OCCASIONAL PAPERS ^^'- "^ ^^'^^

HARVARD
of the UNIVERSITY

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 66, PAGES 1-31 JUNE 24, 1977

THE PHYLOGENY AND SYSTEMATICS OF THE
FUNDULUS NOTTII SPECIES GROUP
( TELEOSTEI : CYPRINODONTIDAE )

By

E. O. Wiley'

The Fundulus nottii species group is composed of five fresh-
water species of North American cyprinodontids distinguished from
other species of the genus by the presence of a subocular teardrop.
The group as a whole is distributed from Virginia to Texas along
the coastal plain and in parts of the Mississippi River drainage
north to Michigan. Agassiz (1854) named and briefly described
five species wliich he placed in the genus Zygonectes: Z. nottii,
Z. guttatus, Z. hieroglyphicus, Z. lineolatus, and Z. dispar. Jordan
and Brayton (1878) recognized Z. nottii but doubted that Z.
guttatus or Z. hieroglyphicus could be recognized from published
descriptions. Five other nominal species were described by other
authors: Z. melanops Jordan (1878), Z. inurus Jordan and Gilbert
(1883), Z. craticula Goode and Bean (1882), Z. zonifer Jordan
and Meek (1884), and Z. escambiae Bollman (1886). In addi-
tion, Jordan (1886) considered Fundulus zonatus Cuvier and Val-
enciennes (1846) a synonym of Z. nottii. Jordan and Gilbert
(1883) recognized Z. megalops and Z. dispar as valid species,
stated that Z. nottii, Z. guttatus, and Z. hieroglyphicus had not
been recognized since Agassiz's descriptions, and reported that
Professor Putnam considered Z. lineolatus and Z. nottii identical.
Hay (1885) recognized the validity of Z. nottii. Hay's recognition
was acknowledged by Jordan (1886) and Jordan and Everaian
(1896). Gilbert (1891) recognized Z. guttatus as valid and

1 Assistant Curator, Museum of Natural History, University of Kansas,
Lawrence, KS 66045.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

synonymized Z. escambiae with it. Carman (1895) revised the
entire complex and recognized a single species, Z. nottii. Jordan
and Evermann (1896) made Zygonectes a junior synonym of
Fundulus and recognized three species: F. nottii, F. dispar, and
F. guttatus. They considered F. escambiae and F. lineolatus as
synonyms of F. nottii. This same arrangement was followed by
Jordan, Evermann, and Clark (1930) except that Zygonectes was
resurrected as a valid genus.

Hubbs and Allen ( 1943 ) recognized a single species, Fundulus
dispar, and three subspecies: F. d. dispar (northern), F. d. nottii
(Mobile Bay area), and F. d. lineolatus (Florida), Carr (1936)
first used this arrangement ( based on consultation with Hubbs and
Allen ) but appHed the trivial specific name notti rather than dispar.
The nomenclature of Hubbs and Allen (1943) was used by most
investigators until Brown's (1958) revision of the group (e.g.,
Frey, 1951; Knapp, 1953; Bailey et al, 1954). Frey (1951) re-
ported on the variation of several meristic characters of North
Carohna F. lineolatus and was the first to report that water quality
has a pronounced effect on color pattern intensity. Knapp (1953)
illustrated Fundulus blairae (Wiley and Hall, 1975) from Texas,
terming this species F. dispar. Bailey et al. ( 1954 ) stated that sev-
eral subspecies of F. dispar could be recognized but deferred any
taxonomic decision pending a thorough study of geographic varia-
tion. Miller (1955a) followed the then unpublished revision of
Brown (1958) in recognizing F. nottii as the correct name of
the species and commented that F. nottii was conspecific with
F. dispar.

Brown (1958) studied populations of the F. nottii complex
from large portions of its eastern range. He recognized Carman
(1895) as first reviser and thus adopted the specific name nottii
(which following recommended nomenclatural practice of that
time he spelled notti ) . Brown ( 1957, 1958 ) recognized three sub-
species. Fundulus notti notti was distributed from the New River,
Florida to eastern Texas. Fundulus n. lineolatus was distributed
from rivers east of the New River south to central Florida and
north to Virginia along the coastal plain. Fundulus n. dispar was
distributed in the Mississippi River tributaries from Tennessee to
Michigan.

Rivas (1966) recognized F. lineolatus as a species distinct from
F. nottii based on their sympatric occurrence in Florida. Criffith
(1974) used the name F. swampinus (Lacepede) for F. lineolatus
on the grounds of priority. This nomenclatural change is unwar-
ranted because Hydrargira swampina Lacepede (1803) differs in
several meristic and color pattern characteristics from Fundulus
lineolatus Agassiz and is best synonymized with F. heteroclitus ( see
Bailey and Wiley, 1976, for discussion). Thus, the name F.
lineolatus is used here as the oldest valid synonym of the species.

FUNDULUS NOTTII SPECIES GROUP 3

Wiley and Hall (1975) described F. hlairae from western Louisi-
ana, eastern Texas, and southeastern Oklahoma. We recognized
the other forms ( nottii, lineolatus, and dispar ) as full species based
on confirmation of Rivas' (1966) conclusions, and analyzed the
phylogenetic relationships between the species of the group and
other Fundulus.

The present study completes the revision of the F. nottii
species group begun by Wiley and Hall (1975). The "eastern"
and "western" populations of F. nottii recognized by Wiley and
Hall ( 1975 ) are recognized here as full species ( F. escamhiae and
F. nottii, respectively). Synonymies, diagnoses, and re-descrip-
tions are given for these species plus F. lineolatus and F. dispar.
A synonomy and re-diagnosis is given for F. hlairae. Synonymies
include only major taxonomic references and state-wide faunal
works where the new usage of names might cause future confusion
in identification. A revised phylogenetic analysis is presented
which incorporates F. escamhiae. Finally, some aspects of the
biogeography of the group are discussed.

Methods and Materials

Body color patterns were determined by visual inspection of
alcohol preserved specimens and terminology follows Wiley and
Hall (1975) with three additions. A counter teardrop is defined as
a discrete aggregation of melanophores similar in shape but
smaller than the subocular teardrop, which runs from the orbit to
the posterior nares where it may blend with general melanophore
development on the top of the head. A cleithral stripe is defined
as a discrete aggregation of melanophores running from the base
of the pectoral fin to the dorsal junction of the operculum with the
body. Flank dots are rounded aggregations of melanophores and
chromophores while dashes are thin elongate melanophore ag-
gregations.

Fin ray counts follow Hubbs and Lagler (1947). Note that
Brown's (1958) and Wiley and Hall's (1975) fin ray counts result
in one more fin ray per fin than the standard method. The modes
and ranges given by Brown (1958) and Wiley and Hall (1975)
are adjusted herein to make them similar to standard counts.

Head squamation patterns (Fig. 1) foUow the terminology of
Hoedeman (1958). The E scale(s) is defined as that medial scale
or pair of scales which is bordered by the fused or separate pore 4
of the supraorbital sensory canal. The single or paired G scale(s)
is that medial scale(s) immediately anterior to the E scale(s). The
A scale is that medial scale immediately posterior to the E scale(s).
Head squamation patterns are defined below.

E-type Squamation. — The single or paired E scale(s) overlap
both the G scale(s) and A scale (Fig. la).

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

G-type Squamation. — The single or paired G scale(s) overlap
both E scales (Fig. lb).

A-type Squamation. — The A scale overlaps the single or paired
E scale(s) which in turn overlaps the single or paired G scale(s)
(Fig. Ic).

E-G Squamation. — One of the E scales (but not both) is over-
lapped by the G scale (Fig. Id).

E-A Squamation. — The A scale overlaps one but not both of
the E scales (Fig. le).

Initial analysis of a limited number of specimens from each lot
of Tulane specimens did not allow for variation within samples to
be assessed for this character (10 specimens per lot). Therefore,
when it was brought to my attention that squamation patterns
varied v^thin species and within populations by Dr. Gerald Smith
(University of Michigan Museum of Zoology), I rechecked for
variation using the UMMZ specimens cited in the appropriate sec-
tions below.

Supraorbital sensory canal pore terminology follows Gosline
(1949). The distance between pores 4a and 4b is defined as dis-
tance "A" in Figure 2, the distance between pores 4a and 3 is
defined as distance "B" in the same figure. A fused pore 4 is
termed 4a+b and is defined as a single oblong or round opening
associated with the E scale between pores 3 and 5. Fused pores
lack a complete septum between 4a and 4b. The ontogeny of the
pore canal system as evidenced by comparison of individuals of
various sizes indicates that in those species in wliich adults usually
have fused 4a4-b the juvenile condition is commonly one of
unfused but close association. Thus there are many specimens of
young ( and occasional adults ) where the distinction between close
but unfused and fused is arbitiary since both conditions are parts
of the same ontogenetic sequence.

Map localities shown in Figure 9 are those for which positive

Fig. 1. — Diagramatic drawing of variation in E, G, and A head squama-
tion patterns in species of the Funduhis nottii species group, dorsal views,
(a) E-type squamation (F. lineolatus); (b) G-type squamation (F. blairae);
(c) A-type squamation (F. escamhiae); (d) E-G variant (F. dispar); (e) E-A
variant (F. nottii). Pores 4a and 4b of the supraorbital sensory canal num-
bered, refer to text for variation within each species.

FUNDULUS NOTTII SPECIES GROUP

geographic locality of specimens examined could be determined
and those localities given by other authors for species that were
positively identifiable by description or figure.

The following abbreviations are used:

Institutional Abbreviations,

AMNH — American Museum of Natural History
CU — Cornell University

KU — University of Kansas Museum of Natural History
MCZ — Museum of Comparative Zoology, Harvard

University
NLU — Northeast Louisiana State University
SHSU — Sam Houston State University, Texas
SSC — Southern State College, Arkansas
TU — Tulane University Museum of Natural History
UMMZ — University of Michigan Museum of Zoology
USNM — U.S. National Museum of Natural History

Other Abbreviations.

N,S,E,W— North, South, East, West

Pi — pectoral fin

Po — ^pelvic fin

SL — standard length

trib. — tributary

Co. — county
cr. — creek
dr. — drainage
Hwy. — highway
jet. — junction

Fig. 2. — Diagrammatic drawing of a portion of the right supraorbital sen-
sory canal of Fundulus lineolatus, dorsal view. Numbers refer to pores of the
supraorbital sensory canal, distances "A" and "B" are defined in the text.

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Systematic Accounts

The Fundulus nottii species group is characterized by the pres-
ence of a subocular teardrop in both sexes and six to eight stripes
on the flanks between the scale rows of females (Wiley and Hall,
1975). A key to the five species recognized here is given below,
la. Mature females with vertical bars or with regular rows
of dots on the flank, mature males with vertical bars
or without vertical bars and without caudal fin pigment
blotches. Pores 4a and 4b of the supraorbital sensory
canal system either fused or their distance less than or
equal to the distance between pores 3 and 4a. Head
squamation usually E-type, occasionally E-type variant
rarely G or A-type except for individuals with regen-
erated head scales 2

b. Mature females without vertical bars, mature males
with thin vertical bars and with flank dots restricted to
the upper half of the body or without vertical bars and
with regular or irregular rows of dots over entire
flank and caudal fin pigment blotches. Pores 4a and 4b
of the supraorbital sensory canal widely separated,
their distance equal to or more than the distance from
pore 4a to pore 3. Usually G-type head squamation
patterns with occasional individuals having one E-scale

overlapping the posterior edge of the G scale 4

2a. Males with vertical bars conspicuously thickened in
the middle and tapering to a point ventrally. Females
without dots on the flank and usually with subdermal
vertical bars extending past the pelvic fin insertion.
Both sexes with some pigment on the pectoral fin base
but without a well developed cleithral stripe. Mature
specimens usually with pores 4a and 4b unfused and
separate, rarely with one side fused. Scales on the
upper opercular without pigment on their edges. Scales
around the caudal peduncle modally 16 (range 16-19,

rarely 18 or 19) F. lineohfus

b. Males with vertical bars of even width, females usually
with dots on flank and with subdermal vertical bars
ending at or before pelvic fin insertion. Mature speci-
mens usually with a single pore 4; 4a and 4b rarely
separate (young with fused or separate pores). Males
and females with a conspicuous cleithral stripe from
pectoral fin to dorso-posterior insertion of operculum.
Scales on the upper half of the operculiun with mela-
nophores around their exposed edges. Scales around

the caudal peduncle modally 19-20 (range 18-20) 3

3a. Males with vertical bars which extend anteriorly to or

FUNDULUS NOTTII SPECIES GROliP 7

near the pectoral fin base and ventrally past the lowest
row of flank dots. Females with flank stripes inter-
connected by heavy melanophore development on the
posterior edges of flank scales giving the impression of
a series of rows of light blotches on the dorsal two-
thirds of the flank. Dot pattern of females irregular,
not forming two or more rows on the flank F. nottii

b. Males with vertical bars ending anteriorly between the
paired fins on the flank, these bars not extending ven-
trally past the lowest row of dots. Some males without
vertical bars. Females without heavy melanophore
development on the posterior edges of flank scales —
flank stripes not interconnected by dark flank pig-
ment on the posterior scale edges. Dots present or
absent on females, in two or more regular rows when
present F. escambiae

4a. Males with three to twelve vertical bars, females with
little or no melanophore development between the
flank stripes, young of both sexes with vertical bars

F. dispar

h. Males without vertical bars, females with numerous
dashes, dots, and less discrete melanophore develop-
ment between the flank stripes. Young of both sexes
lacking vertical bars F. blairae

Fundulus nottii (Agassiz)

Western Starhead Topminnow

Figs, le, 3, 8, 9, 10

Zygonectes nottii Agassiz, 1854:353. — ^Jordan and Brayton, 1878:
48.— Gilbert, 1882:341.— Hay, 1885:557.— Garman 1895:121 (in
part). — Jordan, Evermann, and Clark, 1930:178.

Zygonectes guttatus Agassiz, 1854:353. — Jordan and Gilbert, 1883:

341. — ^Jordan, Evermann, and Clark, 1930:178.
Zygonectes hieroglyphicus Agassiz, 1854:353. — Jordan and Gilbert,

1883:341.

Fundulus craticula (Goode and Bean). — Jordan 1886:527.

Fundulus nottii (Agassiz). — Jordan and Evermann, 1896:656 (in
part). — Smith- Vaniz, 1968:79 (in part, Alabama populations
west of the Styx River).— Wiley and Hall, 1975:2 (in part,
"western populations").

Fundulus guttatus (Agassiz). — ^Jordan and Evermann, 1896:658.

Fundulus notti notti (Agassiz). — Carr, 1936:82. — Brown, 1957:75,
1958:483 (in part, populations west of the Styx River, Ala-
bama).

Fundulus dispar nottii (Agassiz). — Hubbs and Allen, 1943:123.

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fundulus dispar (Agassiz). — Bailey, Winn and Smith, 1954:132

(in part).
Fundulus notti (Agassiz). — Miller, 1955a: 10 (in part). — Cook,

1959:151.— Moore, 1968:112 (in part).

Type material: MCZ 1365 (4 males), MCZ 1366 (5 females,
1 male), and MCZ 1449 (1 male, 1 female). All specimens from
the vicinity of Mobile, Alabama.

Other material examined: ALABAMA. — Escatawpa River dr.: TU 21165
(1). Mobile Bay dr.: MCZ 1365 (4, Types, F. guttatus); TU 1824 (382);
TU 2602 (11); TU 3092 (11); TU 27444 (11); TU 32538 (1); TU 33905
(18); TU 44306 (31). LOUISIANA.— Bayou Chitto dr.: TU 7906 (1); TU
14313 (1). Pascagoula River dr.: TU 28105 (5). Pearl River dr.: TU 93
(77); TU 735 (10); TU 1184 (13); TU 1370 (42); TU 1421 (1); TU 1680
(6); TU 7713 (11); TU 7779 (3); TU 8348 (1); TU 11468 (5); TU 15143
(4); TU 15466 (26); TU 17408 (5); TU 18624 (6); TU 23394 (3); TU
24779(1); TU 25822 (1); TU 28840 (3); TU 32285 (2); TU 39850 (2);
TU 40063 (9); TU 43318 (1); TU 44946 (27); TU 53578 (10); TU 52309
(91); TU 53131 (2); TU 55818 (1); TU 61982 (1); TU 61999 (1); TU
71262 (322). UMMZ 163684 (17, ditch 1 mi NE of Pearl River, St. Tam-
many Parish). Lake Pontchartrain dr.: TU 2938 (32); TU 46100 (2); TU
47699 (3); TU 92350 (10); USNM 124769 (6). Tangipohoa River dr.: TU
1033 (6); TU 3589 (5); TU 45550 (1); TU 80007 (6); TU 83009 (2);
TU 83028 (1);TU 84133 (1);TU 84173 (2); TU 84204 (2); TU 84224 (1).
Tchefuncta River dr.: TU 1990 (11); TU 74882 (1); TU 74899 (1); TU
75092 (1); TU 80959 (5); TU 81027 (1). MISSISSIPPI.— Bay St. Louis dr.:
TU 4520 (4); TU 7662 (3). Jordan River dr.: TU 27075 (1); TU 89141 (1).
Pascagoula River dr.: TU 567 (11); TU 51445 (1); TU 52075 (1); TU
54732 (60); TU 60473 (171); TU 65858 (58); TU 86225 (34). Pearl River
dr.: TU 64 (2); TU 86 (5); TU 4776 (3); TU 4916 (1); TU 1727 (1);
TU 7676 (21); TU 10801 (50); TU 15198 (2); TU 16774 (26); TU 17654
(5); TU 17751 (1); TU 23562 (38); TU 25797 (19); TU 23562 (8); TU
28067 (1); TU 28613 (19); TU 28728 (4); TU 29104 (92); TU 36092 (1);
TU 43464 (1); TU 82093 (1); TU 86403 (1); TU 89718 (1). Wolf River
dr.: TU 59478 (2); TU 64628 (3); TU 62834 (2); TU 68420 (3); TU
62521 (3); TU 79816 (1).

Diagnosis: Males, females, and young with a subocular tear-
drop and a cleithral stripe. Females with flank stripes intercon-
nected by pigment on the posterior edges of the flank scales and
with flank dots usually randomly distributed. Females with vertical
bars not extending anteriorly past the pelvic fin insertion. Males
with vertical bars of even width extending ventrally past the most
venti-al row of flank dots onto the belly, and anteriorly to or near
the pectoral fin insei-tion. Modally with E-type head squamation
and pores 4a and 4b of the supraorbital sensory canal fused.

Description: Color pattern descriptions are based on 1,918
alcohol preserved specimens. Typical male and female shown in
Figure 3. Males and females with the dorsal three or four scale
rows with pigmentation uniformly dark or forming a reticulate
pattern on the exposed edges of the scales. Males with seven to
eight rows of dots on the flank, the most ventral row not extending
anteriorly past the insertion of the pelvic fin. Dots on the flank of

FUNDULUS NOTTII SPECIES GROUP

9

Fig. 3. — Fundulus nottii. Top, male, 45 mm SL; bottom, female, 51 mm
SL (UMMZ 163684, Pearl River drainage).

females usually irregularly placed, occasionally forming one row
ventrally. Females with six to eight flank stripes, all but the most
ventral one or two stripes interconnected by pigment on the
posterior edges of the flank scales. Females over 37 mm standard
length with connecting pigment bars as dark as the stripes whereas
in small females the pigment bars are lighter than the stripes. The
most dorsal stripe in females occasionally fades into general pig-
ment development on the back. Males less than 35 mm standard
length with or without stripes which are lighter than flank dots,
stripes indistinct or missing in larger males. Males with nine to
fifteen vertical bars of dermal melanophores between the base of
the caudal peduncle and to or near the pectoral fin insertion, these
bars of even width, extending ventrally to or past the most ventral
row of flank dots and onto the belly. Females with two to six
subdermal vertical bars between the base of the caudal peduncle
and the insertion of the pelvic fin. Young with five to thirteen
vertical bars. All individuals with a cleithral stripe running from
the pectoral fin base to the junction of the operculum with the
body. The melanophores of the cleithral stripe underlain by
guanine on the lower base of the pectoral fin. Exposed edges of
the opercular scales above mid-eye level with melanophores form-
ing a light reticulate pattern. A teardrop of solidly fused mela-
nophores underlain with guanine extends from the ventral orbital
margin to the preopercular in mature individuals. An oblong coun-
ter teardrop extending from the antero-dorsal orbital border to
the nares. Isthmus overlain with guanine, isthmus either lacking

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

melanophores or having one or two melanophores atop the gua-
nine. Mouth bordered by melanophores. The region between the
mouth and the orbital margin on the side of the head with Httle
or no pigment when preserved and, for those few live specimens
examined, with red-orange pigment in life. The rays of all fins
bordered by separate melanophores. Pelvic and pectoral fins with-
out pigment blotches. Four to six transverse rows of pigment
blotches on the anal fin of males, one to three rows on females.
Dorsal fins of both sexes dusky, males with three or four trans-
verse rows of blotches on the fin membrane, females lacking
blotches or having one or two rows of blotches. Caudal fin of
males with or without pigment blotches, when present blotches are
restricted to the anterior half of the caudal fin. Females without
pigment blotches on the caudal fin.

Head squamation usually E-type (Fig. 8). [Of 62 specimens
examined from the Wolf River, Mississippi (UMMZ 163698) 42
were E-type, 14 the E-G variant, 3 the E-A variant, and 3 were
G-type. Specimens from other parts of the range showed a pre-
dominance of E-type squamation, but sample sizes per lot were
too small to determine a useful frequency distribution.] Mature
specimens usually with pores 4a and 4b of the supraorbital sen-
sory canal fused ( Fig. 8 ) , occasional mature specimens with these
pores close but unfused on one side, rarely with both sides un-
fused. Young with fused or unfused pores 4a and 4b.

Meristic counts on 50 specimens from the Tangipohoa (11),
Pearl (26), and Tombigbee (13) river drainages showed the fol-
lowing modes and ranges: anal fin rays 9 (9 — 10); dorsal fin rays
7 (6—8); pectoral fin rays 12 (11—13); lateral hne scales 33 or 34
(32 — 35); scales around the body 24 or 25 (23 — 27); scales around
the caudal peduncle 18 or 19 (17—21).

Range: From the Mobile Bay drainage of Alabama westward
below the Fall Line along the Gulf coastal plain to the Lake
Ponchartrain drainage, Louisiana (Fig. 9).

Etymology: Trivial name in honor of Dr. Nott, the discoverer
of the species.

Remarks: The type of Zygonectes hieroglyphicus (MCZ 1368,
1 spec.) apparently was discarded in 1938 (R. Schoknecht, MCZ,
pers. comm.).

Fundulus escamhiae (Bollman)

Eastern Starhead Topminnow

Figs. Ic, 4, 8, 9, 10

Zygonectes escamhiae Bollman, 1886:463.
Zygonectes nottii Agassiz. — Carman, 1895:120 (in part).
Fundulus guttatus (Agassiz). — Jordan and Evermann, 1896:658
(in part).

FUNDULUS NOTTH SPECIES GROUP 11

Zygonectes guttatus Agassiz. — Jordan, Evemiann, and Clark, 1930:

178 (in part).
Fundulus notti notti (Agassiz). — Carr, 1936:82 (in part). — Carr

and Coin, 1955:73.— Brown, 1957:75 (in part), 1958:483 (in

part).
Fundulus dispar nottii (Agassiz). — Hubbs and Allen, 1943:123 (in

part).
Fundulus dispar (Agassiz). — Bailey, Winn, and Smith, 1954:132.
Fundulus notti (Agassiz). — Miller, 1955a: 10 (in part). — Rivas,

1966:353.— Moore, 1968:112 (in part).— Relyea, 1975:50.
Fundulus nottii (Agassiz). — Smith- Vaniz, 1968:78 (in part). —

Wiley and Hall, 1975:2 ("eastern population").

Type material: USNM 37994 (1), a male from Flomaton, Ala-
bama.

Other material examined: ALABAMA. — Apalachicola (Chattahoochee)
River dr.: TU 3323 (52). Escambia River dr.: TU 15256 (18); USNM
37994 (1, holotype); USNM 94380 (1). Styx River dr.: TU 87818 (45, 3.6
mi. E. Seminole, Baldwin Co.). GEORGIA. — Apalachicola ( Flint-Chickasa-
watchee) River dr.: TU 6184 (21, 6 mi. E. Leary on Uwy 62, Dougherty
Co.). FLORIDA.— Apalachicola River dr.: TU 2045 (6); TU 2297 (1);
TU 2540 (2); TU 20550 (25); TU 20768 (1); TU 22557 (1); TU 32017
(37); TU 32043 (15); TU 32130 (76); TU 32152 (37); TU 32176 (5);
TU 32225 (24); TU 33331 (24); TU 33870 (44); TU 34823 (31); TU 45606
(4); TU 45780 (19); TU 46272 (4). Blackwater River dr.: TU 18343 (45);
TU 23693 (176); TU 27407 (23); TU 46429 (4); TU 57073 (1); TU 87783
(12); TU 88867 (2); TU 88889 (10); TU 92038 (1). Choctawhatchee River
dr.: TU 309 (3); TU 1060 (3); TU 1081 (9); TU 1105 (3); TU 1592 (1);
TU 1607 (25); TU 1660 (1); TU 1688 (1); TU 2314 (6); TU 2467 (19);
TU 2491 (3); TU 2640 (1); TU 15688 (1); TU 45650 (1); TU 45653 (2);
TU 45874 (12); TU 46087 (12); TU 46229 (24); TU 88840 (15, Black Cr.
at Hwy. 394, Walton Co.). East Bay dr.: TU 21450 (1); TU 41196 (1);
TU 45640 (2); TU 45890 (5); TU 46682 (3). Escambia Bay dr.: TU
27429 (8). New and Carrabelle River drs.: TU 39567 (1), TU 42780 (1),
and TU 42782 (2)-4 mi. NE Carabelle on Duck Lake, Franklin, Co.; TU
44459 (27, ditch 5.5 mi. N. Carabelle, 3 mi. W. of junction Hwys 98 and 67
on Gullie Branch Road, Franklin Co., New River); TU 46895 (36, swamp
2.7 mi. N. Carabelle on Hwy. 67, Franklin Co., Carabelle River). Ochlocknee
River dr.: TU 34934 (1, trib. of Telogia Cr., 1.2 mi. S. Greensboro on Hwy.
12, Gadson Co.). Pensicola Bay dr.: TU 9761 (5); TU 10493 (6); TU
27407 (23). Perdido River dr.: TU 82837 (3) and TU 88923 (43)-jct.
Perdido River and Hwy. 184 at Muscogee, Escambia Co.; TU 85535 (4,
jet. Perdido River and Interstate Hwy. 10, Escambia Co.); TU 87732 (9,
McDavid Cr., 3.7 mi. NNW Barrineau Cr. on Hwy. 99, Escambia Co.); TU
88986 (1, jet. Hwys. 98 and 298, Escambia Co.). St. Andrews Bay dr.: TU
46309 (2). St. Marks River dr.: TU 46689 (1) and TU 69688 (2)-McBrides
Slough, trib. Wakulla River, 2.5 mi. NW of Wakulla on Hwy. 267, Wakulla
Co.); TU 75396 (3, jet. Hwy. 98 and St. Marks River, Wakulla Co.). Su-
wannee (Sante Fe) River dr.: TU 8440 (1, Sante Fe River 2.4 mi. NW of
High Springs on Hwy. 12, Columbia Co.); TU 36514 (8, spring .5 mi. from
mouth of Sante Fe River, Gilchrist Co.). Yellow River dr.: TU 35939 (3);
TU 46104 (1).

Diagnosis: Males, females, and young with a subocular tear-
drop and a cleithral stripe. Females with flank stripes, or when

12

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

rff ;.^..>»V-*:«;:^r*--

^ V b • «

• iSSjit/^^f^^

Fig. 4. — Fundulus escamhiae. Top, male, 52 mm SL; bottom, female, 50
mm SL (TU 88840, Choctavvhatchee River drainage).

flank stripes are very light or missing, then with regular rows of
dots on the flank. Males with vertical bars restiicted to the middle
region of the flank anteriorly, these vertical bars not extending
past the most ventral row of flank dots and not extending anteriorly
to the pectoral fin insertion. Females and young without pigment
on the posterior edges of the flank scales as dark as stripes when
stripes are present. Modally with E-type head squamation and
usually with pores 4a and 4b of the supraorbital sensory canal
fused.

Description: Color pattern descriptions are based on 1,112 al-
cohol preserved specimens from throughout the range of the
species. Typical male and female shown in Figure 4. Dorsum
dark, washing out stripes and dots or forming a reticulate pattern
on most dorsal scale rows. Males with seven to nine rows of dots,
the most ventral row extending anteriorly to one-half the distance
between Pi and P2 fin bases. Larger females from the Perdido to
Choctawhatchee drainages with dots in seven to nine regular
rows whereas smaller females from these drainages and females
from more eastern drainages variable, some lacking dots and some
v^dth rows ending half-way posteriorly on the flank. Stripes when
present in males not as dark as dots. Females with six to eight
stripes, uppermost fading into background coloration anteriorly.
Males with zero to fourteen vertical bars of even width and re-
stricted to the mid flank anteriorly. Females with zero to five
vertical bars not reaching anterior insertion of the anal fin. Most
young with vertical bars of even intensity and height from base of
caudal fin to insertion of the pectoral fin. A vertical stripe from

FUNDULUS NOTTII SPECIES GROUP 13

the dorso-posterior junction of the body and opercular flap along
the edge of tlie cleitlirum onto the base of the pectoral fin in all
individuals. Head dark above. Teardrop of solidly fused mela-
nophores extending from the ventral edge of the orbit to the
preopercular canal on the cheek. Teardrop and isthmus underlain
with guanine. Isthmus without melanophores. An oblong counter-
teardrop from the anterio-dorsal border of the orbit to the nares.
Mouth bordered by melanophores, the space between tliis pigmen-
tation and the orbit dusky in males, Hght to dusky in females and
young. In life, those few specimens examined have red-orange pig-
ment between the mouth and orbit. Under side of the head dusky.
Operculum with melanophores bordering the scales on the dorsal
half, this pigmentation extending ventrally below mid-eye level.
All fins with rays bordered by melanophores. Caudal, pectoral, and
pelvic fins lacking blotches. Distal parts of pectoral fin membranes
dusky. Dorsal fin of males without blotches but with extensive
melanophore development on the fin membrane. Anal fin of males
with two to four rows of blotches where usually the first and occa-
sionally the second rows reach the first fin ray. Females without
blotches on the anal fin or with a single row of blotches.

Head squamation modally E-type (Fig. 8). [Of 81 specimens
examined from the St. Marks River, Florida (UMMZ 163425) 64
were E-type, 3 were E-G variants, 7 were E-A variants, 6 were
G-type, and 1 was A-type. Other specimens examined throughout
the range showed a predominance of E-type squamation, but the
number of specimens examined per lot was too small to determine
useful frequency distributions.] Pores 4a and 4b of the supra-
orbital sensory canal usually fused, occasionally unfused on one
side in adults. These pores frequently separate in young.

Meristic counts on 98 specimens from the Sante Fe (8), Apa-
lachicola (31), Choctawhatchee (25), Blackwater (16), and Es-
cambia ( 18 ) river drainages showed the following modes and
ranges: anal fin rays 9 (8 — 10); dorsal fin rays 7 (6—8); pectoral
fin rays 13 (11 — 14); lateral line scales 34 (33 — 36); scales around
the body 23 — 25 (22 — 27); scales around the caudal peduncle 19
(16—20).

Geographic variation: Males from the Escambia River may
lack vertical bars on the side of the body or have only one or two
bars when preserved. Females and young from the Perdido to
Choctawhatchee rivers may lack vertical bars and have regular
rows of flank dots and light flank stripes whereas females from the
Apalachicola River and eastward may lack dots and have dark
flank stripes. What taxonomic significance this variation indicates
can not be assessed without further studies involving larger
samples (and perhaps different characters) than those used here.

Range: From the Perdido River, Alabama and Florida, east-

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

ward below the Fall Line along the Gulf coastal plain of Florida
and Georgia to the Santa Fe River, Florida (Fig. 9).
Etymology: Named for the Escambia River.

Fundulus lineolatus (Agassi z)

Lined Topminnow

Figs, la, 2, 5, 8, 9, 10

Zygonectes lineolatus Agassiz, 1854:353.

Zygonectes craticula Goode and Bean, 1882:433. — Jordan and Gil-
bert, 1883:892.

Zygonectes zonifer Jordan and Meek, 1884:482. — Jordan, 1886:527.

Zygonectes nottii Agassiz, — Garman, 1895:121 (in part). — Jordan,
Evermann, and Clark, 1930:178.

Fundulus zonatus (Cuvier and Valenciennes). — Jordan, 1886:527.

Fundulus nottii (Agassiz). — Jordan and Evermann, 1896:656 (in
part).

Fundulus nottii lineolatus (Agassiz). — Carr, 1936:82. — Brown, 1957:
76, 1958:497.— Carr and Coin, 1955:77.

Fundulus dispar lineolatus (Agassiz). — Hubbs and Allen, 1943:123.

Fundulus notti (Agassiz). — Miller, 1955a: 10 (in part). — Moore,
1968:112 (in part).

Fundulus lineolatus (Agassiz). — Rivas, 1966:353. — Wiley and
Hall, 1975:1.— Relyea, 1975:50.

Fundulus swampinus (Lacepede). — Griffith, 1974:320.

Types.— UCZ 1451 (1 male and 1 female), MCZ 1344 (2
females). Augusta, Georgia.

Other material examined: GEORGIA. — Alapaha River dr.: USNM 162424
(7). Canoochee River dr.: TU 16475 (3). Suwannee (New) River dr.:
USNM 28505 (1, Type Z. zonifer). Occnce (Attamaha) River dr.: TU 16437
(1); USNM 43459 (5); USNM 61556 (6). Satilla River dr.: TU 21196 (136,
Satilla River, 7 mi. E. of Nahunta on Hwy. 84, Brantle Co.); TU 41272 (6).
Savannah River dr.: USNM 82639 (7); USNM 82640 (4). Suwannee River
dr.: USNM 179194 (2); USNM 196720 (5). FLORIDA.— Econfina River
dr.: TU 5040 (6, Econfina River 2.9 mi. NNE of Otter Cr. on Hwy. 24,
Levy Co.). Fenholloway River dr.: TU 36123 (1, Fenfolloway River 6 mi.
N. of Perry on Hwy. 27, Taylor Co.). Indian River dr.: USNM 31439 (1, type
Z. craticula). Oldawana River dr.: TU 12634 (13); USNM 44863 (2);
USNM 83936 (6); USNM 106438 (13); USNM 133535 (15). Santa Fe
(Suwannee) River dr.: TU 74396 (5); USNM 92851 (27); USNM 94922
(1). St. John's River dr.: USNM 25338 (8); USNM 63949 (1); USNM
125285 (1). St. Mary's River dr.: TU 21213 (4); TU 21314 (7). Suwannee
River dr.: TU 34934 (6). Tampa Bay dr.: USNM 102468 (1); USNM
106968 (1). Waccasassa River dr.: TU 4729 (2). Withlacoochee River dr.:
TU 9833 (3). NORTH CAROLINA.— Fear River dr.: USNM 49147 (1);
USNM 52069 (40); USNM 94235 (4); USNM 94380 (1); USNM 94387 (1).
Neuse River dr.: USNM 191010 (27). SOUTH CAROLINA.— Congaree
River dr.: TU 7347 (1); USNM 21591 (1); USNM 213931 (4); USNM
213938 (1); USNM 213940 (3); USNM 213941 (5); USNM 213942 (1);
USNM 213943 (12); USNM 213944 (2); USNM 213945 (2); USNM 213946
(7); USNM 213947 (2); USNM 213950 (2); USNM 213951-213954 (1

FUNDULUS NOTTII SPECIES GROUP

15

each); USNM 213956-213959 (1 each); USNM 213960 (7); USNM 213961
(18). Edisto River dr.: USNM 213939 (2); USNM 213948 (29); USNM
213949 (11); USNM 213955 (7); USNM 213935 (1); USNM 213957 (15).

Diagnosis: Males, females, and young with a subocular tear-
drop and a pigment patch on the upper half of the pectoral fin.
Flank of females with stripes, without pigment on the posterior
edges of the scales as dark as the stripes, and without dots. Males
with vertical bars distinctly thickened in the middle and tapering
to a point ventrally; females usually with numerous vertical bars
extending anteriorly past the pelvic fin insertion. Head squama-
tion usually E-type, pores 4a and 4b of the supraorbital sensory
canal closely situated, their distance equal to or less than the
distance between pores 4a and 3. Scales around the caudal ped-
uncle modally 16 (16—19).

Description: Color pattern descriptions are based on 383 alco-
hol preserved specimens. Typical male and female shown in Figure
5. Dorsal three or four scale rows as light as flank, but in some
individuals pigment on scale edges forms a reticulate pattern.
Males with three to seven rows of small flank dots, these dots
usually absent ventrally. Females without dots and usually with-
out subdermal dashes. Both males and females with seven or eight
flank stripes. Flank stripes of large males lighter than dots or
dermal vertical bars; flank stripes of females darker than sub-
dermal vertical bars. Females with little or no dark pigment be-
tween the stripes. Some males with the posterior edges of flank
scales pigmented, giving a reticulate pattern. Males with eleven
to fifteen vertical bars distributed from the base of the caudal fin
to the insertion of the pectoral fin. Vertical bars of females three

Fig. 5. — Fundulus lineolatus. Top, male, 43 mm SL; bottom, female, 41
mm SL (TU 21196, Satilla River drainage).

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

to fifteen, more distinct and numerous in smaller females (^38
mm SL). Vertical bars of most males and females distinctly
thicker than comparable bars on other species of the group. Ver-
tical bars of males tapering to a point ventrally and extending past
the last stripe or row of dots onto the belly. Anterior vertical bars
of females restricted to the mid-flank region. Subocular teardrop
of both sexes varying with geograhic region. Southern popula-
tions (Florida, Georgia) have lighter teardrops which "fade" ven-
trally and do not reach the preopercular sensory canal whereas
northern populations have darker teardrops with well defined
ventral edges which reach the preopercular sensory canal. Tear-
drop and isthmus pigmentation underlain with guanine. A counter-
teardrop present. Isthmus without melanophores. Mouth bordered
by fused melanophores. Region from the orbit to the mouth dusky
to unpigmented in preserved specimens. The few live specimens
examined had red-orange pigment between the orbit and the
mouth. Opercular scales without melanophore development on
their edges. All fin rays of both sexes bordered by widely separated
melanophores. Both males and females lacking pigment blotches
on the pectorals, pelvics, and caudal fins. Females without pig-
ment blotches on anal or dorsal fins. Males with three to five
pigment blotches on anal and dorsal fins. Pectoral fin base with a
border or fused melanophores on both sides giving the impression
in most specimens of an open triangle of pigment on the upper
pectoral fin base.

Head squamation usually E-type (Fig. 8). [Of 14 specimens
from Clear Water Lake and 59 specimens from Alton Lake, Florida
(UMMZ 110662 and 166605 respectively) 47 were E-type, 2 were
E-A variants, 1 was an E-G variant, and 3 were G-type. Smaller
samples examined from throughout the range showed a predomi-
nance of the E-type pattern.] Pores 4a and 4b of the supraorbital
sensory canal usually unfused and closely situated, their distance
from each other usually less tlian, rarely equal to, the distance
between pores 4a and 3 (Figs. 2, 8).

Meristic modes and ranges reported by Brown (1958) are given
here for completeness of description. These counts are corrected
counts based on 209 specimens taken from throughout the range
of the species, but not including Brown's specimens from the
Suwannee, Ochlockonee, or New rivers which may represent F.
escamhiae or combined collections of F. escamhiae and F. lineo-
latus. Modes and ranges are: anal fin rays 9 (8 — 10); dorsal fin
rays 7 (7 — 8); pectoral fin rays 13 (11 — 14); lateral lines scales 33
(32 — 36) scales around the body 24 (22 — 27); scales around the
caudal peduncle 16 (16—19).

Geographic variation: Variation in teardrop intensity from
north to south is discussed above. No other clear-cut geographic
variation was noted.

FUNDULUS NOTTII SPECIES GROUP 17

Range: From the Ocklockonee River, Florida and Georgia
( Swift, Yerger, and Parrish, in press ) south to Dade County, Flor-
ida (Relyea, 1975) and north and east to southern Virginia along
the coastal plain (Brown, 1958). Localities plotted in Figure 9
includes Brown's (1958) in North Carolina and Virginia but not
those of Swift, Yerger and Parrish (in press) from the Ocklockonee.

Etymology: The trivial name from the Latin for lined.

Fundulus dispar (Agassiz)

Northern Starhead Topminnow

Figs. Id, 6, 8, 9, 10

Zygonectes dispar Agassiz, 1854:353.— Jordan, 1877a: 67, 1877b :49.
—Jordan and Gilbert, 1883:341.

Zygonectes megalops (Cope). — Jordan, 1878:52. — Jordan and Gil-
bert, 1883:340 (but p. 892 hsts Z. megalops Jordan not of H.
megalops Cope but a synonym of Z. inurus Jordan and Gilbert).

Zygonectes inurus Jordan and Gilbert, 1883:892.

Zygonectes nottii (Agassiz). — Carman, 1895:121 (in part, only
those populations within the range of F. dispar).

Fundulus dispar (Agassiz). — Jordan and Evermann, 1896:658. —
Forbes and Richardson, 1908:212. — Jordan, Evermann, and
Clark, 1930:178.— Kuhne, 1939:75.— Wiley and Hall, 1975:1.

Fundulus notti dispar (Agassiz). — Carr, 1936:82. — Brown, 1957:75,
1958:486.

Fundulus dispar dispar (Agassiz). — Hubbs and Allen, 1943:123. —
Smith, 1965:9 (Illinois).

Fundulus notti (Agassiz). — Miller, 1955a: 10 (in part). — Moore,
1968:112 (inpart).— Pflieger, 1971:404 (in part, Missouri popu-
lations only). — Branson, 1972:76 (in part, Kentucky popula-
tions only). — Clay, 1975:230 (in part, Kentucky populations
only).

Type.: USNM 120298, a male from a creek in Illinois, opposite

St. Louis, Missouri.

Other material examined: ALABAMA. — Tombigbee (Mobile Bay) River
dr.: TU 76844 (9, trib. of the Tombigbee 2.5 mi. SW of Forkland, Green Co.).
ARKANSAS. — Mississippi River dr.: 1 lot SSC (uncat. spec). Ouachita River
dr.: KU 16518 (9, Locust Bayou at Hwy. 4, Calhoun Co.); NLU 11725 (80);
1 lot SSC (4 uncat. spec). INDIANA.— Wabash River dr.: USNM 40781
(1); USNM 40787 (1); USNM 40966 (1); USNM 66637 (1). Bruce Lake:
USNM 69216 (5). Lost Lake: USNM 62120 (15); USNM 65114 (4); USNM
65115 (2); USNM 65116 (25); USNM 65118 (4). ILLINOIS.— Illinois River
dr.: USNM 16935 (1). Mississippi River dr.: TU 22805 (1); UMMZ 105834
(7); USNM 101152 (6). Wolf Lake: UMMZ 162874 (2). LOUISIANA.—
Ouachita River dr.: NLU 248 (4); NLU 798 (4); NLU 836 (3); NLU 840
(6); NLU 843 (6); NLU 844 (13); NLU 852 (20); NLU 939 (15); NLU
940 (39); NLU 1359 (31); NLU 1691 (14); NLU 1693 (7); NLU 1899
(11); NLU 3821 (4); NLU 9523 (2); NLU 9831 (1); NLU 11536 (2);
NLU 13183 (18); NLU 13310 (10); NLU 15157 (1); NLU 1624 (13); NLU

18

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

21003 (1); 1 lot SSU (uncat. spec); USNM 172015 (3); USNM 172917 (1);
USNM 173240 (1). MICHIGAN.— Bawchee Lake: UMMZ 60564 (5); Drag-
on Lake: UMMZ 90149 (5). Fifth Lake: UMMZ 98101 (4). MISSISSIPPI.
— Mississippi River dr.: TU 86075 (6, Legion Lake 5 mi. S. Rosedale, Bolivar
Co.). TENNESSEE.— Mississippi River dr.: UMMZ 161023 (8).

Diagnosis: Males, females, and young with a subocular tear-
drop and without a cleithral stripe or distinct pigment aggregation
on the pectoral fin base. Large females with flank stripes, without
vertical bars, and with few or no flank dots or less discrete pigment
aggregations between the flank stripes. Males with three to
thirteen thin vertical bars, anterior bars restricted to the mid-flank
and not extending ventrally to the belly nor extending anteriorly
to the pectoral fin insertion. Males with flank dots on the upper
half of the flank, "dots" on the ventral half of the flank being dis-
connected flank stripes. Head squamation modally G-type, pores
4a and 4b of the supraorbital sensory canal widely separated, their
distance equal to or more than the distances between pores 4a
and 3.

Description: Color pattern descriptions are based on 423 alco-
hol preserved specimens. Typical male and female shown in Figure
6. Males and females with dorsal three or four scale rows dusky,
almost as light as flank. Males with three to six rows of dots on the
flank. Rows of dots on males indistinct or absent on lower half
of body, lower most rows not extending below pelvic fin base.
Most ventral two or three rows of dots usually not extending pos-

FiG. 6. — Fundulus dis-par. Top, male, 49 mm SL; bottom, female, 59 mm
SL (KU 16518, Ouachita River drainage).

FUNDULUS NOTTII SPECIES GROUP 19

teriorly of pelvic insertion. Females without dots but some large
females with dash marks between the stripes on mid-flank scales
anterior to the pelvic insertion. Females with seven to nine flank
stripes. Female stripes usually continuous dorsally and broken
ventrally. Space between stripes with little pigment except for
anterior subdermal dash marks. Males with flank stripes on ventral
half of the flank, these stripes broken into widely separated pig-
ment blotches giving, in some individuals, the impression of a
series of small dots. Small males with light stripes between the
dots dorsally. Males with three to thirteen vertical bars on the
flank. Vertical bars thin (compared to nottii and lineolatus), an-
terior bars on the mid-flank only, not extending past the most
ventral row of stripe blotches. Some young females with very faint
subdermal vertical bars, others apparently lacking subdermal ver-
tical bars. Larger females without vertical bars. Head with a
subocular teardrop of solidly fused melanophores underlain by
guanine and extending from the ventral edge of the orbital margin
to the preopercular canal. Isthmus not overlain by guanine. Coun-
ter teardrop extending from the anterodorsal orbital margin to the
nares. Anterior orbital margin between the teardrop and the
counter teardrop dusky. Melanophores bordering the mouth sep-
arate ( not fused ) , mouth border dusky. All fin rays bordered with
widely spaced melanophores. Pectorals and pelvics of both sexes
without pigment blotches. Caudal fin of males with variable num-
bers of pigment blotches, caudal fin of females without pigment
blotches. Five or six transverse rows of pigment blotches on the
anal fin of males, present or absent in females, when present in
two to four rows. Males with three to five rows of blotches on
dorsal fins, females usually lacking blotches on dorsal fin, but occa-
sional females have as many as three rows.

Head squation pattern usually G-type, but frequently E-G vari-
ant (Fig. 8). Pores 4a and 4b of the supraorbital sensory canal
widely spaced, their distance usually greater than, rarely equal to
the distance from pores 4a to 3.

Meristic counts for 74 specimens from Arkansas (12), Louisiana
(31), Illinois (9), Michigan (14), and Tennessee (8) show the
following modes and ranges: anal fin rays 10 (9 — 11); dorsal fin
rays 7 (6 — 7); pectoral fin rays 13 (12 — 14); lateral line scales 32
(31 — 34); scales around the body 25 or 26 (22 — 28); scales around
the caudal peduncle 18 or 20 (16—20).

Geographic variation: Populations from the Ouachita River
drainage of Louisiana and Arkansas have lower number of scales
around the body (mode of 25) and scales around the caudal ped-
uncle (mode of 18) than do more northern populations (mode of
26 scales around the body and 20 scales around the caudal
peduncle). Males from the Ouachita drainage are more variable

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

in number of vertical bars (as few as three compared to seven or
more in more northern populations).

Range: From the Ouachita River drainage, Louisiana, north to
southern Michigan, east to the upper Tombigbee River drainage,
Alabama. Localities shown in Figure 9 include reports by Bu-
chanan (1973, Arkansas), Gerking (1945, Indiana), Forbes and
Richardson (1919, Illinois), and Pflieger (1971, Missouri) as well
as localities of specimens examined in this study.

Etymology: From the Latin for dissimilar.

Fundulus blairae Wiley and Hall
Blair's Starhead Topminnow
Figs, lb, 7, 8, 9, 10

Fundulus escambiae (Bollman). — Evermann, 1892:87.

Fundulus guttatus (Agassiz). — Jordan and Evermann, 1896:312

(in part).
Fundulus dispar (Agassiz). — Knapp, 1953:89, fig. 122.
Fundulus notti (Agassiz). — Miller, 1955a: 10 (in part). — Hubbs,

1957:96, 1958:8.— Moore, 1968:112 (in part).
Fundulus notti notti (Agassiz). — Brown, 1957:75, 1958:483 (in

part).
Fundulus blairae Wiley and Hall, 1975:3.

Types: Holotype, AMNH 32721 (1 male). Paratypes, AMNH
32722 (17 specimens including males, females, and juveniles).
Holotype and paratypes from the junction of Neville Bayou and
Texas Highway 105, Liberty County, Texas.

Other material examined: In addition to specimens cited by Wiley and
Hall (1975), the following specimens were examined. ALABAMA. — Mobile
Bay dr.: TU 309 (11, D'Olive Creek .3 mi. S of junction of Hwys. 90 and 89
on 89, Baldwin Co.); TU 35383 (1, Oxbow Lake at Choctaw BliifF camp on
Stimpson Game Reserve, Clark Co.); TU 44212 (1, tributary Blakely River 1
mi. N or Spanish Fort, Baldwin Co.); TU 44306 (31, including specimens of
F. nottii, D'Olive Creek 2.4 mi. Spanish Fort, Baldwin Co.); TU 59400 (1,
Bassett Creek, tributary of Tombigbee River, 7.4 mi. E of Grove Hill, Clarke
Co.); TU 67938 (1, Alabama River at river mile 120.4, Wilcox Bar, Wilcox
Co.). LOUISIANA.— Bayou Cocodrie dr.: TU 87380 (1); TU 87395 (3).
Calcasieu River dr.: TU 41487 (2); TU 41520 (1); TU 43285 (4); TU 50212
(2). Red River dr.: NLU 7985 (21); NLU 13475 (46). SHSU 457 (18);
TU 42349 (2); TU 42387 (1); TU 85449 (2); TU 71432 (13). Tensas River
dr.: USNM 172729 (2); USNM 172787 (1). MISSISSIPPI.— Pascagoula
Rivr dr.: TU 67177 (2, Leaf River 2 mi. N of McLain, Green Co.); TU 7177
(2, same loc. as TU 67177). Pearl River dr.: TU 85816 (3, Pearl River .5
mi. below Hwy. 84, Lawrence Co.). TEXAS.— Neches River dr.: SHSU 454
(15). Sabine River Dr.: KU 6052 (5, Sabine River at Hwy. 59, Panola Co.).
Spring Creek dr.: USNM 118546 (14); SHSU 231 (13); SHSU 288 (2).

Diagnosis: Males, females, and young with a subocular tear-
drop and without a cleithral stripe or distinct pigment aggregation
on the pectoral fin base. Females with flank stripes, without verti-
cal bars, and with numerous dots and less discrete melanophore

FUNDULUS NOTTII SPECIES GROUP

21

Fig. 7. — Fundulus blairae. Top, male, 42 nim SL; bottom, female, 35 mm
SL (KU 6052, Sabine River drainage).

development between the flank stripes. Males without vertical bars
and with flank dots on the lower half of the flank. Head squama-
tion G-type, pores 4a and 4b of the supraorbital sensory canal
widely separate, their distance equal to or more than the distance
between pores 4a and 3 (Fig. 8). Young of both sexes lacking
vertical bars.

Description: See Wiley and Hall (1975). Typical male and

4atb

Fig. 8. — Modal head squamation and supraorbital sensory pore patterns in
(Ifift) Fundulus nottii and F. escambiae (E-type, pores 4a and 4b fused);
(middle) F. Imeolatus (E-type, pores 4a and 4b close but unfused), and
(right) F. dispar and F. blairae (G-type, pores 4a and 4b unfused and widely
separated). Semidiagrammatic, pores numbered, scales lettered, scales and
sensory canals unstippled, body scales not drawn.

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

female shown in Figure 7. M eristic counts made by Wiley and
Hall (1975) followed Brown (1958) whose method of counting
anal and dorsal fin rays differs from the standard. Corrected modes
and ranges of all counts for tliis species are: anal fin rays 10 (8 —
11); dorsal fin rays 7 (6—8); pectoral fin rays 13 (11—14); lateral
line scales 33 (30—36); scales around the body 23—24 (22—27);
scales around the caudal peduncle 17 — 18 (16 — 20).

Range: From the Brazos River drainage, Texas eastward along
the Gulf Coastal Plain to the Mobile Bay drainage, Alabama, and
northeastward to the Red River drainage of southeastern Oklahoma
and northwestern Louisiana. Dr. Henry Robison (Southern State
College, Arkansas) has infonned me that it is also in the tribu-
taries of the Little River ( Red River dr. ) in southeastern Arkansas.
Localities shown in Figure 9 include those of Wiley and Hall
(1975).

Etymology: The trivial name in honor of Ms. Blair Knies.

Phylogenetic Relationships

Wiley and Hall (1975) discussed the phylogenetic relationships
of tlie F. nottii species group as evidenced by the presence of
shared derived (synapomorphic) characters. Our conclusions
were: (1) the species group is a member of the "LSA-V" group
of Chen ( 1971 ) on the basis of the derived characteristics of the
karyotypes of all included species; (2) F. cingulatus is the sister
species of the F. nottii species group based on the color pattern of
males and head squamation pattern; (3) that the nottii species
group is monophyletic based on the presence of female stripes and
a subocular teardrop; (4) that F. dispar and F. blairae are sister
species based on their relatively derived head squamation pattern
and a lack or virtual lack of vertical bars on the flank of females.
We also concluded that there was no evidence which permitted us
to decide the placement of F. nottii and F. lineolatus within the
species group.

The resurrection of F. escambiae as a species has led to a
re-evaluation of the phylogenetic relationships within the species
group. The conclusions reached in the discussion below are sum-
marized in the phylogram presented in Figure 10. This phylogram
is the least rejected of the various possible alternatives considered.

The corroborating characters relating F. dispar and F. blairae
have been discussed by Wiley and Hall (1975). Fundulus lineo-
latus, F. nottii, and F. escambiae are hypothesized more closely
related to each other than to either F. dispar or F. blairae based
on the position of pores 4a and 4b of the supraorbital sensory canal
and pectoral fin base pigmentation.

Fundulus dispar, F. blairae, and most other Fundulus examined
have pores 4a and 4b relatively widely spaced, their distance

FUNDULUS NOTTII SPECIES GROUP

23

Ai

35"

33"

31"

29

Fig. 9. — Geographic distribution of species of the Fundulus nottii species
group. Open circles, F. twttii; closed circles, F. escamhiae; open squares,
F. lineolatus; open triangles, F. hlairae; closed triangles, F. dispar.

usually more than the distance between pores 4a and 3. Species of
Profundulus ( Miller, 1955b ) also have unf used and widely spaced
pores. Fundulus lineolatus usually have pores 4a and 4b closely
situated, their distance usually less than the distance between pores
4a and 3. Most F. nottii and F. escamhiae adults have fused pores
4a and 4b. The only other Fundulus with closely situated pores 4a
and 4b is F. diaphanus (TU 36701), a species usually not con-
sidered a relative of the nottii species group. I conclude that the
pore pattern in F. lineolatus is intermediate between the primitive
pattern of wddely separated pores 4a and 4b and the more derived
pattern of fusion of these pores, thus implying that F. lineolatus is
more closely related to nottii and escamhiae than to dispar and
blairae. The alternate hypothesis, that widely separated pores are
derived, would lead to the conclusion that dispar and hlairae were
more closely related to other Fundulus than to lineolatus, nottii
and escamhiae. This hypothesis is refuted by the characters which
corroborate the monophyletic nature of the nottii species group.

Fundulus lineolatus has an aggregation of melanophores on
the upper part of the pectoral fin base while F. nottii and F.

24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

escamhiae have a cleithial stripe. Fundulus dispar and F. blairae
have a few scattered and unfused melanophores on the pectoral fin
base, but no concentration Hke that seen in F. lincolatus. I inter-
pret the partial cleithral stripe seen in F. lUieolatus as intemiediate
between dispar and blairae (no stripe) and the fully developed
cleithral stripe of nottii and escamhiae. I suggest that this char-
acter provides additional corroboration for the relationship of
lineolatus with nottii and escamhiae rather than dispar and hlairae.
Fundulus nottii and F. escamhiae are hypothesized sister spe-
cies based on their common possession of a cleithral stripe and
their modal tendency for fused pores 4a and 4b in the supraorl)ital
sensory canal.

Examination of several large series of each species at the Uni-
versity of Michigan Museum of Zoology by Dr. Gerald R. Smith
showed that the head squamation patterns discussed by Wiley and
Hall (1975) are subject to the variation defined in the Methods and
Materials section and discussed for the various species of tlie
nottii species group. Fundulus dispar has, modally, a G-t)'pe pat-
tern, but individuals also show tlie E-G variation found in F.
lineolatus, F. nottii and F. escamhiae (who have a modal E-type
pattern). G-type patterns predominate in both F. hlairae and F.
dispar but are rare in F. lineolatus (2 of 77 specimens, UMMZ
110662 and 166605), F. nottii (3 of 62 specimens, UMMZ 163698),
and F. escamhiae (6 of 81 specimens, UMMZ 163425). I ascribe
this variation to ontogenetic plasticity and suggest that the modal,
rather than invariant, nature of the characters does not detract
from their usefulness as phylogenetic indicators. The same reason-
ing is apphed here to the variation on the fusion of pores 4a and
4b of the supraorbital sensory canals.

Discussion

The resurrection of F. escamhiae brings the nmuber of species
of the F. nottii species group to five. Fundulus escamhiae is the
most polymorphic species of the complex, displaying color pattern
variation over the relatively short extent of its range. The sig-
nificance of this variation is unknown and populational studies
utilizing more samples would be useful. Fundulus escamhiae is
recognized here for the same reasons F. hlairae was recognized by
Wiley and Hall (1975): (1) F. nottii and F. escamhiae are diag-
nosable at the 100% level without recourse to geographic data;
(2) neither species shows evidence of color pattern intergradation
in contiguous populations (e.g., the Mobile Bay drainage F. nottii
and the Styx-Perdido drainages F. escamhiae); and (3) the phy-
logenetic and biogeographic relationships of the two species with
other members of the group are not incompatible with their
specific status.

FUNDULUS NOTTII SPECIES GROUP

Fig. 10. — Least rejected phylogenetic hypothesis of the interrelationships
of the species of the Fundulus nottii species group. Black rectangles are syn-
apomorphies uniting sister groups: (1) a subocular teardrop present; (2) fe-
males with numerous flank stripes between the scale rows; (3) pores 4a and
4b of the supraorbital sensory canal not widely separated; (4) a distinct pig-
ment spot and the pectoral fin base; (5) pores 4a and 4b of the supraorbital
sensory canal usually fused in adults; (6) a cleithral stripe present; (7) usually
with G-type head squamation; (8) no vertical bars on the flank of females.

Fundulus hlairae and F. dispar occur farther east than reported
by either Brown (1958) or Wiley and Hall (1975). The F. dispar
from the Tombigbee River drainage (TU 76844) are unquestion-
ably of that species and show no signs of intergradation of color
pattern. I was unable to find any collections of F. dispar between
the Tombigbee and Mississippi rivers. This disjunction may reflect
stream capture between the eastern tributaries of the Mississippi
River drainage and the Mobile Bay drainage as discussed by Smith-
Vaniz (1968) or simply a lack of collections. Fundulus hlairae
occurs in the Tensas, Pearl, Pascagoula, and Mobile Bay drainages.

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Although they both occur in the Mobile drainage, F. dispar is
apparently found upstream while F. hlairae is found downstream.
This is basically the same association found in the Mississippi
River drainage, with F. hlairae being found in the lower reaches
( Red River and southward ) and F. dispar being found farther up-
stream (Ouachita River and northward). The distributional rela-
tionship of the two species in the Tombigbee and Alabama rivers
is unknown because of a lack of collections.

Chen's ( 1971 ) karyotype analysis of F. nottii and "F. lineolatus"
showed no differences between the two species. However, Dr.
Michael Stevenson (Univ. of New Orleans) has informed me that
Chen's "F. lineolatus" came from the range of F. escambiae. Setzer's
(1968) karyotype of "F. notti" was taken from a specimen from
the Neches River drainage and is undoubtably F. hlairae. The
fact that Setzer's karyotype of F. hlairae is different than Chen's
karyotypes of F. nottii and F. escambiae suggests that other karyo-
typic data may be useful for indicating relationsliips within the
complex.

Color in life may also provide additional evidence for relation-
ships. Several small series of each species were collected by Michael
M. Stevenson and I. The F. lineolatus, F. nottii, and F. escambiae
collected had red or orange-red pigment on the side of the head
between the orbit and the mouth whereas the F. dispar and F.
hlairae examined lacked this pigment. In addition, the three former
species have a generally brown or blackish background color and
a single light spot on the head whereas the latter two have a
greenish background coloration and several hght spots on the
head. These characters were not incorporated into the phylo-
genetic analysis because they were observed in only the few live
specimens examined for each species. Workers collecting new
material might examine color characteristics in life for possible
phylogenetic significance.

The distributional patterns of the F. nottii species group repre-
sent a little-altered vicariance pattern (as defined by Croizat, Nel-
son, and Rosen, 1974). Sister pairs such as F. nottii-F. escambiae
and F. dispar-F. hlairae are allopatric and represent unaltered vi-
cariance. This type of distributional pattern is similar to the pat-
tern for closely related species of Xiphophorus platyfishes (as
viewed by Rosen, 1960, and discussed by Croizat et al., 1974).
Sympatry within the species group only occurs among non-sister
groups such as F. hlairae and F. nottii, or F. escambiae and F.
lineolatus. These patterns of sympatry may be interpreted as al-
terations of past vicariance events by dispersal of one or both
sympatric species; but which dispersed and in what direction re-
mains problematical. I suggest the following hierarchy of vicari-
ence events based on the phylogenetic relationships (Fig. 10) and
distributional patterns of the species (Fig. 9). (1) The ancestor of

FUNDULUS NOTTIl SPECIES GROUP 27

the species group was split into the ancestral species of F. dispar
and F. blairae on the one hand and the ancestral species of F.
lineolatus, F. nottii, and F. escamhiae on the other hand. ( 2 ) Fun-
dulus dispar and F. blairae speciated. ( 3 ) Fundulus lineolatus and
the ancestor of F. nottii and F. escamhiae speciated. (4) The
ancestor of F. nottii and F. escamhiae speciated and the vicariance
event which was involved may have been the late Pliocene or
early Pleistocene uplift of the Chattahoochee Arch which sep-
arated the Perdido and other more eastern river systems along the
Gulf coast from the Alabama-Tombigbee drainage (Price and
Whetstone, in press). Vicariance event (2) certainly followed
(1), but whether it was coincident with or time independent of
(3) or (4) can not be determined without a general track analysis
of the biota. Lacking this, most of the vicariance events can be
only indirectly inferred from the phylogenetic hypothesis.

Acknowledgments

A part of this work was submitted as a Masters Thesis, Sam
Houston State University. Later work was done at the American
Museum of Natural History and the Museum of Natural History,
University of Kansas. My thanks go to all institutions for providing
financial and logistical assistance. Drs. Darrell D. Hall (Sam
Houston State University) and Donn E. Rosen (American Mu-
seum of Natural History) provided help and encouragement
throughout the project. Dr. Frank B. Cross (University of Kansas)
and Dr. Camm Swift (Los Angeles County Museum) read and
suggested improvements of the manuscript. Dr. Gerald R. Smith
(University of Michigan, Museum of Zoology) brought the head
squamation and sensory pore variation to my attention. Ms. Deb-
ora Bennett drew figures 1 and 2. The following persons and
institutions are thanked for loaning specimens or for welcoming
me during visits: Dr. Reeve M. Bailey (University of Michigan
Museum of Zoology); Dr. Neil H. Douglas (Northwestern Louisi-
ana State University); Mr. John Heizer (Cornell University); Dr.
Henry W. Robison (Southern State College); Dr. Royal D. Suttkus
(Tulane University); Dr. Stanley H. Weitzman (U.S. National
Museum of Natural History).

Summary

(1) The Fundulus nottii species group consists of five North
American freshwater cyprinodontids distinguished from other
species of Fundulus by the presence of a subocular teardrop in
both sexes of all ages and by the presence of numerous stripes
between the flank scales of females only. These species and their

28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

ranges are: F. nottii (Lake Pontchartrain, Louisiana, drainage east
to the lower Mobile Bay drainage Alabama); F. escamhiae (Styx
River, Alabama, east to the Suwannee River drainage, Florida);
F. Uneolatus (Ocklockonee River drainage, Florida, south to Dade
Co., Florida, north and east to southern Virginia); F. dispar (Oua-
chita River drainage, Louisiana, north to southern Michigan, east to
the upper Mobile Bay drainage, Alabama); and F. blairae (Brazos
River drainage, Texas, north to the Red River drainage, southeastern
Oklahoma, east to the lower Mobile Bay drainage, Alabama). A
key to the species is provided for identification.

(2) Synonymies, diagnoses, and re-descriptions are given for
all species except the recently described F. blairae for which a
synonymy and diagnosis are given.

(3) A revised phylogenetic analysis indicates two monophy-
letic groups within the complex: the blairae-dispar species pair
and the escambiae-nottii-lineolatus species group. Fundulus
blairae is the sister species of F. dispar based on head squamation
pattern and female color pattern. Fundulus escambiae and F.
nottii are sister species based on supraorbital sensory pore pattern
and color pattern. Fundulus Uneolatus is more closely related to
F. nottii and F. escambiae than to F. blairae and F. dispar based
on supraorbital sensory pore pattern.

(4) A brief discussion of the biogeographic patterns of the
species group is included.

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