Designation of a neotype for brook trout, Salvelinus fontinalis

Abstract

The taxonomic status of Salvelinus fontinalis (Mitchill) is problematic. Difficulties in comparison of populations are exacerbated by the lack of type material. Here we designate a neotype from Connetquot River, Long Island, New York. We provide genetic and morphological data for the neotype, conspecifics, and other populations (Swan Creek, Nissequogue Creek) from Long Island, New York. We demonstrate, using molecular markers, that the population from Connetquot River most likely has not been influenced by the major broodstock strains utilized in the Northeast for supplemental and restorative stocking programs. We distinguish the above populations morphologically from lower interior basin populations, represented by fishes from the Pigeon-French Broad drainage, North Carolina and Tennessee. Finally, we position populations from Long Island, New York, within six distinct lineages of S. fontinalis.

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Designation of a neotype for brook trout, Salvelinus fontinalis
Author(s): Jay R. Stauffer, Jr. and Timothy L. King
Source: Proceedings of the Biological Society of Washington, 127(4):557-567.
2015.
Published By: Biological Society of Washington
DOI: http://dx.doi.org/10.2988/0006-324X-127.4.557
URL: http://www.bioone.org/doi/full/10.2988/0006-324X-127.4.557
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Designation of a neotype for brook trout, Salvelinus fontinalis
Jay R. Stauffer, Jr.* and Timothy L. King
(JRS) Ecosystem Science and Management, The Pennsylvania State University, University Park,
Pennsylvania 16802, U.S.A., e-mail: vc5@psu.edu;
(TLK) U.S. Geological Survey, Leetown Science Center, 11649 Leetown Road, Kearneysville,
West Virginia 25430, U.S.A.
Abstract.—The taxonomic status of Salvelinus fontinalis (Mitchill) is
problematic. Difficulties in comparison of populations are exacerbated by
the lack of type material. Here we designate a neotype from Connetquot
River, Long Island, New York. We provide genetic and morphological data
for the neotype, conspecifics, and other populations (Swan Creek, Nisse-
quogue Creek) from Long Island, New York. We demonstrate, using
molecular markers, that the population from Connetquot River most likely
has not been influenced by the major broodstock strains utilized in the
Northeast for supplemental and restorative stocking programs. We distin-
guish the above populations morphologically from lower interior basin
populations, represented by fishes from the Pigeon-French Broad drainage,
North Carolina and Tennessee. Finally, we position populations from Long
Island, New York, within six distinct lineages of S. fontinalis.
Keywords: brook trout, neotype, New York, Salvelinus fontinalis
The brook trout, Salvelinus fontinalis
(Mitchill), evolved a great variety of life
history, developmental and physiological
traits in response to a broad range of
habitats. Populations of brook trout are
native to headwater streams and cold-
water lakes of the Mississippi River
drainage east to the Atlantic Slope drain-
ages; they are found from northeastern
Canada south through the Great Lakes
and into the southern Appalachian moun-
tains (Power 1980). The geological history
of this region reveals numerous events
(Hocutt et al. 1986) that would serve to
isolate fish populations, including exten-
sive glacial impoundment and stream
captures in the northern parts of its range,
and montane glaciations and stream cap-
tures affecting southern populations.
Brook trout, along with most other
salmonids, show exceptional levels of life-
history variation (e.g., resident and migra-
tory types often co-occur). Upon reaching
sexual maturity individuals undergo dra-
matic morphological (e.g., male kype
formation), physiological, and behavioral
adaptations. This life-history variation
appears to be influenced by complex
interactions between genetic and environ-
mental factors (Hendry et al. 2004).
Anthropogenic disturbances and climate
change have resulted in the proliferation of
demographically small isolated popula-
tions, thus, providing the opportunity to
study the accumulation of phenetic and
genetic traits in allopatric populations.
Additionally, the overlapping transloca-
tion of isolated populations into new areas
permits the study of reproductive isolation
of introduced populations in natural envi-
ronments.
Salvelinus fontinalis was described from
Long Island, New York (Mitchill 1814).
Unfortunately, neither type material nor a
* Corresponding author.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
127(4):557–567. 2014.

type locality were designated. Mitchill
(1815:438) further described the brook
trout as a ‘‘most dainty fish’’ that lives in
running waters; however, he also reported
fish that weighed in excess of two kg.
Certainly, there were both resident and
sea-run individuals present on Long
Island during this time. Behnke (1980)
posed the question of whether there were
distinct northern and southern groups of
brook trout, or a single homogenous
stock established since the last glacial
retreat during the Pleistocene. Morgan &
Danzmann (1997, 2001) and Hall et al.
(2002) suggested high mtDNA RFLP
diversity within, and differentiation
among, brook trout populations in the
mid-Atlantic when compared to northern
populations analyzed previously (Jones et
al. 1997, Danzmann et al. 1998). By
contrast, brook trout from the mid-
Atlantic region belong to five of the six
established mtDNA assemblages (Mor-
gan & Danzmann 1997, 2001; Danzmann
et al. 1998, Hall et al. 2002). Additionally,
most allozyme-based studies of popula-
tions from the southern Appalachians
have produced evidence that genetic
diversity is relatively high in this region
of the native range of brook trout
(McCracken et al. 1993, Hayes et al.
1996, Kriegler et al. 1995). Surveys of
microsatellite DNA, however, suggest less
allelic diversity in the southeast portion of
the species’ range (Richards et al. 2008).
In order to determine species status within
this complex lineage, it is necessary to
have a reference point to which these
other native populations can be com-
pared. It is the purpose of this paper to
designate a neotype for S. fontinalis from
Long Island and redescribe the species.
Materials and Methods
Fishes were collected on Long Island
with the aid of a New York Department of
Environmental Conservation crew led by
Charles Guthrie at the following localities:
Nissequogue Creek 40849.610390N,
073813.5930W; Swan Creek 40846.5510N,
072859.62380W; and Connetquot River
40847.17140N, 073810.13340W. We used
brook trout from the Pigeon-French
Broad system, located in Great Smoky
Mountains National Park (GRSM), Ten-
nessee to use as representatives of the
LowerInteriorBasinforcomparative
purposes. Specifically, we collected fishes
from Indian Camp Creek 35844.2650N,
083816.6740W; Cosby Creek 35844.8810N,
083812.0200W; and Greenbrier Creek
35845.92470N, 083815.22720W. All fish
were collected by backpack electro-shock-
ing. All sites in the Lower Interior Basin
were above 680 m altitude. All fish were
anesthetized with clove oil, euthanized in
1%formalin, pinned in trays so that the
bodies were flat and the fins erect, pre-
served in 10%formalin, and placed in
permanent storage in 70%ethanol. Pig-
mentation patterns and color were record-
ed in the field via direct observation.
Counts and measurements follow Stauffer
(1991). All counts and measurements were
taken from non-spawning specimens from
the left side of the body with the exception
of gill-raker counts, which were taken on
the right side. Fin-clips were preserved in
99%ethanol.
Morphometric data were analyzed us-
ing a sheared principal component anal-
ysis, which factors the covariance matrix
and restricts size variation to the first
principal component (Humphries et al.
1981, Bookstein et al. 1985). Meristic
data were analyzed using a principal
component analysis in which the correla-
tion matrix was factored. Differences
among populations were illustrated by
plotting the sheared second principal
components (SPC2) of the morphometric
data against the first principal compo-
nents (PC1) of the meristic data (Stauffer
& Hert 1992).
We brought fin tissue samples to the
United States Geological Survey (USGS)
558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Leetown Science Center, Kearneysville,
West Virginia, for molecular analyses.
Genomic DNA was extracted from tissue
using the Puregene Kit (Gentra Systems,
Minneapolis, Minnesota). All samples were
screened for 13 microsatellite loci designed
specifically for brook trout (SfoB52,
SfoC24, SfoC28, SfoC38, SfoC79, SfoC86,
SfoC88, SfoC113, SfoC115, SfoC129,
SfoD75, SfoD91, SfoD100; King et al.
2012). Details of the master mix composi-
tion, thermal cycling parameters, and mul-
tiplexing are provided in King et al. (2012).
PCR amplifications were performed on
either PTC-200 or PTC-225 thermal cyclers
(Bio-Rad Laboratories, Hercules, Califor-
nia), and microsatellite allele sizes were
determined on an Applied Biosystems
(Foster City, California) ABI 3130. Genetic
Analyzer GeneScan 3.7 and GeneMapper
Fragment Analysis software (Applied Bio-
systems) were used to score, bin, and output
allelic data.
No records indicate fish being stocked
into the Long Island, New York streams.
We used GeneClass (Cornuet et al. 1999)
to determine the probability of each
individual collected having genotypes
found among seven potential hatchery
source populations (Table 1) used for
supplementation in the northern Atlantic
Slope region. Because detailed records of
supplementation were not available for the
Long Island, New York strains, we as-
sumed any hatchery source could have
been stocked in any drainage and, there-
fore, tested for the presence of the most
commonly stocked hatchery strains. Pop-
ulation allele frequencies were estimated in
GeneClass using the Bayesian option
(Rannala & Mountain 1997). The proba-
bility that an individual belonged to one of
the hatchery populations was calculated by
simulating 10,000 genotypes and calculat-
ing the probability of the individuals
genotyped being observed in that simulat-
ed hatchery population. While no fish was
determined to be of stocked origin based
on assignment testing, a principal coordi-
nates analysis, PAST (Hammer et al.
2001), was utilized to compare the propor-
tion of shared alleles distance among all
individuals.
The evolutionary relationships among
brook trout collections from throughout
its range (Table 2) were visualized through
the construction of a Neighbor-Joining
tree (Saitou & Nei 1987). Genetic distances
between each pair of collections were
summarized with genetic distance matrices
calculated using the Cavalli-Sforza &
Edwards (1967) chord distance in MEGA5
(Tamura et al. 2011). The strength of
support for each node in the phylogenetic
tree was tested by bootstrapping over loci
using njbpop (J.-M. Cornuet, INRA,
Montpellier, France).
Results
Populations inhabiting the North Atlan-
tic Slope (represented by fish from Long
Island, New York) were distinguished from
those populations that 1) sea run, 2) inhabit
the North Atlantic slope, 3) inhabit the St.
Lawrence River and the Great Lakes
drainages, 4) inhabit the Upper Interior
Basin (Ohio River), 5) inhabit the southern
Atlantic Slope, and 6) inhabit the lower
interior basin (Ohio River) (Fig. 1). Ordi-
nation of the inter-individual genetic dis-
tance suggested that some degree of
relatedness (overlap) existed between some
putative wild individuals from Nissequogue
Creek and fish from the Bellefonte (Penn-
Table 1.—Hatchery brook trout stocks utilized for
comparisons to putative wild collections.
Hatchery strain State
Year
sampled
Sample
size
Phillips Hatchery Maine 2005 60
Sandwich Hatchery Massachusetts 2003 37
Hyde Pond Strain New York 2005 35
Mountain Pond Strain New York 2005 31
Big Hill Pond Strain New York 2005 55
Rome Hatchery New York 2005 50
Bellefonte Hatchery Pennsylvania 2004 31
VOLUME 127, NUMBER 4 559

Table 2.—List of collections included in a range-wide genetic analysis at 13 microsatellite DNA loci in brook trout (Salvelinus fontinalis). The survey consists of
fish sampled from 18 locations representing at least six of the major phylogeographic groupings within the species’ range. Results of the comparison are presented
in a neighbor-joining tree (Fig. 1).
Major drainage Primary drainage General location Year sampled #fin clips sampled
Atlantic Ocean Freshwater R. Freshwater River 2000 50
Atlantic Ocean Watern Cove Watern Cove 2000 36
Hudson Bay Lake Mistassini Pepeshquasati River 2000 50
Hudson Bay Lake Mistassini Cheno River 2000 43
Gulf of Maine Bass Harbor/Eastern Passage Marshall Brook 2002 56
Gulf of Maine Bracy Cove/Eastern Way Jordan Stream 2002 50
Atlantic Ocean Great South Bay Nissequogue Creek 2010 30
Atlantic Ocean Great South Bay Swan Creek 2010 30
Atlantic Ocean Great South Bay Connetquot River 2010 30
St. Lawrence R. Lake Superior Grace Creek 1994 29
St. Lawrence R. Lake Superior Tobin Harbor 1996 56
Pee Dee R. Yadkin R./Pee Dee R. Harris Creek 2006 20
Pee Dee R. Yadkin R./Pee Dee R. Mitchell River 2004 20
Mississippi R. Youghiogheny R./Monongahela R./Ohio R. Puzzley Run 1998 49
Mississippi R. Youghiogheny R./Monongahela R./Ohio R. Little Bear Creek 1999 49
Mississippi R. Pigeon R./French Broad R./Holston R./Tennessee R./Ohio R. Cosby Creek 2004 48
Mississippi R. Pigeon R./French Broad R./Holston R./Tennessee R./Ohio R. Greenbrier Creek 2004 27
Mississippi R. Pigeon R./French Broad R./Holston R./Tennessee R./Ohio R. Indian Camp Creek 2000 33
560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sylvania) and Rome (New York) hatchery
strains. No overlap of populations from
Swan Creek and Connetquot River with
hatchery stocks existed (Fig. 2). Thus, we
chose the neotype from Connetquot River,
Long Island, New York.
Salvelinus fontinalis (Mitchill)
Neotype.—PSU 11387, 178.8 mm SL,
collected by Charles Guthrie and Rachel
Yoder, Connetquot River, Connetquot
River State Park, 40847.17140N,
073810.13340W, Long Island, New York,
13 Jul 2010.
Material examined.—Nine specimens,
PSU 11388, 103.6–166.1 mm SL; collection
data as for neotype. Ten specimens, PSU
11389, 130.1–206.3 mm SL, collected by
Charles Guthrie and Rachel Yoder, Nisse-
quogue Creek at Blydenburgh County
Park Office, Long Island, New York, 14
Jul 2010. Ten specimens, PSU 11390,
123.9–170.1 mm SL, collected by Charles
Guthrie and Rachel Yoder, Swan Creek at
end of Roberts Street, Long Island, New
York, 13 Jul 2010.
Description of neotype.—The brook
trout is a member of the subgenus Baione;
as such it possesses minute teeth on the
maxillaries and intermaxillaries, a patch of
minute teeth on the vomer, and a series of
teeth on the outer edges of the tongue (De
Kay 1842). Populations from Long Island
have isognathous jaws, which form a
terminal mouth that differentiates it from
Fig. 1. Evolutionary relationships of brook trout Salvelinus fontinalis genotyped at 13 microsatellite loci
inferred using the neighbor-joining algorithm (Saitou & Nei 1987) applied to the Cavalli-Sforza & Edwards
(1967) chord distance for 18 collections representing six phylogeographically distinct assemblages (sea run,
northern Atlantic slope, St. Lawrence-Great Lakes, upper interior basin/Ohio River, southern Atlantic slope,
and lower interior basin/Ohio River). The phylogenetic tree was generated using njbpop (J.-M. Cornuet,
INRA, Montpellier, France). Numbers along branches represent bootstrap support for nodes generated from
5000 randomizations. The tree is drawn to scale, with branch lengths in the same units as those of the
evolutionary distances used to infer the phylogenetic tree. Abbreviations: L. ¼Lake, MD ¼Maryland, ME ¼
Maine, NC ¼North Carolina, NP ¼National Park, NY ¼New York, TN ¼Tennessee.
VOLUME 127, NUMBER 4 561

southern populations represented by fishes
from the Lower Interior Basin, which have
retrognathous jaws that form a slightly
inferior mouth.
Jaws isognathous; teeth on upper and
lower jaws and on vomer. Lateral line
scales 109–138, neotype with 119; pored
scales posterior to lateral-line terminus at
hypural plate 5–10. Gill rakers on first
ceratobranchial 8–10, neotype 10. Princi-
pal morphometric data and meristic data
are shown in Tables 3 and 4, respectively.
Differentiation.—We present data that
Salvelinus fontinalis, genotyped at 13 mi-
crosatellite loci (Table 2), demonstrates at
least six phylogeographically distinct as-
semblages (sea run, northern Atlantic
slope, St. Lawrence-Great Lakes, upper
interior basin/Ohio River, southern Atlan-
tic slope, and lower interior basin/Ohio
River).
We further show morphological distinc-
tion of populations from Long Island from
the lower interior basin populations from
Great Smoky Mountains National Park
(Tables 5, 6). The plot of the sheared
second principal component of the mor-
phometric data versus the first principal
component of the meristic data shows that
the minimum polygon cluster formed by
fishes from Long Island, New York, does
not overlap with that formed by those
collected in the Pigeon-French Broad
Basin in Great Smoky Mountains Nation-
al Park (Fig. 3). Variables with the highest
standardized scoring coefficients for the
meristic data were pored scales posterior
to the lateral line (0.27), teeth on the lower
jaw (0.26), and lateral-line scales (0.26).
Size accounted for 95%of the observed
variance and the second principal compo-
nent for 31%of the remaining variation.
Variables with the highest loadings on the
sheared second principal component were
dorsal-fin base length (0.37), lower jaw
Fig. 2. Principal coordinates analysis depicting the relationship of the pairwise proportion of shared
alleles distances from a survey of 13 microsatellite DNA markers among selected regional brook trout
Salvelinus fontinalis hatchery strains and three wild collections (Nissequogue Creek, Swan Creek, and
Connetquot River) sampled from Long Island, New York.
562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Morphometric data of brook trout, Salvelinus fontinalis, from Long Island, New York streams.
Neotype
Connetquot River Nissequogue Creek Swan Creek
¯
XSD Range ¯
XSD Range ¯
XSD Range
Standard length, mm 170.8 152.8 19.7 103.6–170.8 160.0 24.9 130.1–206.3 149.7 15.3 123.9–170.1
Head length, mm 45.2 39.0 5.3 27.4–46.0 42.1 6.0 32.8–51.9 38.8 4.2 32.9–45.6
Percent standard length
Body depth 25.4 25.1 1.3 23.5–28.1 23.3 1.2 21.1–25.2 25.1 1.2 23.7–27.3
Snout to dorsal-fin origin 47.0 46.0 0.8 44.7–47.0 47.5 1.2 45.2–49.7 46.7 1.3 44.5–48.5
Snout to pelvic-fin origin 52.6 50.8 1.6 48.4–53.6 53.6 1.4 51.8–56.0 51.7 1.7 48.2–53.9
Dorsal-fin base length 17.6 17.3 1.4 15.7–20.1 14.6 0.8 12.7–15.5 16.6 1.2 14.7–19.0
Anterior dorsal to anterior anal 37.7 37.2 1.0 35.7–39.3 36.0 1.0 34.3–38.1 36.6 0.7 35.6–37.7
Anterior dorsal to posterior anal 46.5 44.4 1.0 43.1–46.5 42.5 1.2 41.0–44.3 43.4 0.9 41.8–44.7
Posterior dorsal to anterior anal 24.9 24.2 1.0 22.6–25.9 24.1 0.8 22.8–25.4 23.9 0.8 22.5–25.3
Posterior dorsal to posterior anal 29.9 29.4 0.8 28.1–31.0 29.4 1.0 28.3–31.0 28.6 0.8 27.3–29.6
Posterior dorsal to ventral caudal 43.1 42.6 1.1 41.1–44.0 41.8 1.1 39.3–43.3 42.4 0.9 41.1–44.0
Anterior adipose to posterior anal 15.6 14.4 1.1 13.3–16.1 13.2 0.4 12.4–13.8 13.4 0.6 12.7–14.4
Posterior anal to dorsal caudal 20.6 20.3 0.9 18.6–21.5 18.8 1.1 17.9–21.4 19.6 0.6 18.4–20.6
Anterior dorsal to pelvic-fin origin 27.1 25.6 1.5 23.4–28.4 23.7 0.9 22.5–25.0 25.2 1.1 23.8–27.4
Posterior dorsal to pelvic-fin origin 25.9 24.4 1.3 22.5–26.4 22.2 1.0 20.7–23.7 24.0 0.9 22.7–25.3
Caudal-peduncle length 16.3 16.7 0.8 15.3–18.1 15.2 1.1 13.2–17.2 16.3 0.9 14.9–17.6
Least caudal-peduncle depth 11.6 10.6 0.7 10.0–11.8 10.6 0.5 10.0–11.3 10.6 0.7 9.3–11.8
Percent head length
Snout length 23.3 22.8 1.1 21.4–25.2 23.9 1.6 21.4–26.8 22.9 1.6 20.0–25.2
Postorbital head length 54.1 51.2 1.5 49.4–54.1 49.7 2.0 45.5–52.8 51.3 2.1 47.1–53.3
Horizontal eye diameter 25.6 27.5 1.5 25.6–29.6 28.4 2.0 24.0–31.1 28.5 1.6 26.9–31.6
Vertical eye diameter 22.3 24.6 1.8 21.7–28.1 25.7 1.6 22.8–27.8 24.8 1.9 22.9–29.2
Lower-jaw length 68.3 68.9 2.9 64.8–74.2 70.3 4.0 65.1–75.4 65.1 3.4 60.0–71.1
Head depth 63.7 66.5 3.2 62.8–72.4 66.7 2.7 63.6–71.0 64.8 3.6 57.5–69.5
VOLUME 127, NUMBER 4 563

length (0.36), and vertical eye diameter
(0.28).
Discussion
In Article 75.3 of the International Code
of Zoological Nomenclature (http://www.
nhm.ac.uk/hosted-sites/iczn/code/index.
jsp?article¼75&nfv¼true) it states that a
neotype is validly designated when the
express purpose is to 1) clarify the taxo-
nomic status or the type locality of a
nominal taxon (75.3.1); 2) a statement of
the characters differentiating the neotype
from other taxa (75.3.2); 3) present data
that is sufficient to ensure recognition of
the neotype (75.3.3); 4) statement of
Table 5.—Morphometric data of brook trout, Salvelinus fontinalis, from the French Broad Drainage in
Great Smoky Mountains National Park.
¯
XSD Range
Standard length, mm 104.3 15.6 78.5–156.6
Head length, mm 27.7 5.2 19.7–43.8
Percent standard length
Body depth 24.3 2.2 19.9–30.8
Snout to dorsal-fin origin 47.8 1.2 44.9–49.8
Snout to pelvic-fin origin 51.3 1.0 49.5–54.1
Dorsal-fin base length 15.4 1.2 13.5–17.0
Anterior dorsal to anterior anal 35.8 1.4 32.8–39.7
Anterior dorsal to posterior anal 42.7 1.1 40.7–45.0
Posterior dorsal to anterior anal 22.4 1.1 20.1–24.5
Posterior dorsal to posterior anal 27.8 107 21.8–30.2
Posterior dorsal to ventral caudal 40.4 1.7 37.1–43.8
Anterior adipose to posterior anal 18.0 1.2 15.6–20.6
Posterior anal to dorsal caudal 19.1 1.1 17.0–21.9
Anterior dorsal to pelvic-fin origin 24.0 1.7 21.2–29.9
Posterior dorsal to pelvic-fin origin 21.5 1.5 19.1–24.5
Caudal-peduncle length 16.3 1.3 13.9–19.8
Least caudal-peduncle depth 10.7 5.7 10.0–11.9
Percent head length
Snout length 21.6 1.8 18.6–25.2
Postorbital head length 48.5 2.5 43.9–53.1
Horizontal eye diameter 32.0 2.3 28.0–36.5
Vertical eye diameter 28.5 2.5 24.8–33.9
Lower-jaw length 63.2 5.8 53.1–76.0
Head depth 67.5 4.1 60.2–75.3
Table 4.—Meristic data of brook trout, Salvelinus fontinalis, from Long Island, New York streams.
Counts Neotype
Connetquot River Nissequogue Creek Swan Creek
Mode %Freq. Range Mode %Freq. Range Mode %Freq. Range
Dorsal-fin rays 12 12/13 40 11–14 13 50 9–13 13 40 11–13
Anal-fin rays 10 10 100 10 10 90 10–11 11 50 10–12
Pectoral-fin rays 14 13 60 12–14 14 60 12–14 14 60 13–14
Pelvic-fin rays 10 9 70 9–10 9 70 9–10 10 70 8–10
Lateral-line scales 119 127 20 111–138 117 30 112–130 116 20 109–130
Pored scales posterior to lateral line 6 6 50 5–8 5 60 5–6 6 40 5–10
Gill rakers on first ceratobranchial 10 9 50 8–10 9 60 8–9 8 100 8
Gill rakers on first epibranchial 6 5/6 50 5–6 6 60 6–7 5 60 5–6
Teeth in outer row of left lower jaw 15 16 50 15–17 17 50 17–19 18 40 16–19
Parr marks 7 7 60 7–8 8 80 8–10 8 50 6–9
564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

reasons for believing the name-bearing
type specimen was lost or destroyed
(75.3.4); 5) provide evidence that the
neotype is consistent with the known
name-bearing type from the original de-
scription (75.3.5); and 6) provide evidence
that the neotype came from as near as
possible to the type locality.
Certainly the taxonomic status of pop-
ulations of S. fontinalis must be examined
further. To facilitate these studies, we
provide genetic and morphological data
for the neotype and associated populations
from Long Island, New York. We further
show allelic diversity throughout the range
and have differentiated the Long Island
populations morphologically from Interior
Basin populations. We supply both genetic
and morphological characters to describe
and ensure recognition of the neotype. A
type specimen of this taxon was never
preserved or catalogued into a museum.
Finally, we collected the neotype from
localities where the collections were made
upon which the original description was
based and demonstrated that there was no
discernable evidence that hatchery stocks
had altered the genetic composition of
these populations.
Acknowledgments
We want to especially thank Charles
Guthrie, New York Department of Envi-
ronmental Conservation, who organized a
crew and arranged for the collecting of fish
on Long Island, New York. We thank
Rachel Yoder for aiding in the collection
of fish. Fish were collected under the
approved IACUC 40122 research pro-
gram.
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Teeth in outer row of left lower jaw 13 8–15
Parr marks 9 6–11
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