Miller 1955); E. confluentus, Laguna Madre,
Texas (Breuer 1962); Adinia xenica, San Patricio County, Texas (Hastings
and Yerger 1971); Cyprinodon variegatus, into northeastern Mexico (Kaill
1967); Lucania parva, Tampico, Mexico (Hubbs and Miller 1965). These
distributions range slightly north or south of 250 N Lat., the latitude of the
Florida Keys in the eastern Gulf, and the southern limit in the east for all
species listed above (Fundulus grandis and F. majalis actually extend south
to about 220 N Lat.). Menidia beryllina and M. peninsula are similarly
distributed.
Allopatry in the Florida Keys is also demonstrated by Menidia concho-
rum, and may be equivalent to the Keys Fundulus populations. Syngnathus
floridae mckay; is a Florida Keys endemic isolated from its disjunct Gulf-
Atlantic relatives S. f. floridae (Cape Sable, Florida to Corpus Christi,
Texas) and S. f. hubbsi (Chesapeake Bay to South Carolina) (Herald 1965).
I believe Lucania parva in the Florida Keys may also be an allopatric
population. Hubbs and Miller (1965) did not examine specimens from
south of the St. Johns River on the Atlantic coast. I have specimens south-
ward to the Ft. Pierce area, but none are known between there and the
Florida Keys (Duggins 1976, 1980). The relationship of Cubanichthys
cubensis of Cuba to L. parva should be investigated as well (see Rosen
1975). Cyprinodon variegatus may be isolated in the Keys, too. Jordan
(1884) described Cyprinodon in the Florida Keys and Cuba as C. v.
riverendi. All of these Keys' forms probably constitute glacial relicts, in the
sense of Walters and Robins (1961). Other populations of Cyprinodon in
the Bahamas and West Indies (see Hubbs and Miller 1942) may likewise be
glacial relicts.
I believe that based on my data, Fundulus grandissimus and F. persimi-
lis and possibly Cyprinodon variegatus artifrons, Floridichthys popula-







BULLETIN FLORIDA STATE MUSEUM


tions, and Menidia colei represent glacial relicts (originating from western
Gulf of Mexico populations) that have become isolated in Yucatan. Flori-
dichthys carpio does not now occur in the western Gulf of Mexico, but may
have in the past and did not survive the last glaciation except as a relict in
Yucatan and South Florida. I do not believe that Florida Keys and Yucatan
populations of these are very closely related, nor that a direct, overwater
Keys-Yucatan dispersal has been involved. Rather dispersal on a north-
south basis in the western Gulf or following interglacial-glacial events has
occurred.
The reader is referred to a recent paper on this problem by Rosen
(1975). Rosen's vicariance (allopatric speciation) model of Caribbean zoo-
geography suggests a North American-Caribbean distributional track
(among others) that is the result of tectonic movement that fragmented an
ancestral biota. With respect to my study, the relationship of Cuban (Antil-
lean) and south Florida populations to Yucatan populations is at issue.
Rosen presented an excellent summary (1975: 457, Fig. 21) of cladistic
interpretation of vicariance versus dispersal for Caribbean distributions,
and convincing arguments. My data support Rosen's suggestion that the
distribution and cladistic relationships of populations in Fundulus heterocli-
tus-F. grandis species complex do not fit the vicariance model, and that
dispersal over long periods of time (Pleistocene glacial-interglacial events)
best accounts for observed distribution and relationships. These data do
not support Rosen's suggestion of overwater dispersal from Florida to
Yucatan. Cuban and Floridian populations are more closely related to one
another than either is to F. grandissimus of Yucatan. This suggests Florida-
Cuba dispersal, but not dispersal from Florida (or Cuba) to Yucatan. Other
cases of Cuba-Florida or Keys-peninsula relationships may be cited: Flori-
dichthys carpio, Yucatan and Florida, but not Cuba, with possibly disjunct
Gulf-Atlantic populations; Menidia beryllina (or M. peninsula) is proba-
bly more closely related to M. conchorum of the Florida Keys than either is
to M. colei of Yucatan (personal communication, Dr. Robert R. Miller);
Cyprinodon populations noted earlier; Fundulus majalis of the Florida
peninsula and the Keys are more closely related than either is to Yucatan F.
persimilis.
An intriguing "hybrid" pattern is suggested by L. parva. The seeming
isolation of Florida Keys populations from very closely related peninsular
populations, the close relationship of Gulf coast populations to Lucania
from Mexico (personal communication, Dr. Robert R. Miller), and the
isolation of the more distant sister species, Cubanichthys cubensis on Cuba,
suggest both dispersal and vicariance. Indeed, this may be the critical
point. Obviously, Pleistocene glacial-interglacial alternation occurred, as
did the Caribbean tectonic events Rosen described. Any theory used to
explain current distributions of American north temperate-subtropical


Vol. 29, No. 1







RELYEA: GENUS FUNDULUS


organisms must acknowledge both. Animals and plants exist in certain
ecological regimes that have probably moved with glacial movements.
These are not abrupt movements, but rather gradual shifts or ranges over
tens of thousands of years. These are themselves vicariant events (as
actually acknowledged by Rosen 1975:432). My collection data for Florida
east coast killifish suggest seasonal and yearly distributional shifts. Perhaps
some of the best examples of Pleistocene shifts involve isolation of subter-
ranean populations of animals, notably fishes and crayfishes in North
America, from epigean ancestral populations that were moved either
northward or southward, or brought to extinction by glacial-interglacial
events. Relyea (1976) discussed this situation for Florida cave crayfishes,
and suggested a similar origin as a result of isolation in the springs of Florida
during a glacial period, for the endemic killifish of the Florida peninsula:
Lucania goodei, sister species of L. parva; Fundulus seminolis, sister spe-
cies of F. majalis (?); Jordanella floridae, sister species of Floridichthys
carpio (is Garmanella of Yucatan a close relative of Jordanella, or is it a
species similar in appearance to Jordanella for ecological reasons, that has
been independently derived from Yucatan or western Gulf of Mexico
Floridichthys carpio?). Finally, the geminate species in the Gulf of Mexico
and along Florida's Atlantic coast argue for a dispersal theory as well.
A critical point for further study involves the large gap between north-
eastern Mexico and Yucatan in killifish and silverside distributions. Is this
the result, as I suggest, of north-south dispersal and extinction in the Gulf of
Mexico of a fragmented Antillean-Yucatan biota or a result of tectonic
events or of overwater dispersal from Florida to Yucatan?
The seeming lack of affinity between some greater Antillean and Yuca-
tan fishes, exemplified best by the killifishes noted above, would suggest
the former (dispersal) and is at odds with Rosen's North American-
Caribbean track, but does not invalidate any of his other evidence or
conclusions. My data, I believe, further the gradual elucidation of North
American-Caribbean fish distributional patterns, and additional studies of
complexes such as Floridichthys-Jordanella-Garmanella species group are
needed.









BULLETIN FLORIDA STATE MUSEUM


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Vol. 29, No. 1




TABLE 1.-Frequency distribution of lateral scales in the Fundulus heteroclitus-F. grandis species complex. The t value between asterisked


populations is 3.37. T

Population
F. heteroclitus
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

South of St. Johns River

F. grandis grandis
Florida east coast*

F. grandis saguanus
Florida Keys


he t value following the Florida Keys population is for comparison with Southwest Florida.

30 31 32 33 34 35 36 37 38 39 40 41 42 N X S t


11

3 13

1 5 16

2 3 24

3 22

1 2 6


40 66 36 9 1

42 46 13 2 1

35 32 4 2

49 33 7

50 61 25 5

22 16 1


3 20 59 42 18


4 12

4 4


Cuba


F. grandis grandis
Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River


29 45

2 1


1 3

1 13

2 10

1 3 12


18 1


38 46 14 2

83 95 23 4

57 61 46 10

60 95 25 11 2


34.97

34.53

34.18

34.09

34.59

34.11


142 34.37 .66


32.59

31.00


34.72

34.63

34.91

34.79


4.00

2.69

.64

3.85

3.69 m

2.36
Z
C1
17.12


9.26 r
C


.87

3.11

1.20

15.98


F. grandissimus
Yucatan


Yucatan 2 2844 7 2 29 39.21 1.64


2 2 8 4 4 7 2


29 39.21 1.64





TABLE 2.-Frequency distribute
populations is 3.73. T

Population
F.:heteroclitus
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

South of St. Johns River

F. grandis grandis
Florida east coast*

F. grandis saguanus
Florida Keys


Cuba


F. grandis grandis
Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River

F. grandissimus
Yucatan


)n of predorsal scales in the Fundulus heteroclitus-F. grandis species complex. The t value between asterisked
he t value following the Florida Keys population is for comparison with Southwest Florida.

13 14 15 16 17 18 19 20 21 22 23 24 25 N X S t


2

1 12

4 15

4 27

25

1 4


17 44 43

23 41 27

37 28 7

47 30 7

52 56 25

14 21 4


38 8 0 1

10 3 2

1


16.83

16.12

15.24

15.08

15.67

15.58


9 1

1


2 12 49 57 20 5


8 18

4


6

1 12

5 30

4 9


44 25 6

7


20 45 27

67 100 41

51 65 27

79 74 23


8 1

7 1

7 1

3 2


145 15.66 1.01


15.03

14.64


16.13

15.83

15.56

15.62


1.23

4.90

.56 q

.50


4.85


7.85 C


2.38

2.48

.60

38.52


3 5 13 8 2


31 23.03 1.10




TABLE 3.-Frequency distribution of gill rakers in the Fundulus heteroclitus-F. grandis species complex. The t value between asterisked
populations is 1.22. The t value following the Florida Keys population is for comparison with Southwest Florida.


Population
F. heteroclitus
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

South of St. Johns River

F. grandis grandis
Florida east coast*

F. grandis saguanus
Florida Keys

Cuba


7 8 9 10 11 12 13 14 15 16 17 N X S t


2 22 55 65 19 4

8 24 56 25 7

4 33 40 16 4

3 23 53 38 4

1 21 44 84 16 6

2 7 19 16 5 1


1 42 69 32 2


1 25 52 31

9 2


167 10.53 1.01

120 10.99 .97

97 10.82 .96

121 12.14 .85

0 1 173 12.67 1.01

50 12.36 1.04


146 9.95 .71


111 9.07 .81

11 9.18 --


F. grandis grandis
Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River

F. grandissimus
Yucatan


1 2 31 61 14

1 13 87 110 20

19 73 69 22

13 77 87 25


3 20 5 1 29 9.14 .61


5.05 $

1.85 t

15.06
z

9.77
Z
C
7.06 r
C/1


112 9.84

233 9.61

186 9.56

207 9.68


3 20 5 1


29 9.14 .61




TABLE 4.-Frequency distribution of vertebrae number in the Fundulus heteroclitus-F. grandis species complex. The t value between asterisked
populations is 2.30. The t value following the Florida Keys population is for comparison with Southwest Florida.

Population 31 32 33 34 35 36 37 38 N X S t


F. heteroclitus
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

South of St. Johns River

F. grandis grandis
Florida east coast*

F. grandis saguanus
Florida Keys

Cuba

F. grandis grandis
Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi
F. grandissimus
Yucatan


2 17 18 13

1 10 19 18

5 20 18 5

5 15 20 6

2 11 15 4 1

1 4 9 11 1


1 7 16 14


2 17 23

7 1


50 34.84

57 35.42

49 34.53

46 34.59

33 33.73

26 33.27


41 35.27 .85


8 1


1 4 14 2

1 16 29 12

1 7 25 18 5

7 14 6 1

1 6 1


51 33.78

9 32.00


21 34.81

58 34.89

56 35.34

28 35.04

8 37.00


.38

5.06

2.10 "
0
9.13


7.76 >
5. I


5.15 q
C


.67

1.00

.85

.57








RELYEA: GENUS FUNDULUS


TABLE 5.-Frequency distribution data for total mandibular pores in the Fundulus hetero-
clitus-F. grandis species complex. The t value between asterisked populations
is 44.72.


Species 6 7 8 9 10 11 12 N X S t

F. bermudae 30 30 8.00 -
F. rhizophorae 30 30 8.00 -
F. heteroclitus 2 1 100 103 7.95 .32
58.52
F. grandis grandis* 17 203 1 1 222 9.94 .20
F. grandis saguanus 20 20 10.00 -
F. grandissimus" 3 25 28 11.89 .32




TABLE 6.-Anal sheath length, expressed as thousands of standard length, and standard
length (in mm) of specimens, of females for the Fundulus heteroclitus-F. gran-
dis species complex.


Standard Length Anal Sheath Length
Population N Range Mean Range Mean

F. bermudae 10 53-71 62.8 112-155 140.0
F. rhizophorae 10 46-58 52.0 103-164 143.2
F. heteroclitus
North of Cape Cod 10 67-90 75.6 59-114 86.7
Long Island 10 56-84 66.9 107-145 125.1
Chesapeake Bay 10 58-79 67.6 155-200 173.5
Cape Lookout-Charleston area 10 54-83 63.6 180-233 200.1
South of Savannah River 10 55-89 76.0 166-194 178.8
F. grandis
Florida east coat 10 62-77 69.7 52-80 69.0
Florida Keys 10 59-82 70.4 52-99 79.4
Southwest Florida 10 59-81 67.4 34-80 53.9
Northern Gulf of Mexico 10 57-80 65.4 67-100 85.6
Western Gulf of Mexico 10 58-70 62.9 67-103 77.7
F. grandissimus 3 111-134 124.7 54-75 64.0













TABLE 7.-Summary of meristic data for Fundulus heteroclitus, F. bermudae, and F. rhizophorae. The t value that follows F. rhizophorae
compares the mean for that species and F. heteroclitus.


Dorsal Fin Rays 10 11 12 13 14 15 N X S t
F. heteroclitus 2 188 419 111 10 1 731 11.92 .71
19.20
F. bermudae 1 2 33 36 5 77 13.55 .73
22.78
F. rhizophorae 2 50 51 1 104 11.49 .57
.60
Anal Fin Rays 9 10 11 12 13 14 N X S t
F. heteroclitus 2 178 498 57 735 10.82 .72
17.80
F. bermudae 2 47 26 1 76 12.34 .60
17.91
F. rhizophorae 1 24 72 8 105 10.83 .53
.14
Branched Caudal Rays 13 14 15 16 17 18 19 20 21 N X S t


5 28 105 170

3 30

2 8 42 40 10


231 126 39 4 708 17.62 1.29

34 7 74 17.61 .67

3 105 15.54 .99


F. heteroclitus

F. bermudae


F. rhizophorae


.07

15.64

15.86










TABLE 7 Continued.

Lateral Scales 30 31 32 33 34 35 36 37 38 N X S t
F. heteroclitus 4 16 92 238 254 86 18 2 710 34.50 .92
4.93
F. bermudae 1 6 14 24 19 2 66 33.91 1.03
3.23
F. rhizophorae 1 3 14 35 37 12 1 103 33.40 .99
11.24
Predorsal Scales 11 12 13 14 15 16 17 18 19 20 21 N X S t
F. heteroclitus 10 85 190 220 113 59 12 2 1 692 15.84 1.28
9.99
F. bermudae 2 4 38 19 2 65 14.23 .76
3.05
F. rhizophorae 2 13 27 32 21 4 1 100 13.73 1.17
15.56
Gill Rakers 8 9 10 11 12 13 14 15 16 17 N X S t


2 34 118 212 176 153 25 7 0 1 728 11.55 1.28

27 34 9 1 71 12.77 .80

1 31 50 20 2 1 105 10.94 .87


z






cj


7.87

14.08

4.74


F. heteroclitus

F. bermudae

F. rhizophorae











TABLE 7 Continued.

Caudal Peduncle
Circumferential Scales 15 16 17 18 19 20 21 22 N X S t
F. heteroclitus 5 37 147 270 29 4 492 19.60 .67
7.09
F. bermudae 18 24 16 58 18.97 .65
18.29
F. rhizophorae 4 43 36 20 103 16.70 .81
38.41
Pectoral Fin Rays 15 16 17 18 19 20 21 N X S t
F. heteroclitus 7 91 307 240 50 2 697 18.34 .96
.79
F. bermudae 5 43 26 3 77 18.35 .68
21.75
F. rhizophorae 10 64 27 4 105 16.24 .62
21.65
Number of Vertebrae 31 32 33 34 35 36 37 38 N X S t
F. heteroclitus 1 6 33 88 80 43 10 261 34.57 1.02
5.62
F. bermudae 1 2 14 11 3 1 32 33.50 .98
3.87
F. rhizophorae 1 16 22 4 43 32.67 .87
11.54








RELYEA: GENUS FUNDULUS


TABLE 8.-Frequency distribution of dorsal fin rays in the Fundulus majalis population
complex. The t value between asterisked populations is .63

Population 11 12 13 14 15 16 N X S t
F. majalis ("maialis-type")
North of Cape Cod 1 12 15 2 30 14.60 .67
.44
Delaware River-Long Island 3 62 55 9 129 14.54 .71
2.33
Chesapeake Bay 1 8 53 38 3 103 14.33 .71
4.22
Cape Lookout-Charleston area 1 20 64 15 1 101 13.95 .66
3.00
Savannah River-St. Johns River 1 46 54 12 113 13.68 .70
1.56
Matanzas River, Florida 2 43 94 17 1 157 13.82 .85
4.95
Flagler County, Florida 8 48 29 2 87 13.28 .80
.09
F. majalis ("similis-type")
Cape Canaveral area, Florida,
and south* 11 40 24 4 79 13.27 .69
8.00
Florida Keys 8 56 52 7 123 12.47 .73
8.92
Southwest Florida* 23 90 45 6 164 13.21 .67
3.52
Tampa Bay-Apalachicola Bay 3 60 160 55 1 279 12.96 .82
.36
Cape San Bias-Mississippi River 58 162 40 1 261 12.94 .60
2.00
West of Mississippi River 1 27 91 34 1 154 13.05 .57








TABLE 9.-Frequency distribution of branched caudal fin rays in the Fundulus maialis population complex. The t value between asterisked
populations is 3.89.


Population
F. majalis ("majalis-type")
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

Matanzas River, Florida

Flagler County, Florida

F. majalis ("similis-type")
Cape Canaveral area, Florida,
and south*

Florida Keys

Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River


13 14 15 16 17 18 19 20 N X S t


5 17 8

7 18 78 18

1 2 26 40 21

7 42 36 9

1 33 43 31

1 2 36 64 36

8 55 21 2


1 16 43 9

5 43 64 8

6 71 69 7

20 124 109 21

1 19 154 74 12

5 61 69 15


30 17.10

125 16.96

98 17.05

95 16.53

110 17.00

142 16.99

86 16.20


69 15.87

121 15.65

153 15.50

277 15.51

260 15.30

150 15.63


1.00
C
.71

4.00

4.31 1

.09

7.90 -

3.30
C


2.32

1.88

.14
<
3.82

5.23
o







TABLE 10.-Frequency distribution of lateral scales in the Fundulus majalis population complex. The t value between asterisked populations is
2.50.


Population
F. majalis ("maialis-type")
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

Matanzas River, Florida

Flagler County, Florida


F. majalis ("similis-type")
Cape Canaveral area, Florida,
and south*

Florida Keys

Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River


30 31 32 33 34 35 36 37 N X S t


1 11

10

6 22

4 31

3 38

1 2 8


36 9 1


4 34

2


4

12

6 3

3


1 30 34.77

126 34.39

3 98 34.63

101 33.91

112 33.94

145 33.99

70 34.09


68 33.81

123 31.93

162 34.06

279 34.23

260 34.12

152 34.07


15.41

20.48

2.21

1.31

.53







TABLE 11.-Frequency distribution for caudal peduncle circumferential scales in the Fundulus maialis population complex. The t value between
asterisked populations is 4.35.

Population 15 16 17 18 19 20 21 22 N X S t


F. majalis ("maialis-type")
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

Matanzas River, Florida

Flagler County, Florida

F. majalis ("similis-type")
Cape Canaveral area, Florida,
and south*

Florida Keys

Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River


2 6

3

2 12

4




3 7

6 31

7 23

1 44 67

1 44 39

8 15


9

1 21

21 41

32 31

22 35

27 31




14 14

36 20

22 6

48 20

23 8

30 34


10 15 4

45 27 5

51 11 2

19 1

1

21

2


29 20.79

86 20.33

86 19.91

90 18.80

67 18.45

92 18.66

64 18.48




51 18.53

96 17.82

62 17.63

180 17.23

115 16.94

94 18.18


1.72 r

).41

!.97

.62

.22
.2
.26
K







TABLE 12.-Frequency distribution of number of vertebrae in the Fundulus majalis population complex. The t value between asterisked 2
populations is 2.67.

Population 32 33 34 35 36 37 38 39 N X S t


F. majalis ("maialis-type")
North of Cape Cod

Delaware River-Long Island

Chesapeake Bay

Cape Lookout-Charleston area

Savannah River-St. Johns River

Matanzas River, Florida

Flagler County, Florida

F. majalis ("similis-type")
Cape Canaveral area, Florida,
and south*

Florida Keys

Southwest Florida*

Tampa Bay-Apalachicola Bay

Cape San Bias-Mississippi River

West of Mississippi River


8 12 6 1

1 15 21 11

2 17 20 2

5 20 22 5 2

1 14 14 6

5 21 43 15

2 12 24 6 3


2 9 42 13

3 17 22 8 4

5 14 11 4

14 20 7

1 7 29 24 8

2 10 24 6 1


27 37.00

48 36.88

41 36.54

54 35.61

35 35.71

84 35.81

47 35.91




67 36.03

54 33.87

34 35.41

41 35.83

69 35.45

43 34.86


.71

3.78

.55

.48

.54

.56

.67 C
Z



13.42

7.10

2.13

2.57

3.60















TABLE 13.-Head length, expressed as thous




Population
F. majalis ("majalis-type")
North of Cape Code
Long Island
Chespeake Bay
Cape Lookout-Charleston area
Savannah River-Matanzas River
Flagler County, Florida
F. majalis ("similis-type")
Cape Canaveral area, Florida, and south
Florida Keys
Southwest Florida
Northern Gulf of Mexico
West of Mississippi River


sandths of standard length, and standard lengths (in mm) for Fundulus majalis. t

Females Males
Standard Length Head Length Standard Length Head Length
N Range Mean Range Mean N Range Mean Range Mean

10 72-84 78.0 316-333 323.4 10 62-70 66.7 323-339 330.0
10 51-85 68.3 294-333 314.5 10 56-84 72.8 291-323 307.2
10 55-87 68.6 301-341 316.4 10 63-85 72.5 286-350 304.0
10 62-94 80.7 315-350 334.0 10 60-95 74.4 316-343 329.4
15 57-83 68.6 302-368 332.4 15 53-75 61.1 293-363 328.5
10 58-69 66.0 319-348 337.2 10 53-67 59.0 316-343 332.6

15 60-97 72.5 297-358 330.2 15 51-66 58.7 306-339 321.1
10 64-87 69.6 343-380 358.5 10 55-100 63.7 320-369 342.0
10 61-84 74.6 322-385 346.3 10 53-78 65.6 345-368 353.9
10 59-72 63.4 308-346 323.2 10 55-70 61.2 301-333 323.2
10 63-73 66.1 301-333 322.4 10 53-67 60.5 303-333 319.1




o
z..









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