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2013 SOUTHEASTERN NATURALIST 12(1):161–170
Growth Rates and Age Estimations of the Fuzzy Pigtoe,
Pleurobema strodeanum: A Species Newly Listed under the
Endangered Species Act
Evelyn G. Reátegui-Zirena1,2, Paul M. Stewart1,*, and Jonathan M. Miller1
Abstract - Pleurobema strodeanum (Fuzzy Pigtoe), is listed as threatened under the
Endangered Species Act. The purpose of this study was to estimate growth rate, morphometric
ratios, and ages for this species. One hundred and sixty-one P. strodeanum
were originally tagged and measured in 2004 at Eightmile Creek, Walton County, FL.
In 2011, 28 of those 161 were recovered and re-measured. Growth-rate percentages
were determined based on length, width, height, and volume. Age was estimated using
the von Bertalanffy growth equation. Results indicate that over a period of seven
years, P. strodeanum grew 0.48 mm/year (SD = 0.14) in length. The age estimations in
this study ranged from 48.7 to 74.5 years. The growth constant (K) was 0.018 per year,
and the asymptotic length (L∞) was 71.97 mm. Natural history data such as these will
be useful in understanding the basic biology of P. strodeanum and may assist in successful
North American freshwater mussels (Unionidae) are one of the world’s
most threatened groups of organisms. Mussel diversity is concentrated in
the southeastern US, with Alabama being home to 183 species (Jeff Garner,
Alabama Department of Conservation and Natural Resources [ADCNR], Florence,
AL, 25 January 2012 pers. comm.), more than any other state (Williams
et al. 2008). Longevity, growth, and age estimates are needed to better assess
the risk of extinction of rare and endangered species, and add to an understanding
of basic biology that may lead to successful mussel management
(Williams et al. 1993). Pleurobema strodeanum, Walker (Fuzzy Pigtoe) is
found in the Escambia, Yellow, and Choctawhatchee River drainages. Populations
of this species are decreasing and it is currently listed as threatened under
the Endangered Species Act (US Fish and Wildlife Service 2012).
Management efforts are often based on accurate information regarding lifehistory
traits such as age and growth, as such data can be used in modeling
population dynamics (Haag 2009). Changes in length, width, and height of a shell
over time are measurements commonly used for determining rates of growth in
mussels (Dehnel 1956). Furthermore, the ratios between them are used to study
the change in proportion of individuals as a consequence of growth (Lauzon-
Guay et al. 2005), and these variations in shell morphology can be used as
indicators of environmental change (Widarto 2007).
1Department of Biological and Environmental Sciences, Troy University, Troy, AL
36082. 2Current address - Institute of Environmental and Human Health, Department of
Environmental Toxicology, Texas Tech Univesity, Box 41163, Lubbock, TX 79409. *Corresponding
author - email@example.com.
162 Southeastern Naturalist Vol. 12, No. 1
As a group, freshwater mussels are known to be long lived and slow growing.
The lack of age and growth information for these organisms might be
due in part to methodological issues (Haag and Rypel 2011). Growth rings
occur on the external surface of mussel shells and have been used for studying
growth and age. However, results are inconsistent among individuals and
even different regions of a single shell. Because environmental disturbances,
including handling during sampling, can cause the formation of external and
internal false rings, age is often overestimated and growth rate can be underestimated
(Haag 2009, Haukioja and Hakala 1978). Therefore, validation
of annual ring counts is highly recommended to estimate age (Haag 2009).
However, it is not the best approach for species of conservation concern that
should not be sacrificed. Following tagged mussels under natural conditions
is thought to be the most realistic method for estimating growth rate (Karatayev
et al. 2006). With tag and recapture data, length can be used in the von
Bertalanffy growth function (VBGF), which predicts length of an individual
as a function of its age (Kesler and Downing 1997). Measurements of growth
rate and estimation of K (growth constant), may lead to an understanding of
whether or not a population is under stress and why or if they are in decline.
The objectives of this study were to estimate P. strodeanum age, growth rate,
and morphometric relationships over a seven-year period.
Pleurobema strodeanum was studied in Eightmile Creek, Walton County,
FL. Mussels were collected in 2004 by handpicking using qualitative visual
and tactile searches that covered 250 m downstream from a bridge crossing.
Individuals were measured using digital calipers to the nearest 0.01 mm along
three shell axes (length, width, and height). Mussels were tagged by removing
a small portion of the periostracum on the shell and attaching a numbered
Floy® shellfish tag using cyanoacrylate glue (Sickel et al. 1997). Voucher specimens
were preserved in 70% ethanol and are curated in the Troy University
collection. Seven years later, in 2011, 28 P. strodeanum, of the 161 originally
tagged, were recaptured and measured. Growth rate percentages (G%) were
calculated as follows:
G% = 100 * (Mf - Mi) / Mi,
where Mf and Mi are the final and initial measurements (length, width, height, and
volume) in 2011 and 2004, respectively (Negishi and Kayaba 2009). Morphometric
ratios were determined as follows, width to length (W:L), height to length
(H:L), and height to width (H:W).
Volume was calculated using the formula:
V = (Length x Width x Height) / C,
where the constant (C) was determined with the method used by Martins et al.
(2011). Briefly, 18 P. strodeanum shells between 31.6 mm and 60.4 mm in length
2013 E.G. Reátegui-Zirena, P.M. Stewart, and J.M. Miller 163
were sealed with Parafilm®. Mussels were immersed in water, and the volume
was determined by displacement to the nearest 0.1 ml in a graduated cylinder.
These numbers were plotted against the volume obtained by multiplying the
length, width, and height. The regression line was calculated, and the slope (C)
was 2.51 (R2 = 0.975, P < 0.001). The von Bertalanffy growth equation was used
to determine the age of the individuals. The equation is expressed as:
Lt = L∞ (1 - e - K [t - t0)])
It is usually applied when age and body size are known; however, it can be inverted
to find age as follows:
t = ln[(L∞ - Lt) / (L∞ - L0)] / (-K),
where L∞ (asymptotic length) is the theoretical maximum length at infinite age,
Lt is the length of the organism at time t (age), L0 is the length of the organism
at time 0 since age is not known, and K is Brody’s growth constant. Using mark
and recapture growth data, L∞ and K can be estimated using a linear regression of
the Ford-Walford relationship (Ricker 1975):
L∞ = [a / (1 – β)]
K = -ln β,
where a is the y intercept and β is the slope (Anthony et al. 2001, Haag 2009). Assuming
constant growth, the Brody’s growth constant (K) was divided by seven
to be used in the inverted von Bertalanffy growth equation since the estimated K
was based on a seven-year period.
Data were plotted using Excel 2007 for Windows, and linear regressions
between growth rate (%) and all the measurements taken were performed using
SPSS® version 11.0. Data were tested for normality, and a t-test was applied to
measure differences between smaller and larger individuals’ growth-rate percentages,
and differences between morphometric ratios in 2004 and 2011. If data
were not normal, a Mann Whitney (U) or Wilcoxon (Z) test was used. A t-test
was used for width, width:length, and height:length; Mann-Whitney (U) tests
were used for length, height, and volume; and Wilcoxon (Z) test for height:width.
In addition, ratios in 2004 and 2011 were plotted against length to determine if
samples were biased by the size of the individuals.
In the present study in 2004, maturity for P. strodeanum was determined at
37.65 mm based on reproductive activity. Juveniles were not sampled, and all
(n = 161) but seven individuals were larger than 40 mm. The theoretical maximum
length (L∞) found for this species was 71.97 mm, a close approximation
of the 75 mm reported by Williams et al. (2008). During 2011 resampling, a
presumed juvenile individual (providing limited evidence for recruitment but
not considered for analysis) was collected (length = 21.58 mm, width = 14.25,
and height = 8.55).
164 Southeastern Naturalist Vol. 12, No. 1
In 2004, 161 P. strodeanum were marked, and 28 were recaptured in
2011 (for a recapture rate of 17.4%). Of the 28 P. strodeanum recaptured, 27
were used for analysis (one individual displayed negative growth and was
not used in any analysis). Of the tagged mussels, the smallest P. strodeanum
was 37.65 mm and the largest was 51.62 mm in length in 2004, while the individuals
ranged from 41.89 mm to 53.04 mm in 2011. From 2004 to 2011,
individuals grew an average of 3.37 mm in length, and showed a mean increase
in width of 14.08% in seven years. This width increase was almost double the
percentage of growth in length (7.84%) or height (6.87%). The mean volume
for this species increased 31.44%. For purposes of analysis, it was assumed
that growth was constant each year, and results were divided by seven to estimate
the annual growth rate (Table 1). Individuals grew an average of 0.48
mm in length per year. Pleurobema strodeanum growth rate percentage in
length (r2 = 0.273, P = 0.005), width (r2 = 0.609, P < 0.001), and volume (r2 =
0.490, P < 0.001) decreased with increased initial size (Fig. 1). When growth
rate percentages of the nine smaller and the nine larger individuals were tested,
there was a significant difference in length (U = 10.0, P = 0.006), width (t =
7.33, P < 0.001), and volume (U = 15.0, P = 0.024). From 2004 to 2011, the ratio
of width to length increased and the height-to-width ratio decreased, while
relative height (height:length) remained the same (Fig. 2). When morphometric
ratios were tested, there was a significant change in width-to-length ratio
(Z = -2.600, P = 0.009) and height-to-width ratio (t = 4.094, P < 0.001). Also,
when ratios were plotted against length, linear regressions were not significant,
suggesting that samples were not biased by the size of the individuals.
The Brody’s growth constant (K) was 0.125 in a seven-year period (0.018 per
year). The estimated ages ranged from 41.3 to 70.5 years in 2004, with a mean
of 51.9 (SD = 6.16), and 48.7 to 74.5 years in 2011, with a mean of 58.9 (SD =
6.48) (Fig. 3).
In comparable studies using different species, Kesler et al. (2007) compared
the growth rates of Elliptio complanata Lightfoot (Eastern Elliptio)
among three sites in Rhode Island over 14 years. Individuals at one site grew
4.3 mm, while the other two sites had growth rates that were 3 to 8 times
Table 1. Pleurobema strodeanum growth (length, width, height and volume) over seven years and
Growth (mm) Growth rate (percentage)
Measurements Over 7 years Per year Over 7 years Per year
Length 3.37 0.48 7.84 1.12
Width 3.57 0.51 14.08 2.01
Height 1.21 0.17 6.87 0.98
Volume (mm3) 2.47 0.35 31.44 4.50
2013 E.G. Reátegui-Zirena, P.M. Stewart, and J.M. Miller 165
Figure 1. Regression
f o r P l e u -
(A) init ial
and (C) initial
166 Southeastern Naturalist Vol. 12, No. 1
greater. Kesler et al. (2007) suggested that the great difference in growth rates
could be due to food limitation/availability at the site with less growth. In
P. strodeanum, there was a significant difference in growth rate percentages
Figure 2. Morphometric relationship means (± 1 SE) for Pleurobema strodeanum measured
in 2004 and 2011. Note: W = width, L = length, and H = height.
Figure 3. Mussel estimated ages from the inverted von Bertalanffy growth equation using
tag and recapture growth rates for Pleurobema strodeanum in 2004 and 2011.
2013 E.G. Reátegui-Zirena, P.M. Stewart, and J.M. Miller 167
when considering length, width, and volume between smaller and larger
individuals. The smaller P. strodeanum had a higher growth rate than did
larger individuals. This result was expected since mussels grow more rapidly
during their first few years, and the growth rate decreases over time
(San Miguel et al. 2004). For instance, annual growth rates of Pronodularia
japanensis (Lea) individuals greater than 40 mm (adults) were less than 2%, while
juveniles (10 mm) had an annual growth rate of ≈100% (Negishi and Kayaba
2009), suggesting a strong size-dependent relationship.
Variations in allometric relationships are thought to be due to site-specific
factors, reproductive seasons, sex, and are species-specific (Gimin et al.
2004). Shell length and obesity (width:length) increased with age in Pleurobema
strodeanum, while relative height (height:length) remained the same.
Similar results have been observed in Mytilus edulis L. (Blue Mussel), which
has been widely studied and used as a reference for variations in shell morphology.
In M. edulis, the shell width-to-length ratio increased with age,
though shell height-to-width ratio decreased with age and increased shell
length (Seed 1968). These observations were confirmed by Lauzon-Guay et
al. (2005), who additionally found that the height-to-length ratio decreased
as M. edulis grew. This ratio change is supported by the observation that shell
morphology is greatly influenced by mussel density, and more elongate shells
(lower height:length) are expected in the field if mussels grow in crowded conditions
(Seed 1968). Moreover, relative height might decrease with an increase
in shear stress exposure (Bailey and Green 1988). Similar results were found
for Velesunio ambiguous (Philippi) (The Flood Plain Mussel), which suggested
that the decrease in relative shell height is due to the progressively dorsal arching
of the shell (Widarto 2007).
The growth constant (K) is a major determinant for life span and maximum
shell length. The constant declines as life span and maximum shell length increases,
and it describes the shape of an individual’s growth curve (Bauer 1992).
Growth constants (K) vary widely among mussel species, and reported values
range from 0.063 mm for Pinna nobilis L. (Noble Pen Shell) (Galinou-Mitsoudi
et al. 2006) to 0.655 for Megapitaria squalida (Sowerby) (Schweers et al. 2006).
Members of the tribe Pleurobemini, to which Pleurobema strodeanum belong,
are long-lived and slow-growing, having low K values. Fusconaia cerina (Conrad),
F. ebena (Lea), and P. decisum (I. Lea) (Southern Clubshell) are reported
to reach ages of 45–51 years. The K value was determined for F. cerina (0.173),
P. coccineum (0.134), and P. decisum (0.145), and it was considered that a K =
0.05 is extremely slow growth (Haag and Rypel 2011). However, none of these
species’ growth constants were as low as the K for P. strodeanum in the present
study, which was only 0.018 per year.
The growth constant (K) is an indicator of the energy allocated for growth compared
to other functions such as reproduction (Haag and Rypel 2011). The very
low K found for P. strodeanum in this study might be a response to environmental
factors such as temperature, trophic conditions, and stress. Pilarczyk et al. (2006)
168 Southeastern Naturalist Vol. 12, No. 1
showed a decline from the 1990s to 2004 in the number of sites at which candidate
species were reported in the Choctawhatchee drainage, of which Eightmile
is a tributary. Within this watershed and its tributaries, P. strodeanum showed the
greatest decrease in the number of sites in which it was found. The extremely low
K that we found suggested that this population may be under stress. The decrease
in distribution along with a slow growth rate supports a need for conservation
efforts to identify the limiting factors for growth. Only measuring mature individuals
dictates that K and growth rate are lower, therefore, more work needs to
be done including juveniles so that a more accurate set of growth variables can
This study has provided some important insights into the ecology of this
Federally threatened mussel species, showing that tag-recapture methods can
be used to obtain estimates of important life-history and growth parameters. As
a future recommendation, additional studies are necessary to assess population
demographics such as recruitment and survival to better understand population
trends over time. Our team is currently doing such work at this site for these and
other candidate species through occupancy and detection studies.
We thank Holly Blalock-Herod and Adam Kaeser (USFWS), Megan M. White, and
Sumit Nanda for their assistance in the field, and J. Murray Hyde for his assistance with
laboratory work. In addition, we thank Dr. Wendell R. Haag (USDA Forest Service,
Center for Bottomland Hardwoods Research) who helped with the growth equation to
determine age and Jeff Garner (ADCNR) for his contributions in editing the manuscript.
Anthony, J.L., D.H. Hesler, W.L. Downing, and J.A. Downing. 2001. Length-specific
growth rates in freshwater mussels (Bivalvia: Unionidae): Extreme longevity or generalized
growth cessation. Freshwater Biology 46:1349–1539.
Bauer, G. 1992. Variation in the life span and size of the Freshwater Pearl Mussel. Journal
of Animal Ecology 61(2):425–436.
Bailey, R.C., and R.H. Green. 1988. Within-basin variation in shell morphology and
growth rate of a freshwater mussel. Canadian Journal of Zoology 66:1704–1708.
Dehnel, P.A. 1956. Growth rates in latitudinally and vertically separated populations of
Mytilus californianus. Biological Bulletin 110(1):43-53.
Galinou-Mitsoudi, S., G. Vlahavas, and O. Papoutsi. 2006. Population study of the protected
bivalve Pinna nobilis (Linnaeus, 1758) in Thermaikos Gulf (North Aegean
Sea). Journal of Biological Research 5:47–53.
Gimin, R.L., V. Mohan, V. Thinh, and A.D. Griffiths. 2004. The relationship of shell
dimensions and shell volume to live weight and soft tissue weight in the Mangrove
Clam, Polymesoda erosa (Solander, 1786) from northern Australia. Naga
Haag, W.R. 2009. Extreme longevity in freshwater mussels revisited: Sources of bias
in age estimates derived from mark-recapture experiments. Freshwater Biology
2013 E.G. Reátegui-Zirena, P.M. Stewart, and J.M. Miller 169
Haag, W.R., and A.L. Rypel. 2011. Growth and longevity in freshwater mussels: Evolutionary
and conservation implications. Biological Reviews 86:225–247.
Haukioja, E., and T. Hakala. 1978. Measuring growth from shell rings in populations
of Anodonta piscinalis (Pelecypoda, Unionidae). Annales Zoologici Fennici
Karatayev, A.Y., L.E. Burlakova, and D.K. Padilla. 2006. Growth rate and longevity
of Dreissena polymorpha (Pallas): A review and recommendations for future study.
Journal of Shellfish Research 25(1):23–32.
Kesler, D.H., and J.A. Downing. 1997. Internal shell annuli yield inaccurate growth
estimates in the freshwater mussels Elliptio complanata and Lampsilis radiata. Freshwater
Kesler, D.H., T.J. Newton, and L. Green. 2007. Long-term monitoring of growth in
the Eastern elliptio, Elliptio complanata (Bivalvia: Unionidae), in Rhode Island:
A transplant experiment. Journal of the North American Benthological Society
Lauzon-Guay, J-S., D.J. Hamilton, and M.A. Barbeau. 2005. Effect of mussel density and
size on the morphology of Blue Mussels (Mytilus edulis) grown in suspended culture
in Prince Edward Island, Canada. Aquaculture 249(1–4):265–274.
Martins, I., R.P. Cosson, V. Riou, P-M. Sarradin, J. Sarrazin, R.S. Santos, and A. Colaço.
2011. Relationship between metal levels in the vent mussel Bathymodiolus azoricus
and local microhabitat chemical characteristics of Eiffel Tower (Lucky Strike). Deep
Sea Research Part I: Oceanographic Research Papers 58(3):306–315.
Negishi, J.N., and Y. Kayaba. 2009 Effects of handling and density on the growth of the
unionid mussel Pronodularia japanensis. Journal of the North American Benthological
Pilarczyk, M.M., P.M. Stewart, D.N. Shelton, H.N. Blalock-Herod, and J.D. Williams.
2006. Current and recent historical freshwater mussel assemblages in the Gulf coastal
plains. Southeastern Naturalist 5(2): 205–226.
Ricker, W.E. 1975. Computation and interpretation of biological statistics of fish populations.
Fisheries Research Board of Canada Bulletin 191:2–5.
San Miguel, E., S. Monserrat, C. Fernandez, R. Amaro, M. Hermida, P. Ondina, and C.
Altaba. 2004. Growth models and longevity of Freshwater Pearl Mussels (Margaratifera
margaratifera) in Spain. Canadian Journal of Zoology 82:1370–1379.
Schweers, T., M. Wolff, V. Koch, and F. Sinsel Duarte. 2006. Population dynamics of
Megapitaria squalida (Bivalvia: Veneridae) at Magdalena Bay, Baja California Sur,
Mexico. Revista de Biología Tropical 54(3):1003–1017.
Seed, R. 1968. Factors influencing shell shape in the mussel Mytilus edulis. Journal of the
Marine Biological Association of the United Kingdom 48:561–584.
Sickel, J.B., J.J Herod, and H.N. Blalock. 1997. Potential for the Kentucky Dam tailwater
of the Tennessee River to serve as a mussel refuge from invading Zebra Mussels.
Pp. 214-219, In K.S. Cummings, A.C Buchanan, C.A Mayer, and T.J. Naimo (Eds.).
Conservation and Management of Freshwater Mussels II: Initiatives for the Future.
Proceedings of a UMRCC Symposium, 1995 Oct 16–18. Upper Mississippi River
Conservation Committee, Rock Island, IL.
US Fish and Wildlife Service. 2012. Endangered and threatened wildlife and plants;
Determination of endangered species status for the Alabama Pearlshell, Round Ebonyshell,
Southern Kidneyshell, and Choctaw Bean; and threatened species status for
the Tapered Pigtoe, Southern Sandshell, and Fuzzy Pigtoe, and designation of critical
habitat; Final rule. Federal Register 77(196):61663–61719.
170 Southeastern Naturalist Vol. 12, No. 1
Widarto, T.H. 2007. Shell form variation of a freshwater mussel Velesunio ambiguous
Philippi from the Ross River, Australia. HAYATI Journal of Biosciences
Williams, J.D., M.L. Warren, K.S. Cummings, J.L. Harris, and R.J. Neves. 1993. Conservation
status of freshwater mussels of the United States and Canada. Fisheries
Williams, J.D., A.E. Bogan, and J.T. Garner. 2008. Freshwater Mussels of Alabama and
the Mobile Basin in Georgia, Mississippi, and Tennessee. The University of Alabama
Press, Tuscaloosa, AL. 908 pp.