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Group Title: Crown conch (MelongenaMelongenidae) in Florida and Alabama with the description of Melongena sprucecreekensis, n. sp. (FLMNH Bulletin v.36, no.7)
Title: The Crown conch (MelongenaMelongenidae) in Florida and Alabama with the description of Melongena sprucecreekensis, n. sp.
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00099063/00001
 Material Information
Title: The Crown conch (MelongenaMelongenidae) in Florida and Alabama with the description of Melongena sprucecreekensis, n. sp.
Physical Description: p. 181-203 : ill., map ; 23 cm.
Language: English
Creator: Tucker, John K.
Florida Museum of Natural History
Donor: unknown ( endowment )
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1994
Copyright Date: 1994
Subject: Melongenidae -- Florida   ( lcsh )
Melongenidae -- Alabama   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Includes bibliographical references (p. 202).
General Note: Cover title.
General Note: Abstracts in English and Spanish.
General Note: "Publication date: July 7th, 1994"--P. 2 of cover.
General Note: Bulletin of the Florida Museum of Natural History, volume 36, number 7, 99 181-203
Statement of Responsibility: John K. Tucker.
 Record Information
Bibliographic ID: UF00099063
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 30829704
issn - 0071-6154 ;

Table of Contents
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Full Text

of the


Melongena sprucecreekensis, n. sp.

John K. Tucker

Biological Sciences, Volume 36, Number 7, pp. 181-203 1994




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Melongena sprucecreekensis, n. sp.

John K. Tucker'


The Melongena corona species-group in the United States is reviewed. Three species, one of which is
polytypic, are recognized. These include M. corona from the Gulf Coast, M. bicolor from the Keys and
Atlantic Coast, and a new species from the Spruce Creek Estuary of the Atlantic Coast. M. c. corona ranges
from Cedar Key south to Cape Sable, Florida, and M. c. johnstonei ranges from Little Lagoon, Alabama, to
Deckle-Keaton Beach, Florida. These two subspecies differ in shoulder and anterior end spine counts. An
intergrade zone was found in the area between Deckle-Keaton Beach and Cedar Key, Florida. The second
species, M. bicolor, ranges from Matanzas Inlet to the Dry Tortugas, Florida. This species is shown to differ
in food habits and morphological details from its congeners. Morphometric data are presented to
differentiate a new species ofMelongena from the Spruce Creek Estuary of Florida.


Se revisa el grupo de species de Melongena corona de los Estados Unidos. Se reconocen tres
species, una de las cuales es politipica. Estas incluyen M. corona de la costa del Golfo de M6xico, M.
bicolor de los Cayos y la Costa Atlintica de Florida y una nueva especie del Estuario del Arroyo Spruce en
la Costa Atlaintica de Florida. En Florida, M. c. corona se distribuye desde el Cayo Cedar al norte hasta
Cabo Sable al sur. M. c. johnstonei se distribuye desde Little Lagoon, Alabama hasta la Playa de
Deckleton-Keaton, Florida. Estas dos species se diferencian en el numero de espinas anteriores y del
hombro. Se encontr6 una zona de intergradaci6n en el area entire la Playa de Deckle-Keaton y el Cayo
Cedar, Florida. La segunda especie, M. bicolor, se distribuye desde el Islote Matanzas hasta Dry Tortugas,
Florida. Esta especie difiere de otras species congendricas en habitos alimenticios y detalles morfol6gicos.
Se presentan datos morfomntricos con el objeto de diferenciar una nueva especie de Melongena del Estuario
del Arroyo Spruce en Florida.

I The author is an Assistant Rescarch Scicntist at Illinois Natural History Survey, 1005 Edwandsville Road, Wood River IL 62095, USA

Tucker, J. K. 1994. The Crown Conch (Melongena:Melongenidae) in Florida and Alabama with the
Description of Melongena sprucecreekensis, n. sp. Bull. Florida Mus. Nat. Hist., Biol Sci. 36(7):



The Melongena corona species group contains predatory gastropods that
inhabit shallow subtidal and intertidal marine and brackish water habitats along
the coasts of Florida and Alabama. They were first reviewed by Clench and Turner
(1956) who recognized three species including M. bispinosa (Philippi, 1844), M
bicolor (Say, 1827), and the polytypic M. corona (Gmelin, 1791) (Table 1). Of
these, M bicolor and M. corona occur in the United States and are the subject of
this investigation. Subsequently, Abbott (1974), considered these taxa subspecies
of a single polytypic species.
The purpose of the present paper is to characterize variation in shell
morphology from collections made within the range of the species group in the
United States. The data gathered support a different set of taxa as compared to
those recognized by Clench and Turner (1956) and Abbott (1974) who based their
taxonomic decisions on visual cues alone. In particular, I present evidence that
three species, rather than one or two, make up the species group in the United

Table 1. Comparison of the classification of the Melongena corona species group of Clench and Turner
(1956) and the one adopted in the present paper.



Clench & Turner

M. bispinosa (Philippii, 1844)
M. c. corona (Gmelin, 1791)
M. c. johnstonei Clench & Turner, 1956
M. c. altispira Pilsbry & Vanatta, 1934
M. bicolor (Say, 1827)

Keaton Beach to Cape Sable, Florida
Gulf Shores, Alabama to Panacea, Florida
Cape Sable to Matanzas Inlet, Florida
Biscayne Bay to Key West and the Dry

Present paper

M. bispinosa (Philippi, 1844)
M. c. corona (Gmelin, 1791)
M. c. johnstonei Clench & Turner, 1956
M. bicolor (Say, 1827)

M. sprucecreekensis, n. sp.

Cape Sable to Cedar Key, Florida
Panacea, Florida to Gulf Shores, Alabama
Dry Tortugas to Matanzas Inlet, Florida

Spruce Creek Estuary, Volusia County, Florida



Doug and Louise Compton, and Sue Hutchins provided information on a number of localities visited
during the current study. Neil Avery, Neil Miller, Neil Rushton, Myra J. Tucker, and Moynell M. Tucker
helped with the collection and preparation of specimens for the study. Jeannie L. Tucker assisted with the
photography. Russell Jensen and Fred Thompson read a very early version of this manuscript, and their
comments were most helpful and appreciated. Lauren E. Brown and Angelo Capperella read later versions.


Statistical methods.- Except for maximum shell width to maximum shell
length relationships, I used the SAS ANOVA and MANOVA statistical programs
for the analysis of variance with the Waller-Duncan K-ratio t-test (Waller
grouping) and Ryan-Einot-Gabriel-Welsch multiple range test (REGWQ grouping)
selected for separation of means (SAS Institute 1988). I restrict presentation of
means separations to the REGWQ results because they seem to be the most
conservative. The general linear model was used for each, because sample sizes
were unbalanced. For comparisons of maximum shell width to maximum shell
length relationships among samples, I used analysis of covariance (ANCOVA)
with the width as the dependent variable and maximum shell length as the
covariate. Models were considered significant where p < 0.05. I also subjected
means from Gulf Coast samples to cluster analysis.

Specimens examined.- I used specimens collected specifically for the study
in order to standardize collecting methods. Specimens from museum collections
were not used, because they were collected by methods unknown to me. Since
these gastropods are usually numerous where they occur, the possibility that some
criteria, such as size, appearance of the shell, coloration, or extent of spine
development, used to select the specimens collected by others cannot be eliminated.
Any criteria could bias samples. Consequently, all specimens used were collected
by the author or by the author with the assistance of other collectors using one of
two methods. In all cases, the potential collecting site was first surveyed to
estimate the abundance of specimens. Transects were used at the Courtney
Campbell and Port St. Joe localities, where specimens were exceedingly numerous.
At each collecting point along a particular transect all specimens were collected.
Collecting points were spaced at five meter intervals along the transects all of
which were at least 100 meters long. The number and length of transects and
collecting points varied. Transects were used only to avoid a priori selection of the
collecting points and cannot be considered to produce random collections as the
transect locations were not randomly selected. At the other localities every
specimen encountered was collected.


Figure 1. Coastal Florida and eastern Alabama showing approximate location of samples listed in
Appendix 1.


Besides the type specimens of the new species from Spruce Creek, voucher
specimens from the other localities are deposited in the Florida Museum of Natural

Samples.- The term sample refers to the groupings used in statistical
treatment and has no biological reality. A total of 24 samples were constructed
(Fig. 1). Each was assigned a two letter code and a number (Appendix 1). The
numbers were assigned to allow graphical presentation of the data from each
sample. Gulf Coast samples were numbered 1-16. East Coast samples were
numbered 17, 19, 21, 23, 25, 27, 29, and 32. The numbers were assigned so that
each graphical representation (Figs. 2A-D) devotes about equal space to Gulf and
East coast samples. The numbers do not represent the distance between adjacent
Biological populations are represented by 19 samples (Li, Pj, Ep, Ca, Pa, Hb,
Ck, Cc, Sp, Sa, Pi, Gd, KI, Fl, Sw, Mi, Ho, Ns, and Sc). Five samples were
composites of from two to six populations. Populations of four of these (Pk, Pn, Pc,
and Dk) were from the same immediate geographic area but, from my examination
of the sites, likely contain individuals from more than one population. Spine
counts and ratios and maximum shell width to maximum shell length relationships
for these subsamples were first compared by MANOVA and ANCOVA,
respectively. Since no significant (p > 0.05) differences were found, data from
subsamples were combined. A fifth sample (Ky) was constructed with data from
six more widely spaced populations due to the small sample sizes from each
individual population. No statistically significant differences were found with
MANOVA or ANCOVA among these subsamples.

Characters.- The following data were recorded where possible for each
specimen. The abbreviation in parentheses is used hereafter. Maximum shell
width (W) and maximum shell length (L) was measured to the nearest 0.1 mm
with vernier calipers. Spine counts included both broken and entire spines. These
counts included number of spines on the shoulder (S) and anterior end (AE) when
complete. Occasional specimens have spine rows on the whorl tops and shoulder
slope. Because they were only present in a few specimens, they were excluded
from further consideration. One ratio, shoulder spine count divided by anterior
end spine count (S/A), was derived for each specimen with a completed AE.
Unlike S, which appears in early shell growth stages, AE begins at a late or
intermediate growth stage. Because L at completion and start of AE was thought
to vary geographically, the pattern of development of AE was quantified by
defining three criteria. First, L of the smallest specimen in the sample with AE
completed was determined (= Min spine). Second, L of the largest specimen in the
sample without any trace of AE was determined (= Max 0). Third, the mean L of
specimens in the sample with incomplete AE (= Mean B) was determined.


Sex was determined by presence or absence of a penis. Individuals without a
visible penis were considered females.


Sexual dimorphism.- Collections from four samples (Sc, Mi, Cc, Sp), all
collected within one week of each other, were analyzed for possible sexual
dimorphism in W to L relationships, S, and AE. Models for all MANOVAs and
ANCOVAs were not significant at the 0.05 level.

Size-dependent variation.-- Six samples (Pj, Hb, Ck, Cc, Sc, and Mi)
contained sufficient specimens to look for evidence of size dependent variation in
W to L relationships and S. Specimens from each sample were divided into five
size classes. The size class boundaries were arbitrarily set at 30 mm increments of
L. The localities chosen were selected using two criteria. First, specimens must
have been collected in the same immediate area so that the data represent variation
in biological populations. Second, at least fifteen individuals must be present in
each of two or more size classes. Results from ANOVAs for S and ANCOVA for
W to L relationships were not significant at the 0.05 level in any of the samples.

Variation in food habits.-- During the course of this study, I made
observations of prey selection. Along the Gulf Coast and at Spruce Creek, I found
prey items included pelecypods, primarily oysters. Gunter and Menzel (1957)
made similar observations in Apalachicola Bay. On the other hand, in East Coast
specimens that I observed, all prey items were gastropods, in concurrence with the
observations of Clench and Turner (1956).
I also observed variation in prey preference within the East Coast group. I
found those from the coastal Floridian samples (observations made for Mi, Ho, and
Ns) selecting the sessile vermetid gastropod, Petaloconchus varians (Orbigny
1841), a species not previously reported in the diet of Melongena. Specimens from
the Keys and Biscayne Bay (samples Ky and Kl) were observed feeding on small
species of Cerithium, confirming the observations of Butler reported by Clench and
Turner (1956). I did not observe any predation on pelecypods, though they were

Geographic variation.-- For W to L relationships, results of ANCOVA were
highly significant. Table 2 contains a listing of samples and those samples that do
not differ significantly from them. Results of MANOVA for S, AE, and S/A,
which included all samples, were also significant (p < 0.05). Details from these
analyses are presented separately for Gulf Coast and East Coast samples.


Table 2. Results of ANCOVA for dependent variable width with covariate length by the effect of locality.
Sample abbreviations are explained in Appendix 1.

Results of ANCOVA
Species/subspecies locality w/I n not different from:

Melongena corona johnslonei Li 0.57 57 [none]
Pk 0.56 29 Pn, Pc, Ep
Pn 0.56 32 Pk, Ep
Pc 0.56 113 Pk
Pj 0.59 317 Ca, Pa, Dk, Hb
Ep 0.57 52 Pk, Pn
Ca 0.60 137 Pj, Pa, Dk, Hb
Pa 0.58 32 Pj, Ca, Hb
Dk 0.59 137 Pj, Ca, Pa, Hb
Hb 0.60 152 Pj, Ca, Pa, Dk, Sp
Ck 0.61 159 Cc, Sp, Sa, Pi
Cc 0.61 518 Ck, Sa, Pi
Sp 0.60 48 Hb, Ck, Sa
Sa 0.59 33 Ck, Cc, Sp, Pi
Pi 0.62 19 Ck, Cc, Sa
M. c. corona Gd 0.65 36 [none]
M. bicolor Ky 0.55 35 KI
KI 0.53 8 Ky
FI 0.57 15 Sw, Mi, Ho, Ns
Sw 0.59 72 Fl, Mi, Ho, Ns
Mi 0.60 183 Fl, Sw, Ho, Ns
Ho 0.59 84 Fl, Sw, Mi, Ns
Ns 0.61 26 Fl, Sw, Mi, Ho
M. sprucecreekensis Sc 0.54 144 [none]

Gulf Coast Samples (Figs. 4A-J).-- Of the three subspecies of M. corona
(Table 1) that Clench and Turner (1956) recognized, two of them, namely the
nominate race and M. c. johnstonei were said to occur along the Gulf Coast from
Little Lagoon, Alabama, to Cape Sable, Florida. Clench and Turner (1956)
suggested that specimens from between Keaton Beach and Panacea, Florida, were
intergrades between M. c. corona and M. c. johnstonei. Clench and Turner based
their concepts on the widely used nonnumeric method of simply looking at
specimens from many locations and reporting the differences they observed without
statistical analysis. My conclusions about the Gulf Coast specimens, which are
supported by statistical treatment of numeric data in general, mirrors their
Statistical analysis supported recognition of two Gulf Coast subspecies of M.
corona. Visually, specimens from Li and Gd look very different. The latter are
broad bodied with a few large shoulder spines, whereas the former are narrower

24 -
- 23-
3 22
0 21
0 20-
'0. 19


- a

ohnone Ep spiucecreekensis
johnstonei Ep Sc
Pc Pa


50 Hb sw bicolor
Dk Pi
3 Ck 70

5 Cc 17 Ns

S92 4 1016

2040 1 2 3 4 5 6 735 3

0 1 2 3 4

B Sample number




K bicolor

Sw Mi
Pi Ho Ns


15 72 183 84 1



Sample number



Cc Ns
intergrades Ck Pi
johnstonei 0Hb Sp Mi

Pk Pa 36 Ho
Ep I



35 144

012345678 4 7 9

r. Sample number



johnstone corona bicolor

Pc a Gd MI
Hb Cc Sa
Ep Ck Pi Sw
Ca 3

4 Sc

5 204 4 10
14 50 0 3 92 35 17 16
6 70

0 1 2 3 4 5 6 7 8 9 1 1 1
0 3 4

Sample number

Figure 2. (A) comparison of S (y-axis); (B) comparison of AE (y-axis); (C) comparison of W/L (y-axis); and (D) comparison of S/A (y-axis) of the Melongena
corona species group. Gulf Coast samples are arranged so that the western most sample is on the left side of each figure and the southeastern most sample is in the
right middle of each figure. East Coast samples are arranged so that the southern most sample is in the right middle of each figure and the northern most sample is
at the right side of each figure. Separation of adjacent samples does not correspond in any way to geographic distances between samples. The horizontal line
represents the mean while the vertical line represents the mean plus one standard deviation. Letter codes above the vertical lines are sample abbreviations
(Appendix I contains localities included in each sample). Numbers at the bottom of the vertical lines are sample sizes.


bodied and have many relatively small shoulder spines. However, these two
extremes result from more or less clinal variation in S (Fig. 2A), AE (Fig. 2B), and
W/L (Fig. 2C).
However, the lines producing the end members isolated as subspecies by
Clench and Turner and myself are not simple and considerable interpopulational
variation exists complicating them. If the lines were gradual, then geographically
adjacent samples should not differ significantly in the ANCOVA for W to L
relationships and the REGWQ groupings should align samples from west to
southeast for the spine counts. Deviations from this pattern are discussed
separately for each character.
The cline in S (Fig. 2A) at first rises from LI to Pj and then falls from Pa to
Cc. Means for S from LI, Pk, and Pn do not differ (Table 3A) from each other but
are lower than those from Pj, Ep, Ca, Pa, and Dk (Table 3A). Because mean S
declines from Pa to Cc, they pass through the level of the means of S for the
western four samples. As a consequence, means of S for LI, Pk, Pn, and Pc do not
differ from those of Hb (Table 3A). The area included in the decline in mean S
from Pa to Ck closely corresponds to the intergrade zone reported by Clench and
Turner (1956). Samples from Cc, Sp, Sa, Pi, and Gd have S means that are lower
than any determined for the other Gulf Coast samples (Table 3A). These samples
are all contained in the area Clench and Turner (1956) suggested was occupied by
M. c. corona.
The cline in AE more or less parallels the cline in S. However, the far
western samples contained no (for Pk and Pj) or only one (for LI and Pn) specimen
that have completed AE which makes interpretation uncertain. However, the
pattern for AE (Fig. 2B) resembles that for S (Fig. 2A). Mean AE rises to a high
at Ca (Table 3C) then falls to a low at Cc. If the REGWQ groupings for S and AE
are compared, the rankings for each sample are roughly similar. However, sample
sizes for AE are small for some samples as compared to S.
If the changes in S and AE number parallel each other, then the ratio between
S and AE (S/A) should not show evidence of much variation that can be related to
geography. With one notable exception, this is true (Table 3B). Dk has a relatively
high S/A and is significantly different from all other samples (Fig. 2D). This may
indicate that the lines for S and AE are not closely aligned. However, the sample
size (n = 4) is small for the Dk sample.
The cline in W/L (Fig. 2C) appears more regular with W/L increasing from
LI towards Gd and does not seem to follow the pattern of rise and then fall found
for S and AE. However, the ANCOVA results mirror the pattern in S and AE. Pk,
Pn, and Pc are more similar (ANCOVA not statistically different) to each other
than they are to those of Pj, Ca, and Pa (Table 2). Samples Ck, Cc, Sp, Sa, and Pi
are more similar (ANCOVA not statistically different) to each other than they are
to Hb, Dk, Pa, Ca, Ep, and Pj which in turn are more similar (ANCOVA not
statistically different) to each other than they are to Pk, Pn, and Pc. These three
sample groupings (Li-Pk-Pn-Pc, Pj-Ep-Ca-Pa-Dk-Hb, and Ck-Cc-Sp-Sa-Pi-Gd) are

Table 3. REGWQ Groupings for samples from the Gulf Coast sites possessing the character Manova included characters S, S/A, and AE. Samples with
the same letter have means that are not significantly different.

for S for S/A for AE for mean B for mean B
sample sample sample sample sample (n>20)

Site Mean Group Site Mean Group Site Mean Group Site Mean n Group Site Group

Pj 23.5 A Dk 2.59 A LI 17.0 A Ca 108.4 4 A Pc A B
Ca 22.0 B Pn 2.13 A B Ca 14.6 A B Sc 97.1 69 A B Ck C B
Pa 21.4 B Hb 2.03 C B Ep 13.5 B Pc 95.2 35 A B Dk C
Ep 21.2 B Cc 1.71 D C B Pa 12.3 C B Ck 84.8 52 C B Sp C
Dk 19.8 C Gd 1.70 D C B Pc 12.1 C Pi 82.0 8 C B Hb C
Pc 19.4 C Pc 1.70 D C B Sp 10.2 C D Pn 79.1 11 C B Cc D
LI 18.0 D Sa 1.69 D C B Hb 9.8 E C D Ep 77.6 2 C B
Pn 17.9 D Pa 1.67 D C B Pi 8.9 E D Pk 77.4 7 C B
Pk 16.9 D Pi 1.65 D C B Dk 8.8 E D LI 75.4 3 C B
Hb 16.6 D Ck 1.61 D C B Ck 8.7 E D Pa 72.7 7 C B
Sp 14.2 E Ep 1.61 D C B Sa 8.3 E D Dk 69.4 41 C
Pi 14.0 E Ca 1.52 D C B Pn 8.0 E D Gd 69.0 1 C
Ck 13.6 E F Sp 1.40 D C Gd 7.3 E D Pj 66.2 9 C D
Sa 12.5 G F LI 1.18 D Cc 7.2 E Sp 61.1 21 C D
Gd 12.2 G Hb 61.0 61 C D
Cc 11.9 G Sa 45.4 9 E D
Cc 38.8 208 E


close to the geographic distributions of M. c. johnstonei, johnstonei-corona
intergrades, and M c. corona, respectively, of Clench and Turner (1956).
The presence or absence of AE can also be used as a taxonomic character so
long as it is related to shell length, because AE appears at some stage of growth
and is not present at all growth stages. Table 5 contains data from samples where
at least 30 specimens had begun but not completed AE. It is interpreted by me to
show that specimens from western samples, such as Pk, Dk, and Hb, begin
development of the AE spine row at a larger size (bigger Mean B) than do those
from sample Cc. ANOVA for L among specimens with AE started for Gulf Coast
samples with more than 30 specimens support differences between western and
southern samples in the size at appearance of AE in samples with more than 30
specimens (Table 3E), because samples align geographically by Mean B. If all
Gulf Coast samples are included (Table 3D), then the relationship is less obvious.
The results discussed above suggest much the same conclusions about Gulf
Coast Melongena that were reached by Clench and Turner (1954). These are that
the Gulf Coast Melongena are conspecific with each other and that two subspecies
can be recognized. The nominate race, M. c. corona, ranges from Cedar Key south
to Goodland. I also found dead specimens of M. c. corona at Everglades City and
Flamingo. The second subspecies, M. c. johnstonei, ranges from Little Lagoon,
Alabama, to Panacea, Florida. Intergrades between these two subspecies occur
from Panacea to Cedar Key.
Cluster analysis (Fig. 3) strongly supports recognition of two subspecies,
because all of the samples from areas containing shells identified as M. c. corona
by Clench and Turner (1956) separate out from those that they and I identified as
M. c. johnstonei or as intergrades. Cluster analysis is not as successful in isolating
M. c. johnstonei from intergrades. This is due to the rise and fall in S and AE
counts in the region occupied by M. c. johnstonei and the intergrades previously
This pattern of variation may be attributable to the two major rivers that enter
the Gulf of Mexico within the range of M corona. Variation in S demonstrates a
possible effect on clinal variation by the rivers. Two changes in S can be seen in
Figure 2A. The most westerly of these occurs between Pc and Pj which straddle
Apalachie Bay where the Apalachicola River enters the Gulf. At this point S is
higher than for those samples to the west (Table 3A) but begins a decline to a low
at the second more dramatic change between Dk and Ck which straddle the mouth
of the Suwannee River. The three groupings suggested to exist by the results of the
ANCOVA for W and L relationships also break at these points. Specimens from
Ck south also cluster together (Fig. 3).
These rivers may act as barriers to gene flow in the species. The postulated
effect of these rivers on Melongena is speculative, but it is known that M. c.
johnstonei becomes inactive when exposed to lowered salt concentrations (Gunter
and Menzel, 1957). This snail has no planktatrophic stage and hatches at a
crawling stage rather than as a floating veliger or larval shell (Clench and




johnstonei Li

johnstonei Pn

johnstonei Pk

r johnstonei X corona Hb

sprucecreekensis Sc

johnstonei Pc

johnstonei X corona Dk

johnstonei X corona Ep

johnstonei Pa
- _
johnstonei X corona Ca

johnstonei X corona Pj

Minimum distance 0

Figure 3. Linkages and minimum distances for cluster analysis of all Gulf Coast samples and M.
sprucecreekensis (Sc) for characters S, W/L. AE, and S/A. Sample abbreviations are explained in
Appendix one. Solid lines represent minimum distances while dashed lines represent linkages
between clusters.



Figure 4. Melongena corona. M. c. corona from Ck (Figs. A, G), shell lengths 88.5 (A), and 76.8
mm (G); M. c. corona from Cc (Figs. H-I), shell length 73.1 mm (H-I); B, C, M. c. johnstonei
topotypes from LI, shell lengths 73.2 mm (B) and 69 (C); M. c. corona X M. c. johnstonei from Dk
(Figs. D, E), shell length 89.4 mm (D, E); from Pj (Fig. F), shell length 96.2 mm; from Hb (Fig. J),
shell length 99.0 mm


Turner, 1956). These two observations suggest that it is not unreasonable to
associate areas of possible reduced gene flow as elucidated from changes in shell
characters with influxes of fresh water.

East Coast Samples (Figs. 5A-E, 6A-C).- Clench and Turner (1956)
recognized two entities, M. c. altispira and M. bicolor from the East Coast of
Florida. Clench and Turner (1956) thought the latter differed from the former "by
being smaller, much lighter in color and having more and better developed
shoulder spines." Clench and Turner (1956) did not quantify these suggested
I believe that the data presented herein suggest that these two East Coast taxa
are slightly differentiated forms of a single species. Variation in W/L and S show
more or less gradual clinal changes (Figs. 2A, C). Variation in S/A also may be
clinal (Fig. 2D), a reflection of AE counts that decrease faster than S counts do
(Figs. 2A, B). However, this purported cline is generated by data collected from
just three specimens at Ns that had completed AE.
Of the samples compared for S only the Ky sample differs significantly from
the others (Table 4A). ANCOVA for W to L relationships mirrors the MANOVA
for S. The Ky and KI samples, while not significantly different from each other
(Table 2), differ from the other samples, which in turn do not differ from each
other. Such a result would be expected if a cline existed due to the considerable
distance between the Fl and KI samples.
My data do support Clench and Turner's contention that the specimens from
the Keys have more spines than those from other East Coast areas. However, my
data also show that this difference is rather minute and the result of a cline. This
suggests that the East Coast Melongena are not sufficiently differentiated by the
shell characters I studied to recognize subspecies.
However, I made no effort to quantify coloration, which Clench and Turner
(1956) also used to differentiate the two taxa they recognized. Coloration of Ky
and Kl specimens is very pale. Few specimens were found that had dark banding
on the anterior end, which is usually present in specimens from more northern East
coastal localities. Many but not all of the Ky and KI specimens also have the
shoulder spines greatly reduced in size, and a number had no spines at all. This is
rare among specimens from other East Coast samples. However, at least some
specimens from every other sample were as pale as the Ky and KI specimens.
Consequently, the East Coast Melongena should be identified as M. bicolor.
Whether M. bicolor and M. corona are conspecific or not remains a problem.
Clench and Turner (1956) reported specimens that they identified as M. c. altispira
from the Cape Sable region and in the same area that they recorded M. c. corona.
They made no mention of an intergrade zone. During the current study none of the
Florida Bay coastal areas investigated produced living specimens. Some dead
shells were encountered, all of which were AM. c. corona similar to those from Gd.


Figure 5. Melongena bicolor. A, from KI shell length 40.0 mm; B, from Ky shell length 36.2 mm;
C, from Ns shell length 66.4 mm; D, E from Mi shell length 63.1 mm. Sample abbreviations are
explained in Appendix 1.


Comparison of the my data for the Gd sample and the geographically closest
samples of M bicolor (KI and Ky) demonstrate that significant morphological
differences exist. The relationship for W to L is significantly different, as are spine
counts (Figs. 2A, C). The Gd sample includes shells with higher W/L and lower S
than any sample from the East Coast or Keys. In fact, all of the samples I identify
as M. c. corona (Cc, Sp, Sa, Pi, and Gd) differ significantly from the Ky and KI
samples ofM. bicolor. Besides these statistical differences, clinal variation in W/L
of the two are exactly opposite. In M corona the cline shows an increase in W/L
from north to south, whereas in M. bicolor the cline is one of increasing W/L from
south to north.
Since these two differ morphologically as well as in food habits, the present
paper recognizes two closely related, allopatric or parapatric species, namely M.
bicolor and M. corona.
Finally, a population was discovered in the Spruce Creek Estuary (Spruce
Creek, Strickland Bay, and Turnbull Bay) in Volusia County, Florida, that are
markedly different morphologically from surrounding populations of M. bicolor
and are distinguishable from Gulf Coast populations of M corona as well.

Melongena sprucecreekensis, newv species
Figure 6A-C

Description.-- The shell reaches at least 185 mm in length and is solid and
strongly sculptured. Mean W/L is 0.55. The shell is white in color but that is
overlain by two, or less commonly, three brown to grey bands of varying width.
The color bands are located on the body whorl with one anterior and one posterior
to the midbody region. Occasionally the posterior band is subdivided into two
bands by an intervening light colored area. Since the spire is scalariform, the
posterior color band is also exposed on the spire whorls. There are 7-10 convex
whorls. The aperture is subovate in shape. The widest portion of the body is
located well anterior of the shoulder. The outer lip may be thin or thick and
crenulate at least in specimens that are longer than 90 mm. The columella is broad
and twisted. The sculpture consists of a row of strong, erect to recurved spines
along the whorl shoulder. A row of spines may be present near the anterior end.
Shoulder spines average 16.03 (range 12-22). Anterior end spines number
between 9 and 17 with a mean of 12.43. The S/A ratio is 1.26. At the suture there
are numerous imbrications or frills representing previous growth stages of the anal
canal. In specimens more than 120 mm long, these imbrications may be raised
into spines. The operculum is horny, approximates the shape of the aperture, and
has a terminal nucleus. The periostracum is smooth, dull, and greenish in color.
The radula is identical to that of M. c. johnstonei (see Clench and Turner 1956,
plate 96). Egg capsules are also similar to those of M c. johnstonei.


Types.-- The holotype (UF 40336) and ten paratypes (UF 40337) are
deposited in the Florida Museum of Natural History, University of Florida,
Gainesville, Florida. Fifty other paratypes were also designated with ten each
being deposited in the National Museum of Natural History (USNM 792416),
American Museum of Natural History (AMNH 206086), Delaware Museum of
Natural History (DMNH 155535), Los Angeles County Museum of Natural History
(LACM 2033) and San Diego Natural History Museum (TS 81658).

Type locality.-- The type locality is designated as 7.26 km (4.5 miles) north
of New Smyrna Beach, Volusia County, Florida, where U. S. Route I crosses
Strickland Bay of the Spruce Creek Estuary.

Remarks.- This species is known only from the Spruce Creek Estuary along
the northeastern coast of Florida. An intensive search for further populations of
the species was made in the Halifax River to the north of Spruce Creek and in the
Indian River Lagoon to the south. Although M. bicolor was found both to the
north (Daytona Beach-dead specimens only) and to the south (New Smyrna Beach)
of the Spruce Creek system none was found in the system itself.

Comparison.-- The new species differs from surrounding populations of M
bicolor in shell morphology and food habits.
Among the data collected during the current study, S/A and AE for the Sc
sample are statistically different for those determined for specimens of M. bicolor
(Table 4B, C). Even though W/L ratios of M. sprucecreekensis are numerically
similar to those of the Ky and KI samples, ANCOVA separates the Sc sample from
all M. bicolor samples as well as from all M. corona samples.
Although it has not been tested experimentally, Figures 2B-D suggest that
little or no gene flow exists between the Sc sample and neighboring samples of M.
bicolor. Besides differences in means for spine counts, the developmental pattern
of anterior end spine row also differ. M. sprucecreekensis has a distinctly higher
value of Mean B when compared to M. bicolor. Min Spine is also higher in M.
sprucecreekensis than it is in any population of M. bicolor. Thus specimens of M
sprucecreekensis smaller than 58 mm are unlikely to have any sign of the anterior
end spine row, while most specimens of M. bicolor of this size (and smaller) will
have the spine row started.
These two species also differ in food habits. M. sprucecreekensis feeds
primarily on oysters (Fig. 6C), with some other bivalves also being taken, whereas
M. bicolor feeds primarily on gastropods.
Finally, M. sprucecreekensis reaches a size unknown in M. bicolor. Clench
and Turner (1956) reported a maximum shell length of 72 mm for M.


Table 4. REGWQ Groupings for samples from the East Coast sites possessing the character. Manova
included characters S, W/L, S/A, and AE. Samples with the same letter have means that are not significantly
different. Sample abbreviations are explained in Appendix 1.

For S For S/A For AE
Sample Sample Sample

Sites Mean Group Sites Mean Group Sites Mean Group

Ky 18.2 A Ns 2.27 A Sc 12.6 A
Sc 16.0 B Mi 1.72 B Sw 9.9 B
KI 15.9 B Sw 1.62 B Mi 9.1 B C
Fl 15.6 B Sc 1.28 C Ns 7.0 C
Mi 15.3 B
Sw 15.2 B
Ho 15.1 B
Ns 14.7 B

bicolor. The largest specimen observed in my study was 89 mm in maximum shell
length. M. sprucecreekensis reaches at least 185 mm in maximum shell length.
Absolute size is of course a difficult character to use in determining conspecificity.
It is always possible that utilization of oysters as prey items in some way enhances
growth, and the observed differences are ecological and not genetic. This seems
unlikely to me, because smaller specimens from Sc do not differ from larger ones
in morphological parameters.
M. sprucecreekensis more closely resembles certain populations of M. corona
from the Gulf Coast, even though it is located geographically closest to populations
ofMM bicolor. While it is statistically different in W to L relationships, in AE it is
statistically identical to Pa, Pc, Sp, and Hb samples from the Gulf Coast.
M. sprucecreekensis also resembles the subspecies of M. corona in food
habits. Both primarily predate on pelecypods.
It may seem anomalous that the morphological parameters and food habits of
M. sprucecreekensis more closely resemble those of cross-state M. corona than it
does the surrounding populations of M. bicolor. There are at least three possible
First, the Spruce Creek population could represent a relatively recent
introduction of Gulf Coast specimens to the Spruce Creek area by human or natural
means. This possibility is considered remote as morphometric parameters of M
sprucecreekensis should approximate those of one of the Gulf Coast samples if this
were an introduction. In S, Sc is statistically identical to Hb and Pk of the Gulf
Coast. In AE, Sc is statistically identical to Ep, Ca, Pa, Pc, Sp, and Dk. In Mean



Figure 6. A-C, Melongena sprucecreekensis, n. sp. A, paratype, shell length 151 mm; B, holotype,
shell length 185 mm; C, a specimen in situ on an oyster bar at the type locality. Sample abbreviations
are explained in Appendix 1.


Table 5. Developmental pattern of anterior end spine row in several populations of the Melongena corona
species group. Sample abbreviations are explained in Appendix 1. All measurements are in mm.

sample Max. 0 Min. spine Mean B n

corona johnstonei
Pc 122.1 78.8 97.1 35
corona Xjohnstonei
Dk 103.2 69.3 69.4 41
Hb 93.6 52.7 61.0 61
Ck 104.6 74.3 84.8 52
corona corona
Cc 58.2 29.9 38.8 208
Sc 74.7 61.2 98.5 69

B, Sc is statistically identical to Ca and Pc. Sc clusters with Pk and Hb (Fig. 3).
Although it resembles Gulf Coast specimens in a general way, it does not match up
with any sample from the Gulf Coast.
Second, AlM. sprucecreekensis could represent a phenotypical response to the
particular conditions found in the Spruce Creek area by specimens that are
genotypically similar to the surrounding populations of M. bicolor. M. bicolor
does occasionally prey on bivalves and may be associated with oysters in some
areas of the Indian River (J. Johnson, pers. comm.). A large East Coast
Melongena that feeds on oysters may simply reflect faster growth rates under those
conditions. However, M. sprucecreekensis differs from equal sized M. bicolor in
all counts and ratios. M. sprucecreekensis is significantly narrower bodied at all
growth stages than is M. bicolor. Such consistency in morphological differences
strongly suggests to me that M. sprucecreekensis is not an ecotypic variant of M.
Finally, M. sprucecreekensis may be a Pleistocene relict that reached the East
Coast from the stock now restricted to the Gulf Coast during a period of elevated
sea level (Cooke, 1939). Left behind by falling sea levels, it was isolated by the
colonization of the East Coast by M. bicolor. This hypothesis cannot be tested
from the available data. However, it should prove amenable to allozyme testing, as
was done for the Seaside sparrow (Ammodramus maritimus) by Avise and Nelson
The molluscan fauna of Florida has been extensively studied, and thus it is
unexpected that such a large species has remained undiscovered for so long.
However, the validity of M. sprucecreekensis is supported by morphometric


differences and by differences in food habits as compared to the parapatric M.
bicolor. One could argue that what I identify as a full species should be considered
a subspecies of M. corona because no one trait studied by me will separate every
individual of M. sprucecreekensis from every individual of M. corona. Whatever
the taxonomic status, it is important to recognize that, within a reasonable degree
of certainty from the shell traits reported herein, an entity genetically distinct from
M. bicolor exists along the East Coast.
Unlike most molluscan species along Florida's rapidly developing and already
heavily developed East Coast, AM. sprucecreekensis inhabits a limited range. It also
appears to be dependent on oysters for food. Consequently, any development in the
Spruce Creek Estuary that adversely impacts the oyster populations will adversely
impact M sprucecreekensis. Development has not left this area untouched. Much
of the eastern part of the system is developed, and other projects are planned for
both the upstream and downstream portions of the estuary. Although quantitative
studies of relative abundance have not been completed, preliminary data indicates
that oyster bars, a focus of activity for M. sprucecreekensis, support far fewer
individuals in developed parts of the system than is the case in relatively
undeveloped Strickland Bay area.


Abbott, R. T. 1974. American Seashells second edition. Van Nostrand Reinhold Co., New York, 663 pp.
Avise, J. C., and W. S. Nelson 1989. Molecular genetic relationships of the extinct Dusky Seaside Sparrow.
Science 243:646-648.
Clench, W. J., and R. D. Turner 1956. The Family Melonginidae in the Western Atlantic. Johnsonia
Cooke, C. W. 1939. Scenery of Florida interpreted by a geologist. Florida Dept. Cons. Geol. Bull.
Gunter, G., and R. W. Menzel 1957. The crown conch, Melongena corona, as a predator upon the
Virginia oyster. Nautilus 70(3):84-87.
SAS Institute. 1988. SAS/STAT user's guide. SAS Institute, Inc., Cary, North Carolina, 1028 pp.



The abbreviations and locality numbers used in the tables and figures are listed with each followed by
the locality or localities from which specimens making up that particular sample were collected. The
number in parenthesis is the number of specimens collected at each locality.

Gulf Coast. 1-Li, Little Lagoon, 7 miles west of Gulf Shores, Alabama on Alabama route 180 (57);
2-Pk, Alabama end of Perdido Key (4), Cotton Bayou, Orange Beach, Alabama (11), Big Bayou, Orange
Beach, Alabama (14); 3-Pn, U. S. route 98, north of Gulf Breese, Pensacola, Florida (13), U. S. route 98,
north of Pensacola Beach, Santa Rosa Sound (15), Florida route 292, 3 miles east of Pensacola, Santa Rosa
Sound (4); 4-Pc, Florida route 77 bridge across bayou George (49), U. S. route 98 bridge to Tyndall Air
Force Base, along bridge abutment on the base (64); 5-Pj, Port St. Joe, along U. S. route 98, St. Joseph Bay
(315); 6-Ep, U. S. route 98, 1 mile east of East Point, Florida (52); 7-Ca, halfway between Carabelle and
Lanark Village on U. S. route 98 (26); 8-Pa, end of Florida route 372A near Panacea (29); 9-Dk, outside
Deckle Beach on Florida route 361 (86), outside Keaton Beach on Florida route 361 (32), Jug Island off
Florida route 361 (19); 10-Hb, outside Horseshoe Beach on Florida route 351 (149); 1 l-Ck, both sides of
Cedar Key along Florida route 24 (164); 12-Cc, along both sides of Courtney Campbell Causeway (Florida
route 60) in Tampa Bay (518); 13-Sp, rest area on south end of the Sunshine Skyway bridge, St. Petersburg
(48); 14-Sa, causeway between Sanibel and Captiva Island, on the inland side (33); 15-Pi, Pine Island Sound
off Pineland (19); 16-Gd, east side of Marco Island, Goodland (36).

East Coast and the Keys. 32-Sc, junction of Spruce Creek and U. S. route 1 (144); 29-Ns, along the
Indian River Lagoon where U. S. route AlA crosses, west side of bridge (26); 27-Ho, Indian River, 1/4 mile
north of Haul-Over canal, Merritt Island, on the east side of river (84); 25-Mi, Florida route 520 bridge
between Merritt Island and Cocoa Beach, in the Banana River (183); 23-Sw, causeway between Port Sewall
and Sewall's Point on U. S. route A1A (72); 21-Fl, at the end of Florida route 84 (15); 19-KI, east end of
Key Largo on both Florida Bay and Atlantic sides (18); 17-Ky, along U. S. route 1 on the Gulf of Mexico
side of the road at the following keys: Ramrod Key (13), Bahia Honda Key (16), Little Duck Key (1),
Marathon Key (4) and Lower Matecumbe Key (2).

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