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2011 SOUTHEASTERN NATURALIST 10(1):133–144
Novel Phylogeographic Patterns in a Lowland Fish,
Etheostoma proeliare (Percidae)
Nicholas J. Lang1,2,* and Anthony A. Echelle1
Abstract - Etheostoma proeliare is distributed in the Gulf Coastal Plain of southeastern
North America from the Colorado River of eastern Texas through the Choctawhatchee
River of eastern Alabama, north to the Fall Line, and upstream along the Arkansas River
valley into eastern Oklahoma. Parsimony and Bayesian analysis of the ND2 gene from
28 populations recovered a monophyletic E. proeliare (PP:1.0) containing a basal split
between a novel clade of populations in the northern apex of the Mississippi embayment,
from the Black and St. Francis rivers to the west of, through the Yazoo River to the east
of, the main channel of the Mississippi River, and all other populations (PB:99, PP:1.0).
Southern populations are resolved into three clades: Trinity plus Neches Rivers (PB:100,
PP:1.0), Sabine River east through western tributaries to the Mississippi River plus the
Lake Pontchartrain drainage (PP:1.0), and eastern tributaries to the Mississippi River
east through the Escambia River (PP:0.99). The eastern clade is further divided into Pearl
plus Big Black rivers (PB:97, PP:1.0) and Mobile Basin plus Escambia River clades
(PP:0.98). These results indicate that although the mainstem of the Mississippi River
corresponds to some phylogeographic breaks in a lowland species, it is not an absolute
barrier. Future studies of species in the region should further explore the placement of
Lake Pontchartrain populations and the existence of divergent populations in the northern
Gulf Coastal Plain.
The North American temperate freshwater ichthyofauna is relatively rich
compared to that of temperate Eurasia (Berra 2001). Most species are found
east of the Continental Divide, where a large portion of the diversity is found in
two clades, the minnows (Cyprinidae) and darters (Percidae) (Lee et al. 1980).
Both clades consist of sympatrically distributed but ecologically divergent lineages
(e.g., subgenera of Etheostoma in darters [Page and Swofford 1984]) that
have given rise to multiple allopatric species across the region’s topographically
diverse and temporally plastic drainages (e.g., members of the Notropis
rubellus (Agassiz) (Rosyface Shiner) species group [Berendzen et al. 2008]).
Endemism is highest within the Ozark, Ouachita, and Eastern Highlands, three
disjunct regions of relatively high relief drained by tributaries to the Mississippi
River. Most biogeographic studies of North American fishes have focused on
species or clades found in one or more of these regions (Mayden 1988, Wiley
and Mayden 1985).
1Department of Biology, Oklahoma State University, Stillwater, OK. 2Current address
- Division of Fishes, Department of Zoology, Field Museum of Natural History, 1400
South Lake Shore Drive, Chicago, IL 60605-2496. *Corresponding author - oligocephalus@
134 Southeastern Naturalist Vol. 10, No. 1
The ichthyofauna of the Mississippi River system is especially diverse because
its tributaries encompass a wide variety of habitat types in which highland
endemics are isolated by unsuitable lowland and/or big river habitat. Within
linear freshwater systems, the presence of intervening unsuitable habitat can
severely limit gene flow among populations, and processes such as long-term
environmental change or drainage rearrangement may be required for the establishment
of new populations (Echelle et al. 1975).
In lowland areas, the relatively modest topographic relief between drainage
systems, along with the presence of connecting lakes and/or deltas, may allow
for greater connectivity among populations in separate drainages over time. Although
many lowland freshwater fishes are widespread, biogeographic breaks do
still occur (Bermingham and Avise 1986). In the lowland area of southeastern
North America known as the Gulf Coastal Plain (GCP), the major biogeographic
break is coincident with the channel of the Mississippi River (Robison 1986).
Although some species are found on both sides of the Mississippi River, widespread
clades generally contain distinct species on either side (Birdsong and
Knapp 1969, Lee at al. 1980, Pramuk et al. 2007). A second biogeographic break
delineates distinct faunal assemblages west of, within, and east of the Mobile Basin
(Swift et al. 1986). Although populations within the GCP have been included
in molecular studies on the phylogeography of both lowland (Bermingham and
Avise 1986) and highland (Berendzen et al. 2003, Ray et al. 2006) fishes, no study
has explored the phylogenetic structuring of a small stream species endemic to,
and distributed across, the entire region.
Etheostoma proeliare (Hay) (Cypress Darter), is found in lowland portions
of GCP drainages from the Colorado River, TX (Auburn University Museum
26406) through the Choctawhatchee River, AL and fl(Burr 1978). Although
largely restricted to the area below the Fall Line, which demarcates the border
between upland regions and the GCP, the species crosses this boundary
in transitional zones in southeastern Missouri and southern Illinois and along
the Arkansas River valley upstream to eastern Oklahoma (Fig. 1). Cypress
Darters can be found in the vegetated margins of swamps and lakes as well
as near cover in slow, sand/silt-bottomed streams (Burr and Page 1978). Burr
(1978) concluded, based on morphological data, that the Cypress Darter is the
sister species of E. fonticola (Jordan and Gilbert) (Fountain Darter), which is
endemic to springs of the Guadalupe River drainage, just west of the range of
the Cypress Darter. Although Burr (1978) found variation among populations
of the Cypress Darter, he considered the variation to be randomly distributed
and not significant enough to warrant taxonomic recognition. The sister relationship
between the Cypress Darter and the Fountain Darter was further
supported by allozyme data in a study that found little divergence among
Cypress Darter populations east of, but did not include any populations west
of, the Mississippi River (Buth et al. 1980). Herein, we present the results of
a phylogeographic study of the Cypress Darter and discuss our results in the
broader context of Gulf Coastal Plain biogeography.
2011 N.J. Lang and A.A. Echelle 135
Materials and Methods
Specimens were collected by seine, dipnet, and/or backpack electroshocker.
Whole specimens were either frozen or preserved in 95% ethanol, or fin clips were
placed in 95% ethanol and voucher specimens preserved in formalin. Specimens
and tissues were accessioned into the collections at the University of Alabama and
Saint Louis University, or are held at Oklahoma State University (Appendix 1).
Attempts were made to collect specimens from all major drainages throughout the
range, but failed in the Colorado, Choctawatchee, and Pascagoula rivers, where the
Cypress Darter is known from one, two, and fewer than ten localities, respectively.
Ingroup sampling includes one specimen per locality from at least one locality
in all other major drainages within the range of the Cypress Darter. Because uncertainty
regarding the relationships among subgenera of Etheostoma makes it
Figure 1. Distribution of the Cypress Darter indicated with shading. The heavy dashed
line indicates the approximate position of the Fall Line, the white lines within the northern
portion of the shaded distribution indicate the approximate position of Crowley’s
Ridge (after Robison 1986), and hollow symbols represent approximate Cypress Darter
sampling localities. Symbols are clade-specific and the numbers correspond to populations
within each clade.
136 Southeastern Naturalist Vol. 10, No. 1
difficult to choose a darter outgroup for this study, the dataset includes all ND2
sequences from Lang and Mayden (2007) (EF027169 through EF027233) plus one
Fountain Darter, which occurs naturally in a single spring system.
Whole DNA was extracted using either a standard phenol-chloroform method or
the DNEasy Kit (QIAGEN, Valencia, CA). The complete mitochondrial ND2 gene
was amplified (35 cycles of 94 °C for 40 sec, 56 °C for 60 sec, and 72 °C for 90 sec)
with Taq DNA Polymerase (PROMEGA, Madison, WI) using the external primers
of Lang and Mayden (2007). Amplification products were gel extracted using a Gel
Extraction Kit (QIAGEN, Valencia, CA). Sequencing utilized both the external
amplification primers and Cypress Darter-specific internal primers (Epro539L:
5-ACTCATCCATCGCCCACCTT-3’, Epro739H: 5’-AGACCTCCTAATGAAAGAAG-
3’) with chemistry specific to various visualization systems. Products were
visualized using either the CEQ 8000 Genetic Analysis System (Department of
Biology, Saint Louis University, St. Louis, MO) or an ABI 3700 (Auburn University
Genomics Laboratory, Auburn, AL, Oklahoma State University Recombinant
DNA Core Facility, Stillwater, OK, and Macrogen, Inc., Seoul, Korea). Sequence
files were edited, contigs were assembled, and sequences were aligned by eye using
Geneious Pro v. 3.5.6 (Drummond et al. 2007).
Maximum Parsimony analyses were implemented in PAUP*4.0b10 (Swofford
2002), with 100 repetitions of random stepwise addition and TBR branch
swapping, with Perca flavescens (Mitchill) (Yellow Perch) and Sander marinus
(Cuvier) (Estuarine Perch) designated as outgroups. Support for nodes (PB: parsimony
bootstrap) was estimated using 1000 “fast-stepwise” addition replicates
of bootstrapping. MrModelTest 2.2.1 for Classic (Nylander 2004) was used to
hypothesize the best-fit model for codon-position specific partitions of the dataset.
These models were used in two independent mixed-model Bayesian analyses
in MrBayes3.1 (Ronquist and Huelsenbeck 2003) that ran for 5,000,000 generations.
Priors for these analyses were flat, four chains were utilized, and trees were
sampled every 100 generations. Yellow Perch was designated as the outgroup for
Bayesian analyses, which only allows a single outgroup. Stationarity of negative
log-likelihood values was evaluated by plotting these values against generation,
and all trees before the value stabilized were discarded as burn-in. Support for
recovered nodes (BPP: Bayesian posterior probability) was calculated by creating
a majority-rule consensus tree using all post-burn-in trees for each analysis
and averaging the resulting values.
The data matrix consists of 1047 base pairs. Sequences were generated for 29
specimens of Cypress Darter from 28 localities. The following samples are based
on incomplete data: Ol-821, missing bp 1–537; Ol-832, missing bp 418–537;
Ol-1043, -1044, -1045, -1046, and -1047, missing bp 1028–1047. Each sampled
locality yielded a unique haplotype.
There were 454 constant and 532 parsimony-informative characters, and
maximum parsimony analysis yielded two most-parsimonious trees (5807 steps).
MrModelTest selected the GTR+I+G model for the second and third codon
2011 N.J. Lang and A.A. Echelle 137
positions. For the first position, the GTR+I+G and SYM+I+G models were each
selected by a single likelihood ratio test (LRT), but the remaining LRT and the
Akaike information criterion selected the HKY+I+G model, which was used in
the analyses. The burn-in period for both Bayesian analyses was designated as
the first 50,000 generations, and average BPP values were calculated from those
in the run-specific majority-rule consensuses of trees 52–5001.
The results of the parsimony and Bayesian results were highly concordant,
disagreeing only on the resolution of nodes with little support, and the amount of
support for consistently recovered nodes. Relationships were identical to those in
Lang and Mayden (2007) except for the addition of a strongly supported sisterspecies
relationship (PB: 100, BPP: 1.0) between the Fountain Darter and the
monophyletic Cypress Darter (PB: *, BPP: 1.0; not shown).
Among populations of Cypress Darter, there is a basal split between those from
the northern part of the range and all others (Fig. 2). This northern clade (PB: 99,
Figure 2. One of two most parsimonious phylograms of the relationships among populations
of Etheostoma proeliare recovered in our analysis. Numbers above nodes are
parsimony bootstrap values and those below are Bayesian posterior probability values.
Nodes found in both most-parsimonious trees, but not receiving bootstrap support, are indicated
by an asterisk. Symbols and population numbers correspond to those in Figure 1.
138 Southeastern Naturalist Vol. 10, No. 1
BPP: 1.0, circles in Fig. 1) encompasses all sampled populations from streams
north of and including the St. Francis River to the west, and the Yazoo River to the
east, of the main channel of the Mississippi River, as well as the upper Black River
in southeastern Missouri. Although the Black River is a tributary to the White
River, the population sampled from the lower White River (Tarleton Creek) is not a
member of the northern clade. The clade encompassing the remaining populations
(PB: *, BPP: 1.0) comprises three well-supported clades distributed as follows:
1. east of the Mississippi River except the Lake Pontchartrain drainage (PB: *,
BPP: 0.99, squares in Fig. 1); 2. the Neches and Trinity river drainages (PB: 100,
BPP: 1.0, stars in Fig. 1); and 3. the Red River drainage, the Lake Pontchartain
drainage, and the Sabine River drainage, exclusive of the Neches River (PB: *,
BPP: 1.0, triangles in Fig. 1). The clade of populations east of the Mississippi River
is divided into two well-supported clades: one comprised of populations from the
Mobile Basin and Escambia River (PB: *, BPP: 0.98) and the other of populations
from the Pearl and Big Black rivers (PB: 97, BPP: 1.0). Percent divergences within
and among these major clades are presented in Table 1.
Within the clade comprised of populations from the Lake Pontchartrain drainage
and those drainages between the Neches and Mississippi rivers, there are a
number of well-supported sub-clades. Single-drainage clades comprise the populations
from the Lake Pontchartrain (PB:98, BPP: 1.0) and lowland Red River
(PB: 100, BPP: 1.0) drainages, and the sampled population from the Sabine River
was recovered in a clade with that from the neighboring Calcasieu River (PB:
92, BPP: 1.0). A population from the lower White River drainage was recovered
as sister to the lowland Red River drainage clade (BPP: 1.0), and a clade uniting
populations from the middle Arkansas (Canadian River) and upland Red (Yanubbe
Creek) river drainages was also recovered (PB: 91, BPP: 1.0).
Our well-supported phylogenetic hypothesis of relationships among regional
populations of the Cypress Darter suggests several novel biogeographic patterns.
The most interesting of these is the resolution of a distinct clade comprised of
drainages in the northern apex of the GCP. West of the main channel of the Mississippi
River, the southern limit of this northern clade (circles in Fig. 1) is found
among tributaries to the Black River (White River drainage) and the St. Francis
River. The sampled individual from the lowland portion of the White River
drainage is in the Red River/Lake Pontchartrain clade (triangles in Fig. 1). Most
Table 1. Uncorrected mean percent sequence divergences within (upper diagonal) and between
(below diagonal) major clades identified among Cypress Darter populations. Symbol designations
correspond to those in Figures 1 and 2.
Circles Triangles Squares Stars
Northern clade (circles) 1.51%
Red River/Lake Pontchartrain/Sabine River clade (triangles) 5.57% 1.70%
Eastern clade (squares) 5.39% 3.06% 2.38%
Trinity and Neches rivers clade (stars) 5.17% 2.90% 3.03% 0.30%
2011 N.J. Lang and A.A. Echelle 139
populations recovered in the northern clade are from transitional areas between
the lowland GCP and the Ozark or Eastern Highlands. Many highland species
are shared between the upland portions of the St. Francis and Black (including
Current and Spring rivers) river drainages, and the Cypress Darter is abundant
in the Missouri portion of the Black River drainage at the transition between the
GCP and the Ozark Highlands (Pflieger 1997, Robison and Buchanan 1988). It
is possible that the northern clade gained access to this transitional area of the
upland Black River drainage via former outlets that flowed east through the current
St. Francis River drainage rather than south to the White River drainage. The
route of the main channel of the Mississippi River has varied over time, and the
present-day configuration of drainages in the northern GCP is likely too recent to
explain the distribution of many species (Robison 1986).
The presence of haplotypes from the northern clade so far downstream in the
St. Francis River drainage may be due to the presence of Crowley’s Ridge, an isolated
highland geological feature in the GCP that runs along the western edge of
the lower St. Francis River drainage, and is drained by the sampled creek. It has
long been recognized that streams draining the ridge are home to isolated populations
of highland fishes such as Campostoma anomalum (Rafinesque) (Central
Stoneroller), Etheostoma caeruleum Storer (Rainbow Darter), and Chrosomus
erythrogaster (Rafinesque) (Southern Redbelly Dace) (Robison and Buchanan
1988), but our data indicate that the ridge may also harbor cryptic biodiversity
within typically lowland species. Future work on the Cypress Darter should incorporate
specimens from farther upstream in the White River drainage and from
streams in the St. Francis River drainage that do not drain Crowley’s Ridge.
East of the main channel of the Mississippi River, the northern clade extends
as far south as the Yazoo River. Although there are no clear biogeographic breaks
between the Big Black and Yazoo rivers in widespread lowland species, the
Yazoo River is the southern limit of Snubnose Darters (Etheostoma, subgenus
Ulocentra). Also, several species, such as Cyprinella whipplei Girard (Blacktail
Shiner), Lythrurus roseipinnis (Hay) (Cherryfin Shiner), and Notropis texanus
(Girard) (Weed Shiner), are common on one side of the Big Black/Yazoo divide,
but rare on the other (Ross 2001). Within Noturus hildebrandi (Bailey and Taylor)
(Least Madtom), the subspecies N. h. hildebrandi is restricted to streams
from the Big Black River south, and N. h. lautus is found from the Hatchie River
north, while populations in the Yazoo River were considered intergrades (Taylor
1969). It is possible that, as in the White River drainage, members of the northern
clade are restricted to relatively upland headwater habitats, where the Cypress
Darter is distinctly more abundant, in the Yazoo River and drainages in western
Tennessee (Etnier and Starnes 1993, Ross 2001). Our samples came from small
streams in the relative headwaters of these drainages, and future work should
include samples from more lowland areas in eastern tributaries to the Mississippi
River in order to more fully explore the range of the northern clade.
Within the southern clade, we recovered both predicted and novel relationships.
The split between the Mobile Basin and GCP drainages between the
Mobile Basin and the main channel of the Mississippi River is predicted by both
140 Southeastern Naturalist Vol. 10, No. 1
the composition of the respective faunas (Swift et al. 1986) and the distribution
of endemic species, such as those in the Notropis dorsalis (Agassiz) (Bigmouth
Shiner) species group (Raley and Wood 2001) and the southern clade of Lythrurus
(sensu Pramuk et al. 2007). The recovery of an unresolved trichotomy among
the Mobile Basin drainages and the Escambia River drainage may be an artifact
of our sampling, which does not include the Choctawhatchee River population.
Although a phylogeographic break at the mainstem Mississippi River was
predicted based on the inability of small-stream fishes to utilize its large-stream
habitat (Robison 1986), the recovery of Lake Pontchartrain drainage populations
with populations west of the main channel of the Mississippi River was novel. It
is difficult to hypothesize a connection between the Lake Pontchartrain drainage
and those of the western GCP because the basin that became Lake Pontchartrain
roughly 4000 years ago has always been east of the Mississippi River delta (Frazier
1967). It is tempting to suggest that a lowland species, such as the Cypress
Darter, may be able to utilize the dynamic environment of a large delta to facilitate
crossing a large barrier such as the main channel of the Mississippi River.
However, the current distribution of the Cypress Darter indicates that it does not
inhabit such regions in either the Mississippi River or Mobile Basin deltas (Boschung
and Mayden 2004, Douglas 1974).
The Cypress Darter’s possible aversion to delta conditions may also explain
the resolution of a clade composed of populations from the Neches and Trinity
rivers to the exclusion of the Sabine River drainage. Conner and Suttkus (1986)
described a fairly uniform fauna within what Knapp (1953) referred to as the
Eastern Area of Texas, with no distinction between the fauna of the Neches
and Sabine river drainages. It is possible that the distinct Neches/Trinity clade
evolved in eastern Texas prior to integration of the Neches and Sabine rivers
into a single outlet, and that uninhabitable delta habitat has kept the populations
separated. Given the presence of the Fountain Darter on the western periphery of
the range of the Cypress Darter, it is possible that the western edge of the GCP
has long harbored significant genetic variation within the clade comprising both
species, as was recently suggested for the clade comprised of Percina apristis
(Hubbs and Hubbs) (Guadalupe Darter) and P. sciera (Swain) (Dusky Darter)
(Robins and Page 2007). Inclusion of the recently discovered Cypress Darter
population from the intervening Colorado River drainage should be a goal of
future analyses. The remaining relationships among sampled populations west
of the main channel of the Mississippi River are largely unresolved and will require
finer scale sampling if phylogeographic structure is to be resolved within
or among the Red and Arkansas river drainages.
Although this analysis is based on a single mitochondrial gene, there is no
evidence of hybridization, which is often readily apparent when using mitochondrial
data due to the sharing of identical, or very similar, sequences among species
(Ray et al. 2008), and the well-supported clades are not only significantly
divergent (Table 1) but also geographically restricted. Reinterpretation of Burr’s
(1978) morphological data in light of our results is difficult due to the presentation
of statistical measures rather than raw data for some characters and the
2011 N.J. Lang and A.A. Echelle 141
grouping of populations from drainages containing clade boundaries (e.g., Burr
 grouped data from the Sabine and Neches Rivers and presented data from
the Black River drainage as part of the White River). It does not appear, however,
that the distributions of our novel clades conform to any of the patterns he identified. Discussion of cryptic speciation within the Cypress Darter will require finer
sampling of genetic and possibly ecological data. Finally, although the novel
biogeographic aspects of our results are difficult to interpret without additional
data from co-distributed species, our analyses clearly indicate that significant
phylogeographic structure can exist in widespread Gulf Coastal Plain fishes.
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(030720073 and 052720082), Illinois (A07.5059), Kentucky (SC0711079), Louisiana
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144 Southeastern Naturalist Vol. 10, No. 1
Appendix 1: Material Examined. Institutional abbreviations follow Leviton et al. (1985),
except STL, which indicates the Ichthyological Tissue Collection of Saint Louis University,
Each record indicates State: Catalog Number (individual DNA extraction identification
number, GenBank accession number), Locality (Drainage), County.
Texas: STL 949.01 (Ol-0114, HM595746), San Marcos River, Hays.
Alabama: UAIC 12411.08 (Ol-0832, HM593884), Limestone Creek (Alabama River),
Monroe; UAIC 13418.08 (Ol-0885, HM593889), Conecuh River, Escambia; UAIC
15590.01 (Ol-1165, HM593872), Wards Mill Creek (Tombigbee River), Tuscaloosa.
Arkansas: STL 413.01 (Ol-0826, HM593880), Bridge Creek (Arkansas River), Faulkner;
STL 427.08 (Ol-0830, HM593883), Tarleton Creek (White River), Arkansas; STL
428.01 (Ol-0827, HM593881), Tuni Creek (St. Francis River), St. Francis. Illinois:
UAIC 13100.12 (Ol-0890, HM593891), Max Creek (Ohio River), Johnson. Kentucky:
OSUS 27599 (Ol-0873, HM593887), Richland Creek (Cumberland River), Livingston.
Louisiana: STL 94.05 (Ol-0076, EF027214), Natalbany River (Lake Pontchartrain),
Tangipahoa; STL 326.07 (Ol-0822, HM593876), unnamed creek (Calcasieu River), Vernon;
STL 331.03 (Ol-0821, HM593875), Clark Creek (Ouachita River), Ouachita; STL
669.09 (Ol-0825, HM593879), unnamed creek (Red River), Rapides; UAIC 15185.01
(Ol-1043, HM593896), Little Natalbany River (Lake Pontchartrain), St. Helena; UAIC
15191.01 (Ol-1044, HM593897), Old River (Sabine River), Beauregard. Mississippi:
OSUS 27600 (Ol-914, HM593893), unnamed stream (Yazoo River), Lafayette; OSUS
27601 (Ol-915, HM593894), Besa Chitto Creek (Pearl River), Choctaw; OSUS 27602
(Ol-916, HM593895), Bala Chitto Creek (Lake Pontchartrain), Pike; STL 870.10 (Ol-
0820, HM593874), Poplar Creek (Big Black River), Choctaw. Missouri: STL 380.04
(Ol-0889, HM593890), unnamed ditch (Black River), Butler; STL 663.04 (Ol-0828,
HM593882), Ditch Number 10 (Mississippi River), New Madrid. Oklahoma: STL 277.02
(Ol-0834, HM593885), Yanubbee Creek (Red River), McCurtain; STL 288.04 (Ol-0824,
HM593878), tributary to Gaines Creek (Canadian River), Latimer. Tennessee: OSUS
27597 (Ol-0877, HM593888), Cash Creek (Hatchie River), Hardeman; OSUS 27598
(Ol-0893, HM593892), Big Sandy River (Tennessee River), Henderson; UAIC 15216.02
(Ol-1047, HM593900), Kings Branch (Tennessee River), Hardin. Texas: STL 323.03
(Ol-0823, 593877; -0844, HM593886), Little Cypress Bayou (Red River), Upshur; UAIC
15195.06 (Ol-1045, HM593898), Hickory Creek (Neches River), Polk; UAIC 15196.02
(Ol-1046, HM593899), Bluff Creek (Trinity River), Polk.