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Ecology and Conservation of the

Diamond-Backed Terrapin Willem


Roosenburg
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Ecol­ogy and Conservation of the
Diamond-­backed Terrapin
This page intentionally left blank
ECOL­O GY and
CONSERVATION
of the
DIAMOND-­B ACKED
TERRAPIN
Edited by
Willem M. Roosenburg and Victor S. Kennedy

Johns Hopkins
University Press | Baltimore
© 2018 Johns Hopkins University Press
All rights reserved. Published 2018
Printed in the United States of Amer­i­ca on acid-­free paper
987654321

Johns Hopkins University Press


2715 North Charles Street
Baltimore, Mary­land 21218​-­4363
www​.­press​.­jhu​.­edu

Library of Congress Cataloging-­in-­Publication Data

Names: Roosenburg, Willem M., editor. | Kennedy, Victor S., editor.


Title: Ecol­ogy and conservation of the diamond-­backed terrapin /
Willem M. Roosenburg and Victor S. Kennedy, eds.
Description: Baltimore : Johns Hopkins University Press, 2018. |
Includes bibliographical references and index.
Identifiers: LCCN 2018000279 | ISBN 9781421426266
(hardcover : alk. paper) | ISBN 1421426269
(hardcover : alk. paper) | ISBN 9781421426273
(electronic) | ISBN 1421426277 (electronic)
Subjects: LCSH: Diamondback terrapin—­Ecology. | Diamondback
terrapin—­Conservation.
Classification: LCC QL666.C547 E26 2018 | DDC 597.92/59—­dc23
LC rec­ord available at https://­lccn​.­loc​.­gov​/­2018000279

A cata­log rec­ord for this book is available from the British Library.

Special discounts are available for bulk purchases of this book. For
more information, please contact Special Sales at 410-­516-­6936 or
specialsales@press​.­jhu​.­edu.

Johns Hopkins University Press uses environmentally friendly


book materials, including recycled text paper that is composed of
at least 30 ­percent post-­consumer waste, whenever pos­si­ble.
To Bill and El, who showed us that science and friends make
good partners, and to Kate and Deb for unstinting support
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Contents

Preface ix

1 Introduction and History 1


J. Whitfield Gibbons

PART I BIOLOGY AND ECOL­O GY

2 Capture, Mea­sure­ment, and Field Techniques 7


Willem M. Roosenburg and Russell L. Burke

3 Evolutionary History and Paleontological Rec­ord 27


Dana J. Ehret and Benjamin K. Atkinson

4 Taxonomy: A History of Controversy and Uncertainty 37


Jeffrey E. Lovich and Kristen M. Hart

5 Molecular Ecol­ogy and Phylogeography 51


Paul E. Converse and Shawn R. Kuchta

6 Life History with Emphasis on Geographic Variation 63


Jeffrey E. Lovich, J. Whitfield Gibbons,
and Kathryn M. Greene

7 Reproductive Be­hav­ior and Ecol­ogy 81


Joseph A. Butler, Russell L. Burke,
and Willem M. Roosenburg

8 Hatchling Be­hav­ior and Overwintering 93


Patrick J. Baker, Ralph E. J. Boerner,
and Roger C. Wood

9 Osmoregulation 111
Leigh Anne Harden and Amanda Southwood Williard
10 Temperature-­Dependent Sex Determination 127
Thane Wibbels, Taylor Roberge, and Allen R. Place

11 Foraging Ecol­ogy and Habitat Choice 147


Anton D. Tucker, Russell L. Burke, and Diane C. Tulipani

12 Environmental Toxicology 161


Dawn K. Holliday, Rusty D. Day, and David Owens

PART II FISHERIES AND CONSERVATION CHALLENGES

13 History of Commercial Fisheries and Artificial


Propagation 187
Victor S. Kennedy

14 Conservation in Terrestrial Habitats:


Mitigating Habitat Loss, Road Mortality,
and Subsidized Predators 201
John C. Maerz, Richard A. Seigel, and Brian A. Crawford

15 Interactions with Motorboats 221


Lori A. Lester, Harold W. Avery, Andrew S. Harrison,
and Edward A. Standora

16 Bycatch in Blue Crab Fisheries 231


Randolph M. Chambers and John C. Maerz

17 Conservation through Environmental Education 245


George L. Heinrich, Timothy J. Walsh, and ­Will Williams

18 Habitat Restoration and Head-­starting 253


Willem M. Roosenburg

19 The ­Future for Diamond-­backed Terrapins 265


Joseph A. Butler and Willem M. Roosenburg

Contributors 269
Index 271

viii  Contents
Preface

The diamond-­backed terrapin has had an impor­tant role


in North American history as a ­human food source that developed
­great economic value but quickly waned due to overexploitation.
Watermen and entrepreneurs, first in Chesapeake Bay and then through-
out the species’ range, made the terrapin an epicurean delicacy in
the late nineteenth and early twentieth centuries. The resulting har-
vest pressure on a species with a life history of slow growth, delayed
maturity, and low fecundity resulted in range-­wide population de-
clines, starting in Chesapeake Bay and expanding to other coastal
regions, as demand outpaced supply. By the end of the nineteenth
­century, stocks w­ ere depleted greatly, and Prohibition and the elimi-
nation of sherry as the key ingredient in terrapin soup helped reduce
the fishery. In the modern day, terrapins encounter numerous an-
thropogenic threats ranging from toxins to capture in commercial
crab pots and habitat loss due to development of coastal regions.
However, t­ here is some pro­gress, as most states have recognized the
unsustainability of terrapin harvesting and have banned commercial
take. Thankfully, through research and outreach, terrapin conserva-
tion is making slow and steady headway that generates optimism
among ­those who study t­ hese turtles that they w
­ ill be h
­ ere for f­ uture
generations. This book reviews the rapidly growing body of lit­er­a­
ture that contributes to our understanding of terrapins throughout
their range. We hope it ­will serve as a reference for terrapin enthusi-
asts and researchers, but also for ­those interested in turtles in
general.
The book also represents a shared life-­long interest in Chesapeake
Bay, its denizens, and their natu­ral history. The terrapin was a spe-
cies much celebrated but ­little known in the Bay ­until Willem Roosen­
burg began gradu­ate school at the University of Pennsylvania in
1986. But the beginnings of this book precede his start in gradu­ate
school by 20 years, when Vic Kennedy was him- mental in initiating this pro­cess, which Tiffany
self a gradu­ate student working at the Hallowing Gasbarrini saw through to completion. We are
Point Field Station of the University of Mary­land grateful to the authors who wrote chapters and
Chesapeake Biological Laboratory. Willem Roosen- ­were patient with our comments and editorial
burg, Sr., collaborated with Vic on several studies of changes. We also are grateful to the many review-
the effects of thermal addition to the Patuxent es- ers who provided detailed comments for individ-
tuary, and Willem Jr. accompanied his dad to work ual chapters, including Kimberly Andrews, Aaron
during summer vacations to fish, crab, and partici- Baxter, Donna Marie Bilcovic, Ruth Boettcher,
pate in the science underway at the field station. Bruce Bury, John Byrd, Kristen Cecala, Andrew
Willem and Vic remained in touch over the years Coleman, Justin Congdon, Paul Converse, Jon
as their ­careers unfolded, eventually agreeing to Costanzo, John Davenport, Peter Paul van Dijk,
unite Willem’s expertise on terrapins with Vic’s Michael Dorcas, Whit Gibbons, George Heinrich,
experience serving as a volume editor to produce John Iverson, Dale Jackson, Peter King, Chris Ma-
this book. nis, Suzanne McGaugh, Brian Mealey, Timothy
Thirty years ago, in the summer of 1987, Willem Muir, James Parham, Greg Pauly, Turk Rhen,
began one of the few long-­term studies of terrapins Christopher Rowe, Rich Seigel, Kyle Selcer, W ­ ill
and the first of its kind in Chesapeake Bay. Al- Selman, Arun Sethuraman, Brad Shaffer, James
though in most other areas of their range terrapins Spotila, David Steen, Tony Tucker, Gordon Ultsch,
are cryptic and prove challenging to catch, in Ches- and Richard Vogt. Copy editor Linda Strange did
apeake Bay they ­were common in some areas and an excellent job of ensuring that issues of style
proved to be relatively easy to catch with the right and format ­were attended to, as well as ferreting
gear. Over the years he has worked to develop the out errors and inconsistencies in the narratives.
terrapin as a model species for the study of turtle We thank Elizabeth Freedlander, of Horn Point
life history, temperature-­dependent sex determi- Laboratory, who helped raise funds to support
nation, chelonian conservation, and more. publication of this book and the private donors
that contributed to helping defer publishing costs.
Many individuals have been instrumental in the Fi­nally, we are grateful to our families—­Deb, Kate,
development and completion of this book. Vincent Dirk, and Selinde—­who have been patient and
Burke, formerly of Johns Hopkins University Press, supportive as we usurped time from other activi-
came to Willem with the idea and the desire to pub- ties to complete the text.
lish a book on terrapins. His guidance was instru-

x  Preface
Ecol­ogy and Conservation of the
Diamond-­backed Terrapin
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1
J. WHI TFI ELD GI B B O N S
Introduction and History

M any researchers proclaim that their study species has quali-


ties that set it apart from other species and make it the “ideal
model” for research pursuits. Some even define their study organism
as “unique,” which becomes meaningless when considering that
­every species on earth is unique when we take into account its evo-
lutionary history and current ge­ne­tic makeup coupled with its geo-
graphic range, ecol­ogy, and life history traits. Nonetheless, a primary
feature that sets the diamond-­backed terrapin apart from all other
turtles is that it resides permanently in brackish ­water. No other tur-
tle can claim more than a passing association with the relatively thin
interface between the full saltwater of the ocean and the freshwater
lakes and tributaries associated with the mainland on ­every turtle-­
inhabited island, isthmus, and continent. Other turtles also are
called “terrapins,” but of the 335 recognized extant species, all but
the diamond-­backed terrapin live mostly on one side or the other of
the brackish ecotone. Thus, the diamond-­backed terrapin truly is
unique among turtles in making this habitat its home.
The history of the species can be viewed from many perspec-
tives. Its evolutionary history began sometime in the Miocene, a
half-­dozen to dozen million years ago, when some freshwater an-
cestor began to thrive in an unexploited niche that excluded com-
petitors. Its greatest biological competitors resided in fresh or ma-
rine w­ aters but not in between. The historical importance of the
species to h ­ umans began a few thousand years ago, when Native
Americans living along the mid-­Atlantic coastal regions of North
Amer­i­ca first began using terrapins as a food source, along with
freshwater turtles on the mainland. When Eu­ro­pean settlers reached
the New World, having never experienced an estuarine turtle, they
simply called them “torope,” the moniker be- threats. This book adds considerably to our
stowed by the Algonquian Indians of the region. knowl­edge of the diamond-­backed terrapin and
The scientific history of the species began in discloses how many capable researchers are now
1793, the year that George Washington held his first involved in acquiring the critical facts needed
cabinet meeting as president of the United States for its conservation.
and Johann David Schoepff described the species One of the first practical questions someone
and gave it its initial binomial, Testudo terrapin. In ­doing ecological research might ask is, how do you
1844, the year James K. Polk defeated Henry Clay catch a terrapin and what do you do once you have
for the presidency, John Edward Gray placed it in hand? The history of techniques for catching
the terrapin in the correct genus, Malaclemys. But turtles extends into the distant past to the Old
turtle taxonomy of the times was muddled and World, before the earliest settlers arrived on the
convoluted, and Gray’s contribution distinguished banks of the Atlantic and Gulf of Mexico where ter-
terrapins from Graptemys but did not recognize rapins thrived. No single capture technique can be
Malaclemys as a monotypic genus. The current sci- deemed “the best,” for it depends on season and cir-
entific name, Malaclemys terrapin, was not used cumstances of when turtles w ­ ill be where. The
­until Outram Bangs put the two together in 1896, collection of thousands of adults during the Kiawah
the last year of Grover Cleveland’s second term Island studies speaks to the advantage of basic sein-
in office. How many of ­these po­liti­cal figures liv- ing and setting trammel nets across tidal creeks.
ing in Mary­land and ­Virginia dined on terrapins is But many other approaches have been effectively
unknown, but Cleveland’s first term was marked employed, before and ­after. The cumulative num-
by the publication of The White House Cookbook. ber of techniques for catching turtles that have
Among the featured ­recipes was “Stewed Terra- been published in books, papers, and scientific
pin with Cream,” a foreboding of the conservation notes, when added to t­ hose that have been used
prob­lems that lay ahead for the diamond-­backed and not published, is enormous. A ­ fter capturing a
terrapin. Terrapins are no longer thrown alive into turtle, the researcher must of course focus on appro-
a pot of boiling ­water, as recommended in the priate mea­sure­ments, which may include the use
­recipe, but drowning in a crab pot could be viewed of digital photography and collection of molecular
as an equally gruesome fate. Have we solved one data. Virtually all standardized techni­ques that a
prob­lem of terrapin conservation while being un- researcher needs to know for studying terrapins
able to curtail another? are referred to in this book.
Our current knowledge of the ecol­ogy and con- How did terrapins come to be one of the few tet-
servation of terrapins began in the early 1900s with rapods in the world to become permanent resi-
classic papers by Robert Coker and Samuel Hil- dents in a singular habitat that has existed on earth
debrand that addressed both the natu­ral history of since elevated land masses first appeared? How
terrapins and their commercial use. Since the pre–­ does an organism live in the brackish interface be-
World War II work by Coker, Hildebrand, and tween freshwater runoff and the sea? An obvious
­others, most research on terrapins has been pub- consideration for organisms living in habitats with
lished in the two de­cades since the mid-1990s. The high salinity is the necessity for physiological ad-
longest continual population studies w­ ere initiated justments to allow salt removal. Osmoregulation by
on Kiawah Island, South Carolina, in 1983. Other terrapins has thus been of some research interest.
studies soon began to encompass a broader portion Yet, approximately half of the published research
of the geographic range. The breadth of questions focusing on salt elimination and electrolyte balance
on ecol­ogy, physiology, and be­hav­ior expanded, expressly in M. terrapin was completed before the
and the work is now escalating at an impressive mid-1970s. Chapter 9, on osmoregulation, draws on
rate. Conservation remains a central theme in ­these earlier works and more recent publications to
many of t­ hese efforts—as it should for such ele- place terrapins in the context of other reptiles for
gant animals that face so many human-­caused which salt elimination is a critical physiological

2   J. W. Gibbons
i­ssue. The consolidation of what was previously young hatching in less than three months. Hatch-
known, complemented by current research find- lings usually emerge in autumn and sometimes
ings, ­will serve as a baseline for ­future research ­remain in the nest cavity or in the surrounding
into the behavioral and physiological strategies habitat for the duration of fall and winter. In e­ ither
­required for M. terrapin to survive in an estuarine case, ­after entering the aquatic habitat they begin
habitat. feeding, approximately one year ­a fter being de-
Into how many genet­ically identifiable units can posited as eggs; ­ whether this is true for the
M. terrapin be partitioned? Many North American southernmost populations in Florida remains
turtles that systematists accepted as single species equivocal. Growth rates are presumably influential
for de­cades, if not centuries, have now been sepa- in the attainment of maturity, and like many
rated into two or more species by warriors armed emydids and trionychids, the males in a region tend
with ge­ne­tic tools. Malaclemys terrapin has the op- to reach maturity at a smaller size than females,
posite situation: no one is proposing that more which remain measurably larger as adults. Sexual
than one species is represented. In fact, separation dimorphism is unequivocal, but are growth rates
into the seven putative subspecies traditionally re- among individuals within some populations faster
ported is now challenged—­ a refreshing move than ­t hose in ­others, as observed in some turtles?
­toward lumping rather than splitting. Discussions If so, do differential growth rates affect age at ma-
of ge­ne­tic variation, phylogeography, paleontology, turity and ultimate adult size? Age-­size relation-
and taxonomy and systematics in this book all con- ships to maturity in the sexes are highly complex
tribute to current and ­future understanding of the in turtles. Many ecological questions are expertly
evolutionary history of this linearly distributed addressed in the chapters on ecol­ogy, including
species that has no natu­ ral barriers between nesting and overwintering of hatchlings, foraging
present-­day populations. Collectively, all topics be­hav­ior and diet, environmental sex determina-
pertinent to the phylogeny, ecol­ogy, and conserva- tion, and basic life history. Refining our under-
tion of the diamond-­backed terrapin are addressed standing of ­these ecological f­actors for terrapins
herein. ­will be an exciting challenge.
Geographic variation is evident among terra- Effective conservation programs historically
pins—hence the early descriptions of subspecies— have involved not only scientific study of the tar-
but patterns of consistency and regional conformity get species but complementary education of the
have been elusive, even for the few ge­ne­t ic pro- public, who elect the officials that can champion a
files that have been accomplished. One of the cause with legislation. Both approaches ­will be cru-
enigmas surrounding the distribution pattern of cial to ensuring that productive populations of ter-
terrapins that continue to intrigue investigators is, rapins ­will persist, or in many cases recover, from
how did terrapins get to Bermuda? This issue is Mas­sa­chu­setts to Texas. This book does an admi-
addressed and opinions, supported by data, are rable job of covering both categories.
offered in this book. In taking a position that ter- Among the conservation chapters is an excellent
rapins reached Bermuda by natu­ ral dispersal, overview of the effect that overharvesting has had
should we also ask why terrapins are not found in historically on ­these easy to find and catch animals
Cuba’s much closer tidal brackish habitats, to which (Chapter 13). Terrapins ­were the first reptile species
US salt marsh snakes (Nerodia clarkii) success- in Amer­i­ca whose steady and alarming decline in
fully emigrated? The Bermuda terrapin case is not the wild can be attributed to commercial exploita-
closed, and hints of titillating debates are on the tion for food. An account of the astounding num-
horizon. ber of ­recipes for preparing terrapin and the list of
Like the life cycles of freshwater turtles in tem- restaurant menus on which they appeared provides
perate regions everywhere, the basic life cycle of M. sober testimony on how quickly a common, wide-
terrapin is one in which eggs are laid in early spring spread species can decline in numbers when conser-
in a terrestrial nest dug by the female, with the vation programs and controls are not properly

Introduction and History   3


administered. ­Human disregard for the welfare impacts, salt marsh development, and other ­human
of individual terrapins was commonplace, and activities that vary regionally is also addressed and
the population status of t­hese animals was not appropriately emphasized. Research leading to
taken into consideration ­until their numbers and ­scientifically sound conservation approaches for
availability noticeably began to diminish. Even beleaguered species such as terrapins must also
then, government programs that ultimately led to include restorative efforts to replace and sustain
artificial propagation through terrapin farming what has been lost. Discussion of a variety of
­were designed more to ensure a continuing supply ­regulatory approaches, including harvest limita-
of the popu­lar dietary item than to lessen the rate tions, head-­starting, and control of nesting area
of dwindling numbers and protect natu­ral popu- predation, provides the background necessary to
lations. Understanding the pro­cess of how public pursue t­ hese conservation tactics.
attitudes can drive a species from abundance to Attention must be paid to all of ­these issues if
rarity in only a few de­cades is impor­tant. The ter- effective conservation programs are to be imple-
rapin story offers insight that can be applied to mented that w ­ ill return terrapins to their natu­ral
­developing effective conservation practices for state throughout their range and maintain them
any overexploited species. both ecologically and behaviorally. Fi­nally, the im-
A conservation program that is scientifically portance of environmental education in concert
based is key to the ultimate success of any species with terrapin conservation programs is highlighted
struggling ­under human-­caused environmental in Chapter 17.
threats, and the terrapin serves as a model for other The chapters on conservation threats that im-
species, especially turtles. Descriptions of the peril the existence of terrapins throughout their
many dangers terrapins face ­because of commer- range, coupled with recommendations for educa-
cial development and activities in coastal habitats tional approaches, ­will serve as a vanguard for
are telling about the potential for continued de- anyone wanting to aid in the protection of this pre-
clines. Collectively, ­these chapters set the stage for possessing species. Continuation of the history of
much-­needed ­future research. Loss of terrapins as this venerable turtle ­will be marked by what is
bycatch from both recreational and commercial ­reported in this book, which w ­ ill unequivocally
crab trapping, as well as other fisheries activities, be recognized as a milestone by t­hose pursuing
is clearly a key area of concern. Injuries and deaths ­future studies on the diamond-­backed terrapin’s
caused by outboard motors in terrapin w ­ aters are ecol­ogy and conservation.
documented and need attention. Road systems sep-
arating nesting habitats from aquatic areas are a
ACKNOWL­E DGMENTS
bane to many turtle species, including terrapins.
The value of examining geographic variation in the I appreciate the reviews of drafts of this chapter by
responses of terrapin populations to agricultural Jeff Lovich, Willem Roosenburg, and Vic Kennedy.

4   J. W. Gibbons
PART I BIOLOGY AND ECOL­O GY
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2
W ILLE M M. RO O SEN B U RG
Capture, Mea­sure­ment, and
Field Techniques
RUSSELL L. BU RKE

T urtles are compelling animals to study b ­ ecause they are charis-


matic, many species are easy to catch, and their life histories are
both diverse and unusual. Turtles also serve as model long-­lived
organisms and are the focus of several renowned long-­term ecologi-
cal studies. The turtle’s shell allows the use of s­ imple and inexpen-
sive methods to mark and identify individuals (Cagle 1939) and is an
excellent platform for attaching recording and transmitting devices.
­Simple scientific tools (balances, calipers, and a file) combined with
hard work and per­sis­tence have resulted in many excellent popula-
tion studies of a variety of turtle species (reviewed in Wilbur and
Morin 1988), and more recent studies include the diamond-­backed
terrapin (Malaclemys terrapin). However, the population ecol­ogy of ter-
rapins has unique and in­ter­est­ing challenges when compared with
that of most freshwater turtles. To supplement two recent reviews
of techniques for studying reptiles (McDiarmid et al. 2012; Graeter
et al. 2013), we review h­ ere the specialized field techniques used with
terrapins.
Comprehensive studies (­those that catch more than just nesting
females) require substantial investment in boats and traps and the
tolerance of pesky or potentially dangerous bycatch that can include
alligators, alligator gar, blue crabs, sharks, sting rays, and sea nettles.
Thorough mark-­recapture studies should employ a variety of capture
techniques to minimize potential bias associated with dif­fer­ent
trapping methods (Roosenburg et al. 1997) and to sample a range of
estuarine habitats, life cycle stages, and sexes (Roosenburg et al. 1999).
Daily tidal fluctuation requires precautions to ensure that terrapins
have access to air if they remain in traps for more than a few hours.
­Because of the variation in tidal amplitude (from <1 to >3 m) and the
diversity of habitat/marsh structure throughout the terrapin’s
range, techniques that work well in some areas bait is not required. Fykes with two wings can be
are not as effective in o ­ thers. Fi­nally, terrapins set parallel to the current in small tidal creeks,
can be sympatric with other turtle species, includ- blocking the entire channel, but ­these ­will catch
ing sea turtles, snapping turtles, kinosternids, and only with the tidal flow entering the funnel (Sel-
emydids, that may be caught in the same gear man and Baccigalopi 2012; Selman et al. 2014).
used for terrapins. Fyke nets require several modifications to catch
We describe a set of methods for catching ter- terrapins. First, if the cod end is submerged, terra-
rapins and a standardized set of mea­sure­ments pins ­will drown when held underwater for more
to facilitate comparability of data among studies. than 2 h in warm w ­ aters. Thus, a daily tidal ampli-
We then describe successful techniques to catch, tude of <1 m and a w ­ ater depth of <2 m are ideal.
mark, mea­sure, and track terrapins; highlight some Stakes or metal poles are preferred over the tradi-
methodological issues that require attention; and tional anchors ­because they can hold the cod end
introduce some new techniques with g­ reat poten- up to help maintain an airspace. Tying 1 or 2 large
tial for terrapin research.

A
Trapping and Capture Methods
Checking traps and catching turtles can be the
most exciting part of a population study. Traps that
catch terrapins also reveal the diversity of their
habitat by catching crabs, fish, other turtle species,
and even sharks and rays. We describe ­here ways
to catch terrapins (see Chapter 13 for an account of
methods used in the early fisheries). Although our
list is fairly comprehensive, we recommend asking
local commercial fishermen both where and how
to catch terrapins when initiating a study. Their ex-
perience can help identify the techniques that B
work best at your study site.

Fyke or Hoop Nets


Fyke nets are effective devices to catch riverine
turtles in large open bodies of w ­ ater or near nest-
ing beaches (Vogt 1980). They consist of a series of
mesh nets (a.k.a. hedgings or wings) attached to
several hoops (fiberglass is preferred), nested within
one another, and usually a double-­throated mesh
Fig. 2.1. (A) Fyke net. Stakes instead of anchors are used to
trap. They are most effectively configured with a
hold the net in place. The float in the cod end of the net
central lead deployed perpendicular to the shore- maintains a permanent air space so that terrapins have
line starting near shore, as well as two wings set at access to air. The net is set perpendicular to the shoreline,
a 45° ­angle to the central lead, forming a funnel to and the wings form a V shape that directs the catch to
the trap (cod end) (Fig. 2.1, A). Most mobile estuarine the cod end. (B) A bank trap (peeler pound) is similar in
princi­ple to a fyke net but with a net made of galvanized
species, including terrapins, move parallel to the
chicken wire. The wings are modified to a heart-­shaped
shoreline, and when they encounter an obstruc- structure (the pound), and the cod end is a ­simple box that
tion such as the central lead they move offshore to maintains an airspace above mean high tide. (B) Courtesy of
get around it, thus being directed to the cod end; ­Virginia Institute of Marine Science

8   W. M. Roosenburg and R. L. Burke


floats in the cod end w
­ ill also maintain the airspace. rapin aggregations and then pulled the net ­toward
Second, a net mesh size of 2.5 cm should be used in shore like a seine or pounded the ­water with pad-
areas with high blue crab densities b ­ ecause larger dles or buckets to drive the terrapins ­toward the
meshes entangle blue crabs, which damage the net, net. Nets should be removed within 2 h to prevent
requiring mending. Dipping the webbing in net terrapins from drowning if they get entangled near
dip stiffens it and reduces crab damage. Floats the bottom. A 7.5 cm mesh is very effective for ter-
and weights on the lead(s) keep the nets vertical in rapins. T
­ hese nets w­ ill generate bycatch of blue
the ­water column, both preventing terrapins from crabs and fish that can be difficult to untangle; in
swimming u ­ nder or over the net and further re- one instance, we caught 17 cow-­nosed rays (Rhinop­
ducing crab damage. Third, nets should be checked tera bonasus) along with 5 terrapins.
at least once a day to ensure that the airspace is
available and to reduce stress on entrapped ter-
Turtle or Three-­Ring Hoop Traps
rapins and other organisms; less frequent check-
ing ­will increase bycatch mortality. Fyke nets can Turtle traps work well with numerous species of
be easily moved between locations and should be freshwater turtles (McDiarmid et al. 2012) but are
cleaned and dried once ­every 5 to 10 days to pre- not as effective for terrapins. Traps typically are
vent the buildup of estuarine fouling organisms, set in shallow w­ ater to maintain a permanent air-
which reduces a net’s effectiveness. space and are baited with fish, crabs, or clams for
terrapins (unbaited traps have caught terrapins in
Barnegat Bay; H. Avery, Drexel University, pers.
Bank Traps or Peeler Pounds
comm.). They can be successful in narrow, shallow
Bank traps or peeler pounds (Fig. 2.1, B) are essen- marsh channels or while set submerged, but they
tially fyke nets made of galvanized chicken wire. must be checked ­every 3 h to avoid terrapin drown-
They are not commercially available and require ing. The traps catch better during the early spring
about 4 to 6 h to build. However, they are consid- when terrapins are emerging from their hibernac-
erably cheaper than fyke nets and are immune ula and are attracted to bait.
to damage by blue crabs. They require a substantial
effort to set and therefore are practical only in
Crab and Eel Pots
long-­term trapping (>2 weeks) proj­ects. Terrapin
drowning is prevented by building the trap’s box The ability of wire-­mesh crab or eel pots to capture
to be at least 30 cm above the mean high tide. terrapins is of conservation concern (reviewed
Bank traps exploit the movement of animals par- in Roosenburg 2004; see also Chapter 16). How-
allel to the shoreline and thus do not need bait. ever, ­these pots can also catch terrapins for study
Checking of bank traps requires a boat and a (e.g., Davis 1942; Bishop 1983; Roosenburg at al.
strong back or winch to lift the boxes. 1997; Hoyle and Gibbons 2000; Avissar 2006). The
use of short pots that remain submerged requires
checking ­every 3 h, but in areas of low tidal ampli-
Gill or Trammel Nets
tude, tall crab pots (Roosenburg et al. 1997) can be
Both gill and trammel nets entangle terrapins un- used to catch terrapins and need to be checked
derwater and therefore must be attended while only once per day. The tall traps can be built by
they are fished. They can catch terrapins in a vari- cutting the bottom and top out of two pots and us-
ety of habitats (e.g., Seigel 1984; Simoes and Cham- ing hog rings to attach the two together. Tall crab
bers 1999; Levesque 2000; Broyles 2010) but are pots (Fig. 2.2) should be tied to a stake to prevent
most effective when terrapins are aggregated in strong wind or waves from knocking them over.
open ­water near basking or nesting areas, or in Care should be taken to cut holes in the “church”
tidal creeks during ebb tide (D. Owens, College of that leads to the top section of the pot so that terra-
Charleston, pers. comm.). We have surrounded ter- pins can retreat to the ­water during low tide. Crab

Capture, Mea­sure­ment, and Field Techniques   9


Fig. 2.2. Comparison of
standard and tall crab
pots. The tall crab pot
contains a second
“church” (or baffle) with
upward and downward
funnels that allow
terrapins to move up
and down with the rise
and fall of the tide.
Courtesy of Conserva-
tion Biology

pots checked regularly can be used in most habitats (Hurd et al. 1979); smaller trawls are most effective
and can be baited with fish, crabs, or clams. in small tidal creeks where their gape can span the
We have used round wire traps that are easier width of the creek. The trawls should be emptied
and cheaper to build than crab pots. They are ap- regularly (­every 20 min) to prevent drowning ter-
proximately 1 m in dia­meter and 1.33 to 1.67 m tall rapins or damaging them with accumulated debris
and are made of 2.54 × 5.08 cm wire built on a frame and bycatch. Fishing with otter trawls requires
of old fyke net hoops. They stand on end and have attention to the position of the “doors,” as their
a single funnel on the side, near the bottom. Baited proper deployment is critical to the net fishing
traps catch juvenile terrapins in small guts and tidal correctly.
creeks late in the season.
Eel pots are another commercial gear that
Seines
catches terrapins as bycatch. Cloth-­funnel eel pots
baited with razor clams capture terrapins at high In the absence of a boat with a power­f ul motor,
rates in creek and nearshore river habitats, but they hand seines can effectively catch terrapins in small
fish submerged and thus must be checked ­every 3 h tidal creeks (e.g., Grosse et al. 2011). This requires
(Radzio and Roosenburg 2005). walking the banks of tidal creeks, where research-
ers can sink deeply in mud and have their shoes
sucked off. Seines are most effective when pulled
Otter Trawls
against the out­going tide with a net that spans the
Otter trawls are tapered mesh nets with two wings width of the creek. A bag in the seine ­will reduce
attached to “doors” that spread the net while being the escape of terrapins, and a mesh size between
towed. They require a motor boat to pull the net at 1.25 cm and 2.5 cm w ­ ill ensure the capture of most
a speed of 3 to 5 knots to prevent terrapins from size classes. Dorcas et al. (2007), Gibbons et al.
avoiding or swimming out of the trawl. Larger (2001), and Grosse et al. (2011) have seined in the
trawls can be effective in open ­water with high narrow channels of salt marshes that lie b ­ ehind the
tidal amplitude or where terrapins are concentrated barrier islands of South Carolina and Georgia.

10   W. M. Roosenburg and R. L. Burke


all of the hatchlings within a nest (Roosenburg
Terrestrial Drift Fences
et al. 2014). Predator excluders (Riley and Litzgus
The traditional herpetological technique using ter- 2013; Roosenburg et al. 2014) are devices designed
restrial drift fences (Graeter et al. 2013) can capture to protect developing eggs from predators and can
adult female terrapins when they come ashore to catch hatchlings if left in place long enough. Rings
nest (Coleman 2011; Roberge 2012) and hatchlings or excluders should be buried a minimum of 2.5 cm
as they emerge from nests (Muldoon and Burke and centered on the nest. Hardware cloth (1.25 cm
2012). Fences can be made of metal flashing, plas- mesh) placed over the rings can prevent avian pre-
tic UV-­resistant screening, or even corrugated pipe dation of emerging hatchlings. On hot days (>29°C),
(P. Baker, Wetlands Institute, pers. comm.). Typi- rings should be checked several times daily or
cally, buckets (or smaller containers for hatch- shade should be provided to prevent hatchling
lings) are sunk in the sand along the fence to catch mortality from overheating or desiccation. Rings
the animals as they search for passage across the and predator excluders may influence nest temper-
barrier. The hot conditions on most nesting areas atures and/or nest moisture, but this has not been
require that the fences be checked several times per explored sufficiently (Riley and Litzgus 2013).
day to prevent mortality due to overheating. This
is particularly true for hatchlings, which are vul-
Rolling or Raking Wrack
nerable to desiccation ­ because of their smaller
size. Wet sponges can be placed in the bottoms of Rolling or raking wrack uses a rake or cultivator to
buckets to reduce the risk of hatchling desiccation, ­gently roll the accumulated wrack at the high-­tide
and covers over buckets can protect terrapins from line in salt marshes. Wrack can also be lifted by
sun exposure (Roberge 2012). hand (Pilter 1985). Hatchling and juvenile terrapins
are active ­under the wrack, which is lifted above
the substrate by emerging grasses, and this may be
Nest Rings and Predator Excluders
impor­tant terrestrial habitat in their life cycle. King
Nest rings are 50 to 100 cm dia­meter rings made of (2007) found hatchling terrapins in both low and
flashing (Fig. 2.3) that catch hatchlings emerging high Spartina marsh by searching ­under debris and
from the nest; they are useful to determine within-­ vegetation. In Chesapeake Bay, this technique
nest survivorship ­because they can account for found more young of the previous year ­under

Fig. 2.3. Nest ring


encircling emerged
terrapin hatchlings.

Capture, Mea­sure­ment, and Field Techniques   11


wrack over Spartina patens than that over S. alterni­ (Wetlands Institute, pers. comm.) followed tracks
flora (Roosenburg, pers. obs.). Raking wrack can left in the mud of tidal ponds and mud flats to catch
catch juveniles and adults in some Florida marshes terrapins by hand. Butler (2002) reported using cast
and can yield more than 60 terrapins per day (B. nets for terrapins, but this required considerable
Turner, Florida Fish and Wildlife Conservation skill and had limited success. Dip netting is used to
Commission, pers. comm.). capture many turtle species (Graeter et al. 2013) but
is not commonly used for terrapins. However, Hart
and McIvor (2008) dip-­netted mangrove terrapins
Winter Dredging or Scraping
during a 2 h win­dow around diurnal and noctur-
Winter dredging was developed by commercial wa- nal low tides. Hibernacula in shallows (<1 m) also
termen in Chesapeake Bay and then a­ dopted by can be searched for depressions on the bottom, and
scientists to catch terrapins (Haramis et al. 2011). the terrapins can be excavated using a dip net from
A crab scrape modified to dig hibernating terrapins a boat (D. Marshall, Smith Island waterman, pers.
from the mud is pulled along the bottom by a comm.).
power­ful boat. The dredge is 1 to 1.3 m wide and
has a twine net suspended from a metal frame. The
Census Techniques
bottom of the frame has 10 to 15 cm teeth that dig
terrapins from the mud and into the net. Dredging Most of the capture techniques described above
requires knowledge of hibernacula where terrapins lend themselves well to standard mark-­recapture
are concentrated (Haramis et al. 2011); catch rates analyses of populations (Hurd et al. 1979; Hoyle and
can exceed 60 terrapins per hour. Haramis et al. Gibbons 2000; Tucker et al. 2001; Mitro 2003;
(2011) also tested post-­catch mortality and docu- Witczak et al. 2014). However, some proj­ects do not
mented 100% survival of animals disturbed from require precise estimates of population size but
their hibernaculum and returned to the ­water on instead focus on site occupancy (presence/absence)
the same day. or relative abundance. Several techniques are avail-
able for ­these purposes.
Miscellaneous Capture Techniques
Cyst Counts
Other techniques that catch terrapins include
SCUBA, hand capture, and dip netting. In areas Terrapins are the unique obligate final host for the
with clear ­water or in colder w­ aters where terra- small parasitic trematode Pleurogonius malaclemys
pins do not move quickly, snorkeling and SCUBA (Hunter, 1961, 1967; Chodkowski et al. 2016). The
diving can yield terrapins (Morreale 1992). This parasite’s intermediate host is a common terrapin
method can be supplemented with the use of dip prey species, the eastern mudsnail (Ilyanassa obso­
nets to extend the reach of the catcher. However, leta). One of the juvenile stages is a <1 mm metacer-
SCUBA is in­effec­tive in most estuaries throughout carial cyst that is affixed to the outside of the snail
the terrapin’s range b
­ ecause of low visibility in tur- (Fig. 2.4), where it remains ­until the snail is eaten
bid ­waters. Collection of nesting terrapins by by a terrapin. Mudsnails are easily collected along
hand is employed by many researchers (e.g., Butler the beach at low tide, and the number of cysts cor-
2002; Feinberg and Burke 2003), and in many stud- relates well with local terrapin population size in
ies hand capture is the sole method of capture. This coastal Georgia (Byers et al. 2011), providing esti-
usually involves ­either patrolling nesting areas or mates of terrapin populations. Counting cysts is
watching from an inconspicuous location while fe- quick and has the potential for low-­cost terrapin
males come ashore. This approach requires no population estimates. Many questions remain
equipment but a knowledge of nesting areas and throughout the terrapin’s range, however, such as
times. Selman and Baccigalopi (2012) and R. Wood how far the trematodes disperse, the trematodes’

12   W. M. Roosenburg and R. L. Burke


Crawl and Nest Surveys
When only presence/absence data are needed, the
simplest technique for identifying terrapin popula-
tions is to explore likely nesting areas and search
for tracks and intact or depredated nests and their
egg shells. Terrapin tracks are easily distinguished,
but intact nests are more difficult to locate. Wind,
rain, and dry sand w ­ ill obscure tracks and nests
within 24 h, making it virtually impossible to de-
tect the presence of terrapins. Butler et al. (2004)
documented crawls and nests on Florida beaches
Fig. 2.4. Mud snail with Pleurogonius malaclemys meta­ where they suspected terrapin presence. Where
cercarial cysts, indicating that a terrapin population is nearby. raccoons and foxes are common, nest predation
The arrows indicate cysts on the snail’s operculum and rates are high. Depredated nests are easily identi-
around the aperture. fied by egg shells, which can remain on nesting
beaches for more than a year. In areas where nest
range (to both north and south), and the strength predators are absent (Roosenburg et al. 2014) or
of the relationship between cyst counts and popu- predators consume egg shells (Burke et al. 2009), de-
lation size. Burke’s laboratory is exploring ­t hese tection probabilities decrease.
questions in New York with a pi­lot program involv-
ing citizen scientists who are collecting snails and
Marking Terrapins
counting cysts throughout the Northeast.
Turtles are easy to mark (Graeter et al. 2013), and
several identification technologies, each with its
Visual Counts
advantages and limitations, can be incorporated
Visual counts of basking and swimming terrapins, into demographic studies. We highly recommend
taken from kayaks, canoes, and skiffs up to 8 m in using a double tagging system when pos­si­ble, so
length (Butler 2002; Harden et al. 2009) or from that tag loss can be confirmed—­for example,
shore (Muehlbauer 1987; Estep 2005), can establish combining the use of marginal notching and PIT
occupancy and identify terrapin concentrations. tags.
Harden et al. (2009) counted terrapins on standard-
ized boat routes and compared the counts with
Noninvasive Methods
terrapin population sizes estimated through mark-­
recapture. Head counts w ­ ere highest at low tide Imaging combined with pattern recognition soft-
and ­under partly cloudy conditions. Visual surveys ware could identify individuals based on the dis-
estimated the number of terrapins and detected tinctive color patterns on terrapins’ shells and skin
inter-­annual trends over large areas at a fraction of (e.g., ­Gamble et al. 2008). One researcher (M. Farina,
the time and cost required for mark-­recapture stud- Ocean­side, NY, pers. comm.) identifies individual
ies. Detailed studies are needed to document how adult female terrapins by plastron scute seam pat-
head counts change seasonally and how weather terns, photocopying plastrons and matching them
and wave height affect detection probabilities. Pref- visually to the cata­log of previous captures. This
erably such studies would take place in an area system is limited to small populations, and scute
with ongoing mark-­recapture to explore the rela- seams may change through ontogeny; photocopy-
tionship between head-­count data and a­ ctual pop- ing of adults works well, but it needs to be validated
ulation size. for efficacy in juveniles.

Capture, Mea­sure­ment, and Field Techniques   13


commercial trade. Disadvantages include the high
Scute Notching
cost ($2 per tag) and increased probability of tag
Cagle (1939) developed the scute notching tech- loss if not properly injected.
nique used in many turtle demographic studies.
Using a triangular file, drill, or Dremel tool, one can
Binary Coded Wire Tags
notch or drill unique combinations of marginal
scutes, based on a coding system. In older juveniles Binary coded wire tags (BCWT; Northwest M ­ arine
and adults the marks are permanent, and hatchling Technologies, Inc.) are miniature tags (1.1 × 0.25 mm)
marks are recognizable as many as 8 to 10 years that can be injected into the terrapin. T ­ hese tags
­later; however, fast growth of hatchlings and juve- are inscribed with a unique number that can be
nile turtles can obscure the marks. Scute marking is read only before injection or ­after removal. There-
inexpensive and long lasting and can identify dead fore, detection is only presence/absence. We inject
individuals. Drilling and marking is invasive, how- tags into the muscle tissue of the rear legs, using a
ever, particularly if made deep enough to be long 25 gauge syringe. T ­ hese tags can mark large num-
lasting, and large population studies (>4,000 indi- bers of hatchlings in studies where cohort scute
viduals) require marking of at least 5 scutes. Fur- notches can assist in providing age data. The ad-
thermore, careful bookkeeping of used and unused vantage of BCWT over PIT tags to mark hatchlings
ID codes is necessary to prevent duplicates. Fi­nally, (Duncan 2013) is their lower cost ($0.33 each). How-
when marked animals incur carapace damage from ever, the equipment necessary to locate inserted
boat injuries or other sources, the identification of tags costs >$5,000. T­ hese tags w
­ ere found to easily
notch codes can become confusing. identify adults marked as hatchlings, and the co-
hort notch code confirmed age.
Passive Integrated Transponder Tags
External Tags
Passive integrated transponder (PIT) tags are excel-
lent for marking turtles (Buhlmann and Tuberville Two types of external tags can mark terrapins.
1998; Szerlag-­Egger and McRobert 2007). ­These tags Sequentially numbered Monel tags (National Tag
come in several sizes, and the smallest can be im- and Band Com­pany) are miniature forms of sea
planted into hatchlings (Rowe and Kelly 2005). turtle flipper tags. T ­ hese small tags require the
Such tags provide a 10-­to 15-­digit (depending on drilling of a 3 mm hole in one of the posterior
type) alphanumeric ID unique to the turtle. Loca- marginals. However, the accompanying pliers do
tions of PIT tag injection vary among terrapin not close the tags well, and we recommend using
­researchers. Some inject at the ventral base of a pair of needle-­nose pliers to apply the tag. Floy
the tail, in the muscle just above the plastron or spaghetti tags (Floy Tag) are applied similarly
(K. Buhlmann, University of Georgia, pers. comm.); but with a smaller hole that matches the dia­
we inject in the loose flap of skin cranial to the meter of the injector and holds the tag. T ­ hese also
rear leg along the plastron and have a <0.05% tag bear a number that allows the identification of
loss rate. We recommend Betadine to disinfect the individuals.
injection site and New-­Skin liquid ban­dage or cy- Both types of tags are temporary b ­ ecause im-
anacrolite glue to seal the wound. Tags identify proper attachment and growing/breaking out of
individuals from hatching to adulthood without the shell result in tag loss. However, they provide
the ambiguity caused by shell growth or damage, a good secondary mark when coupled with PIT
are minimally invasive, can often identify dead in- tags to facilitate clear identification of individuals.
dividuals, have no possibility of duplicate IDs, and They also are vis­i­ble, and thus tag return data can
enable recognition of individuals from other stud- be obtained from individuals not associated with
ies. An additional benefit is that t­hese tags can the study, e.g., commercial fishers or interested citi-
identify wild terrapins that have ended up in the zens who catch or find turtles.

14   W. M. Roosenburg and R. L. Burke


Toe Clipping Plastron and Carapace Length
Cagle (1939) suggested toe clipping for juvenile Plastron length (PL) and carapace length (CL) are
and hatchling turtles. Vogt (Instituto Nacional mea­sured as the straight-­line length of the plastron
de Pesquisas da Amazonia, pers. comm.) toe-­ along the midsagittal plane. Mea­sure­ments differ
clipped hatchling mud turtles (Kinosternon sp.) among researchers—­primarily ­whether to mea­
and South American river turtles (Podocnemis sp.) sure into the anterior and posterior notches
that w
­ ere ­later recognized when adults. No one (= midline PL/CL) or flush with the ends of the
has toe-­clipped terrapins, but it may be an effec- shell (= maximum PL/CL). Use of both techniques
tive way to mark hatchlings. Studies testing the ensures comparability among studies. Inserting the
effects of toe clipping on turtle locomotion or caliper tips into the notches is more consistent and
be­hav­ior are needed, and the technique is not provides less variability than maximum lengths.
recommended for adults. However, the removed Calipers should be deployed parallel to the shell;
toes could be used to age adults (Thomas et al. angling the calipers dramatically affects the mea­
1997). sure­ment. Use of flexible rulers and tape accounts
for the curvature of the shell but is more prone
to error created by variation of the doming of the
Paint
shell (shorter individuals with a higher shell may
Fingernail polish or paint pens can mark terrapins have a longer carapace than a longer individual
temporarily to allow identification from a distance, with a shorter shell). Doming can be accounted for
such as during nesting (Roosenburg and Dunham by using calipers to mea­sure shell height. Curved
1997). Nail polish lasts about 7 to 10 days, which is carapace mea­ sure­
ments can also be useful for
shorter than the inter-­nest interval (15 days; Roos- mea­sure­ments of shell volume.
enburg 1992). Paint markings or stick-on numbers
also can identify individuals basking or on the
Carapace Width and Shell Height
beach.
Carapace width (CW) is mea­sured at the widest
point of the carapace. Pinch the terrapin between
Mea­sur­ing Techniques
the beaks of the calipers to ensure they are perpen-
Turtle researchers have used a variety of mea­ dicular to the animal’s sagittal plane. Shell height
sure­ments to quantify the morphology of turtles. (SH) is mea­sured at the deepest part of the terrapin.
Most researchers use calipers; however, cloth or This is usually at the third vertebral scute on the
plastic tape mea­sures are low-­cost alternatives. carapace. The calipers should be perpendicular to
Digital scales and calipers greatly reduce the mea­ the animal’s longitudinal plane. Some prob­lems
sure­ment error associated with analog scales and may arise with individual terrapins that have knobs
verniers, but they do not correct for differences on vertebral scutes. Should one mea­sure the maxi-
among researchers’ mea­sur­ing of turtles. Although mum SH (including knobs) or SH without knobs?
digital scales and calipers are considerably more The individual study questions w ­ ill determine
expensive than analog tools, they greatly reduce which mea­sure­ment is more appropriate (e.g., mea­
mea­sure­ment errors in data sets. Flexible rulers sure maximum SH if a study is related to bycatch
or tape can be used to mea­sure terrapins, but the reduction devices [BRDs]).
results are not comparable to straight-­line mea­
sure­ments ­because of curvature of the shell. We
Head Width
recommend calipers, and below we describe mea­
sure­ments to standardize across population stud- Head width (HW) varies among terrapin popula-
ies so as to improve comparisons throughout the tions, but it is unknown ­whether ge­ne­tic differen-
terrapin’s range. tiation or plasticity driven by local diet c­ auses the

Capture, Mea­sure­ment, and Field Techniques   15


variation. Head width is mea­sured by g­ ently pinch- Permanent rec­ords of growth rings can be made
ing with calipers perpendicular to the sagittal using dental casts (Fig. 2.5) (Snider and Vano 1987).
plane at the surangular bone, which is the widest Examination of the casts ­under magnification facil-
point of the head. Tricks to encourage terrapins to itates precise observation of year-­to-­year changes in
extend their heads include grabbing the head when the number of growth rings and often reveals rings
extended and ­gently preventing retraction into the not vis­i­ble to the naked eye. Using pencil and paper
shell or holding the terrapin vertical, facing the to make an imprint of the annuli also can reveal
ground. The latter method generally results in head faint annuli and create a permanent rec­ord.
extension within a few seconds. Pushing in on When growth slows at the onset of maturity,
both forelimbs once the head is extended ­will pre- growth rings become compressed and difficult to
vent head retraction. count. Furthermore, the shedding of scutes elimi-
nates younger growth rings, and a­ fter 10 to 15 years
rings may no longer be used to precisely age a ter-
Mass
rapin. At this point, experienced researchers can
We recommend using portable digital top-­loading estimate an age range (e.g., <10, <15 years) from faint
balances or spring scales. A quick, vigorous shake rings. We recommend carefully examining the rings
can calm terrapins while the balance equilibrates. on several plastron scutes to determine age, particu-
larly when false annuli are suspected. Time of year
also can affect the number of annuli. Animals
Age
caught in spring or early summer prob­ably have not
Growth ring (annulus) counting (Sexton 1959) can yet grown that year, in which case the ring at the
age terrapins, but uncertain annularity requires midline seam should be included in the count. In
validation for e­ very population (Litzgus and Brooks contrast, the midline seam ring should not be in-
1998; Wilson et al. 2003). Arrested growth in win- cluded for terrapins caught in late summer; summer
ter forms distinct ridges or growth rings on the growth is lighter in color and sometimes softer than
shell. Although the ridges form on both the plas- the rest of the plastron.
tron and the carapace, the plastron is more reliable
for aging. Rings on the carapace, though frequently
Reproductive Status
deeper and larger, also are often less distinct and
blend together more in older individuals, result- Demographic studies require identification of
ing in “false annuli.” False annuli can sometimes which females are reproducing and quantification
be identified by comparing the preceding and fol- of age of first reproduction, clutch frequency, inter-
lowing rings; if the ­middle ring is less distinct, clutch interval, egg size, and egg and hatchling
then it may be false. We also suggest that age data survivorship—­data that are impor­tant for life his-
include both the number of annuli counted (age in tory analyses and population modeling. Inguinal
years) and the year of hatching. palpation (anterior to the hind leg) can detect the
The annularity of growth rings is confirmed presence or absence of shelled eggs, which feel like
when recaptures over several years document an small lumps or ping pong balls inside the terrapin.
increase in annuli consistent with the years passed. An experienced researcher may detect follicles, but
Recaptures over several years and recaptures of eggs are distinct and often dimple inside the fe-
marked hatchlings have confirmed that counting of male. Palpation can underestimate the number of
growth rings works well to estimate age in terra- gravid females in earlier egg stages, with less calcifi-
pins of Chesapeake Bay (Roosenburg, unpub. data) cation than is detected using radiographic methods
and South Carolina (Gibbons et al. 2001). However, (Keller 1998). X-­rays provide data on reproductive
we found very dif­fer­ent results for Jamaica Bay, status, number of eggs, and, if all individuals are
New York (Burke, unpub. data). X-­rayed similarly, egg size and width (Gibbons and

16   W. M. Roosenburg and R. L. Burke


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tristezas; sus placeres y regocijos
en suspiros y gemidos; su dulce
conversación en una soledad tan
triste que siempre anda huyendo
de aquellos que lo podrían hacer
compañía. En verdad te digo,
Grisaldo, que las veces que con
él me hallo, en verle cual le veo,
con gran lástima que le tengo, me
pesa de haberle encontrado,
viendo el poco remedio que á sus
males puedo darle.
Grisaldo.—Mal se puede
remediar el mal que no se
conoce; pero bien sería procurar
de saberlo dél, si como amigo
quisiesse manifestarnos lo que
siente.
Filonio.—Muchas veces se lo he
preguntado, y lo que entiendo es
que él no entiende su mal, ó si lo
conoce, no ha querido declararse
conmigo; pero lo que yo solo no
he podido, podría ser que
entrambos como amigos
podiésemos acabarlo. Y si su
dolencia es tal que por alguna
manera podiese ser curada, justo
será que á cualquiera trabajo nos
pongamos para que un zagal de
tanta estima y tan amigo y
compañero de todos no acabe tan
presto sus días, trayendo la vida
tan aborrida.
Grisaldo.—¿Pues sabes tú por
ventura dónde hallarlo
podiésemos? que assí goce yo de
mi amada Lidia, no procure con
menor cuidado su salud que la
mía propia.
Filonio.—No tiene estancia tan
cierta que no somos dudosos de
encontrarle, porque siempre se
aparta por los xarales más
espesos y algunas veces en los
valles sombríos, y en las cuevas
escuras se encierra, donde sus
gemidos, sus lamentaciones y
querellas no puedan ser oídas;
pero lo más cierto será hallarle á
la fuente del olivo, que está
enmedio de la espesura del
bosque de Diana, porque muchas
veces arrimado á aquel árbol lo
he visto tañer y cantar estando
puesto debaxo de la sombra y
oteando de allí su ganado, el cual
se puede decir que anda sin
dueño, según el descuido del que
lo apacienta.
Grisaldo.—Pues sigue, Filonio,
el camino, que cerca estamos del
lugar donde dices. Y para que
menos cansancio sintamos,
podremos ir cantando una
canción que pocos días ha
cantaba Lidia á la vuelta que
hacia del campo para la aldea
trayendo á sestear sus ovejas.
Filonio.—Comienza tú á decirla,
que yo te ayudaré lo mejor que
supiere.

GRISALDO
En el campo nacen flores
y en el alma los amores.
El alma siente el dolor
del zagal enamorado,
y en el alma está el amor
y el alma siente el cuidado;
assí como anda el ganado
en este campo de flores,
siente el alma los amores.

Filonio.—Calla, Grisaldo, no
cantemos: que á Torcato veo
adonde te dixe, y tendido en
aquella verde yerba, recostado
sobre el brazo derecho, la mano
puesta en su mexilla, mostrando
en el semblante la tristeza de que
continuamente anda
acompañado, y á lo que parece
hablando está entre sí. Por
ventura antes que nos vea
podremos oir alguna cosa por
donde podamos entender la
causa de su mal.
Grisaldo.—Muy bien dices; pues
no nos ha sentido, acerquémonos
más, porque mejor podamos oirle.
Torcato.—¡Oh, claro sol, que
con los resplandecientes rayos de
la imagen de tu memoria
alumbras los ojos de mi
entendimiento, para que en
ausencia te tenga presente,
contemplando la mucha razón
que tengo para lo poco que
padezco! ¿Por qué permites
eclipsar con la crueldad de tu
olvido la luz de que mi ánima
goza, poniéndola en medio de la
escuridad de las tinieblas
infernales, pues no tengo por
menores ni menos crueles mis
penas que las que en el infierno
se padecen? ¡Oh, ánima de
tantos tormentos rodeada! ¿cómo
con ser inmortal los recibes en ti
para que el cuerpo con el fuego
en que tú te abrasas se acabe de
convertir en ceniza? Si el uso de
alguna libertad en ti ha quedado,
sea para dexar recebir tanta parte
de tus fatigas al miserable cuerpo
que con ellas pueda acabar la
desventurada vida en que se vee.
¡Oh, desventurado Torcato, que tú
mesmo no sabes ni entiendes lo
que quieres, porque si con la
muerte das fin á los trabajos
corporales no confiesas que
quedarán en tu ánima inmortal
perpetuamente! Y si han de
quedar en ella, ¿no es mejor que
viviendo se los ayude á padecer
tu cuerpo en pago de la gloria que
con los favores pasados de tu
Belisia le fue en algún tiempo
comunicada? ¡Oh, cruel Belisia,
que ninguna cosa pido, ni desseo,
ni quiero, que no sea desatino,
sino es solamente quererte con
aquel verdadero amor y aficción
que tan mal galardonado me ha
sido! Ando huyendo de la vida por
contentarte y pienso que no te
hago servicio con procurar mi
muerte, porque mayor
contentamiento recibes con hacer
de mí sacrificio cada día y cada
hora que el que recebirías en
verme de una vez sacrificado del
todo, porque no te quedaría en
quién poder executar tu inhumana
crueldad, como agora en el tu sin
ventura Torcato lo haces; bien sé
que ninguna cosa ha de bastar á
moverte tu corazón duro para que
él de mí se compadezca; pero no
por esso te dexaré de manifestar
en mis versos parte de lo que
este siervo tuyo, Torcato, en el
alma y en el cuerpo padece.
Escuchadme, cruel Belisia, que
aunque de mí estés ausente, si
ante tus ojos me tienes presente,
como yo siempre te tengo, no
podrás dexar de oir mis dolorosas
voces, que enderezadas á ti
hendirán con mis sospiros el aire,
para que puedan venir á herir en
tus oídos sordos mis tristes
querellas.
Filonio.—Espantado me tienen
las palabras de Torcato, y no
puedo ser pequeño el mal que tan
sin sentir lo tiene que no nos haya
sentido; pero esperemos á ver si
con lo que dixere podremos
entender más particularmente su
dolencia, pues que de lo que ha
dicho se conoce ser los amores
de alguna zagala llamada Belisia.
Grisaldo.—Lo que yo entiendo
es que no he entendido nada,
porque van sus razones tan llenas
de philosofías que no dexan
entenderse; no sé yo cómo
Torcato las ha podido aprender
andando tras el ganado. Mas
escuchemos, porque habiendo
templado el rabel, comienza á
tañer y cantar con muy dulce
armonía.

TORCATO
¡Oh, triste vida de tristezas
llena,
vida sin esperanza de alegría,
vida que no tienes hora buena,
vida que morirás con tu porfía,
vida que no eres vida, sino
pena,
tal pena que sin ella moriría
quien sin penar algún tiempo
se viese,
si el bien que está en la pena
conociese!
Más aceda que el acebo al
gusto triste,
más amarga que el acíbar
desdeñosa,
ningún sabor jamás dulce me
diste
que no tornase en vida
trabajosa;
aquel bien que en un tiempo
me quesiste
se ha convertido en pena tan
rabiosa,
que de mí mismo huyo y de mí
he miedo
y de mí ando huyendo,
aunque no puedo.
Sabrosa la memoria que en
ausencia
te pone ante mis ojos tan
presente,
que cuando en mí conozco tu
presencia,
mi alma está en la gloria
estando ausente,
mas luego mis sentidos dan
sentencia
contra mi dulce agonía, que
consiente
tenerte puesta en mi
entendimiento
con gloria, pues tu gloria es
dar tormento.
¡Oh, quién no fuese el que
es, porque no siendo
no sentiría lo que el alma
siente!;
mi ánima está triste, y
padeciendo;
mi voluntad, ques tuya, lo
consiente;
si alguna vez de mí me estoy
doliendo
con gran dolor, es tal que se
arrepiente;
porque el dolor que causa tu
memoria
no se dexa sentir con tanta
gloria.
Mis voces lleva el viento, y
mis gemidos
rompen con mis clamores
l'aire tierno,
y en el alto cielo son más
presto oídos,
también en lo profundo del
infierno;
que tú quieres que se abran
tus oídos
á oir mi doloroso mal y eterno;
si llamo no respondes, y si
callo
ningún remedio á mis fatigas
hallo.
También llamo la muerte y
no responde,
que sorda está á mi llanto
doloroso;
si la quiero buscar, yo no sé á
dónde,
y ansí tengo el vivir siempre
forzoso;
si llamo á la alegría, se me
asconde;
respóndeme el trabajo sin
reposo,
y en todo cuanto busco algún
contento,
dolor, tristeza y llanto es lo que
siento.

TORNA Á HABLAR TORCATO


¡Oh, desventurado Torcato! ¿á
quién dices tus fatigas? ¿á quién
cuentas tus tormentos? ¿á quién
publicas tus lástimas y angustias?
Mira que estás solo; ninguno te
oye en esta soledad; ninguno
dará testimonio de tus lágrimas, si
no son las ninfas desta clara y
cristalina fuente y las hayas y
robles altos y las encinas, que no
sabrán entender lo que tú
entiendes. Das voces al viento,
llamas sin que haya quien te
responda, si no es sola Eco que,
resonando de las concavidades
destos montes, de ti se duele, sin
poder poner remedio á tu pasión.
¡Ay de mí, que no puedo acabar
de morir, porque con la muerte no
se acaban mis tormentos;
tampoco tengo fuerzas para
sustentar la miserable vida, la
cual no tiene más del nombre
sólo, porque verdaderamente está
tan muerta que yo no sé cómo me
viva! ¡Ay de mí, que muero y no
veo quién pueda valerme!
Grisaldo.—¡Filonio, Filonio; mira
que se ha desmayado Torcato!
Socorrámosle presto, que,
perdiendo la color, su gesto ha
quedado con aquel parecer que
tienen aquellos que llevan á meter
en la sepoltura.
Filonio.—¡Oh, mal afortunado
pastor, y qué desventura tan
grande! ¿Qué mal puede ser el
tuyo que en tal extremo te haya
puesto? Trae, Grisaldo, en tus
manos del agua de aquella
fuente, en tanto que yo sustento
su cabeza en mi regazo; ven
presto y dale con ella con toda
furia en el gesto, para que con la
fuerza de la frialdad y del miedo
los espiritus vitales que dél van
huyendo tornen á revivir y á
cobrar las fuerzas que perdidas
tenía; tórnale á dar otra vez con
ella.
Grisaldo.—¡Ya vuelve, ya vuelve
en su acuerdo! Acaba de abrir los
ojos, Torcato, y vuelve en ti, que
no estás tan solo como piensas.
Torcato.—El cuerpo puede tener
compañía; pero el alma, que no
está conmigo, no tiene otra sino la
de aquella fiera y desapiadada
Belisia, que contino della anda
huyendo.
Filonio.—Déxate deso, Torcato,
agora que ningún provecho traen
á tu salud esos pensamientos.
Torcato.—¿Y qué salud puedo
yo tener sin ellos, que no fuese
mayor emfermedad que la que
agora padezco? Pero decidme:
ansí Dios os dé aquella alegría
que á mí me falta, ¿que ventura
os ha traído por aquí á tal tiempo,
que no es poco alivio para mí ver
que en tan gran necesidad me
hayáis socorrido, para poder
mejor pasar el trabajo en que me
he visto; que bien sé que la
muerte, con todas estas
amenazas, no tiene tan gran
amistad conmigo que quiera tan
presto contarme entre los que ya
siguen su bandera?
Filonio.—La causa de nuestra
venida ha sido la lástima que de ti
y de tu dolencia tenemos; y el
cuidado nos puso en camino,
buscándote donde te hemos
hallado, para procurar como
amigo que vuelvas al ser primero
que tenías, porque según la
mudanza que en tus condiciones
has hecho, ya no eres aquel
Torcato que solías; mudo estás de
todo punto, y créeme, como á
verdadero amigo que soy tuyo,
que los males que no son
comunicados no hallan tan presto
el remedio necesario, porque el
que los padece, con la pasión
está ciego para ver ni hallar el
camino por donde pueda salir
dellos; así que, amigo Torcato,
páganos la amistad que tenemos
con decirnos la causa de tu dolor
más particularmente de lo cual
hemos entendido, pues ya no
puedes encubrir que no proceda
de amores y de pastora que se
llame Belisia, á la cual no
conocemos, por no haber tal
pastora ni zagala en nuestro
lugar, ni que de este nombre se
llame.
Grisaldo.—No dudes, Torcato,
en hacer lo que Filonio te ruega,
pues la affición con que te lo
pedimos y la voluntad con que,
siendo en nuestra mano, lo
remediaremos, merecen que no
nos niegues ninguna cosa de lo
que por ti pasa; que si conviene
tenerlo secreto, seguro podrás
estar que á ti mesmo lo dices,
porque los verdaderos amigos
una mesma cosa son para sentir
y estimar las cosas de sus
amigos, haciéndolas propias
suyas, así para saberlas encubrir
y callar como para remediarlas si
pueden.
Torcato.—Conocido he todo lo
que me habéis dicho, y aunque yo
estaba determinado de no
descubrir mi rabioso dolor á
persona del mundo, obligado
quedo con vuestras buenas obras
y razones á que como amigos
entendáis la causa que tengo
para la triste vida que padezco. Y
no porque piense que ha de
aprovecharme, si no fuere para el
descanso que recibiré cuando
viere que de mis tribulaciones y
fatigas os doléis, las cuales
moverán á cualquier corazón de
piedra dura á que de mí se duela
y compadezca. No quiero
encomendaros el secreto, pues
me lo habéis offrecido, que nunca
por mí vaya poco en que todo el
mundo lo sepa. Es tanto el amor
que tengo á esta pastora Belisia,
que no querría que ninguno
viniesse á saber el desamor y
ingratitud que conmigo ha usado,
para ponerme en el extremo que
me tiene.
Filonio.—Bien puedes decir,
Torcato, todo lo que quisieres,
debaxo del seguro que Grisaldo
por ambos te ha dado.
Torcato.—Ora, pues, estad
atentos, que yo quiero comenzar
desde el principio de mis amores
y gozar del alivio que reciben los
que cuentan sus trabajos á las
personas que saben que se han
de doler dellos.

COMIENZA TORCATO Á
CONTAR EL PROCESO DE
SUS AMORES CON LA
PASTORA BELISIA
En aquel apacible y sereno
tiempo, cuando los campos y
prados en medio del frescor de su
verdura están adornados con la
hermosura de las flores y rosas
de diversas colores, que la
naturaleza con perfectos y lindos
matices produce, brotando los
árboles y plantas las hojas y
sabrosas frutas, que con gran
alegría regocijan los corazones de
los que gozarlas después de
maduras esperan, estaba yo el
año passado con no menor
regocijo de ver el fruto que mis
ovejas y cabras habían brotado,
gozando de ver los mansos
corderos mamando la sabrosa
leche de las tetas de sus madres
y á los ligeros cabritos dando
saltos y retozando los unos con
los otros; los becerros y terneros
apacentándose con la verde y
abundante yerba que en todas
partes les sobraba, de manera
que todo lo que miraba me
causaba alegría, con todo lo que
veía me regocijaba, todo lo que
sentía me daba contento,
cantando y tañendo con mi rabel
y chirumbela passaba la más
sabrosa y alegre vida que contar
ni deciros puedo.
Muchas veces, cuando tañer me
sentían los zagales y pastores
que en los lugares cercanos sus
ganados apacentaban,
dexándolos con sola la guarda de
los mastines, se venían á bailar y
danzar con grandes desafíos y
apuestas, poniéndome á mí por
juez de todo lo que entre ellos
passaba; y después que á sus
majadas se volvían, gozaba yo
solo de quedar tendido sobre la
verde yerba, donde vencido del
sabroso sueño sin ningún cuidado
dormía, y cuando despierto me
hallaba, contemplando en la luz y
resplandor que la luna de sí daba,
en la claridad de los planetas y
estrellas, y en la hermosura de los
cielos y en otras cosas
semejantes passaba el tiempo, y
levantándome daba vuelta á la
redonda de mi ganado y más
cuando los perros ladraban, con
temor de los lobos, porque ningún
daño les hiciessen.
Y después de esto, pensando
entre mí, me reía de los
requiebros y de las palabras
amorosas que los pastores
enamorados á las pastoras
decían, gozando yo de aquella
libertad con que á todos los
escuchaba, y con esta sabrosa y
dulce vida, en que con tan gran
contentamiento vivía, pasé hasta
que la fuerza grande del sol y la
sequedad del verano fueron
causa que las yerbas de esta
tierra llana se marchitassen y
pusiesen al ganado en necesidad
de subirse á las altas sierras,
como en todos los años
acostumbraban hacerlo; y ansí,
juntos los pastores, llevando un
mayoral entre nosotros, que en la
sierra nos gobernase, nos fuimos
á ella. Y como de muchas partes
otros pastores y pastoras también
allí sus ganados apacentassen,
mi ventura, ó por mejor decir
desventura, traxo entre las otras á
esa inhumana y cruel pastora,
llamada Belisia, cuyas gracias y
hermosura así aplacieron á mis
ojos, que con atención la miraban,
que teniéndolos puestos en ella
tan firmes y tan constantes en su
obstinado mirar, como si cerrar, ni
abrir, ni mudar no los pudiera,
dieron lugar con su descuidado
embovescimiento que por ellos
entrase tan delicada y
sabrosamente la dulce ponzoña
de Amor, que cuando comencé á
sentirla ya mi corazón estaba tan
lleno della que, buscando mi
libertad, la vi tan lexos de mí ir
huyendo, con tan presurosa ligera
velocidad, que por mucha
diligencia que puse en alcanzarla,
sintiendo el daño que esperaba
por mi descuido, jamás pude
hacerlo, antes quedé del todo sin
esperanza de cobrarla, porque
volviendo á mirar á quien tan sin
sentido robádomela había, vi que
sus hermosos ojos, mirándome,
contra mí se mostraban algo
airados, y parecióme casi conocer
en ellos, por las señales que mi
mismo deseo interpretaba,
decirme: ¿De qué te dueles,
Torcato? ¿Por ventura has
empleado tan mal tus
pensamientos que no estén mejor
que merecen? Yo con grande
humildad, entre mí respondiendo,
le dije: Perdonadme, dulce ánima
mía, que yo conozco ser verdad
lo que dices, y en pago de ello
protesto servirte todos los días
que viviere con aquel verdadero
amor y affición que á tan gentil y
graciosa zagala se debe.
Y ansí, dándole á entender, con
mirarla todas las veces que podίa,
lo que era vedado á mi lengua,
por no poder manifestar en
presencia de los que entre
nosotros estaban el fuego que en
mis entrañas comenzaba á
engendrarse, para convertirlas
poco á poco en ceniza,
encontrándonos con la vista
(porque ella, casi conociendo lo
que yo sentίa, también me
miraba), le daba á conocer que,
dexando de ser mía, más
verdaderamente estaba cautivo
de su beldad y bien parecer. Y
mudando el semblante, que
siempre solίa estar acompañado
de alegrίa, en una dulce tristeza,
también comencé á trocar mi
condición, de manera que todos
conocían la novedad que en mí
había.
Y todo mi deseo y cuidado no era
otro sino poder hablar á la mi
Belisia, y que mi lengua le
pudiese manifestar lo que sentía
el corazón, para dar con esto
algún alivio á mi tormento; y
porque mejor se pudiese encubrir
mi pensamiento, determiné en lo
público mostrar otros amores, con
los cuales fengidos encubriese los
verdaderos, para que de ninguno
fuesen sentidos, y así me mostré
aficionado y con voluntad de
servir á una pastora llamada
Aurelia, que muchas veces
andaba en compañía de la mi
Belisia, y conversaba con mucha
familiaridad y grande amistad con
ella. Y andando buscando tiempo
y oportunidad para que mi deseo
se cumpliese, hallaba tantos
embarazos de por medio, que no
era pequeña la fatiga que mi
ánima con ellos sentía. Y
habiéndose juntado un día de
fiesta algunos pastores y pastoras
en la majada de sus padres de la
mi Belisia, después de haber
algún rato bailado al son que yo
con mi chirumbela les hacía, me
rogaron que cantase algunos
versos de los que solía decir otras
veces, y sin esperar á que más
me lo dixesen, puestos los ojos
con la mejor disimulación que
pude á donde la afición los
guiaba, dando primero un
pequeño sospiro, al cual la
vergüenza de los que presentes
estaban detuvo en mi pecho, para
que del todo salir no pudiese,
comencé á decir:

Extremos que con fuerza así


extremada
dais pena á mis sentidos tan
sin tiento,
teniendo al alma triste,
fatigada,
Causáisme de continuo un
tal tormento
que mi alma lo quiere y lo
asegura,
porque viene mezclado con
contento.
Si acaso vez alguna se
figura
á mi pena cruel que se fenece,
ella misma el penar siempre
procura.
Cuando el cuidado triste en
mí más crece,
mayor contento siento y mayor
gloria,
porque el mismo cuidado la
merece.
De mal y bien tan llena mi
memoria
está, que la razón no
determina
cuál dellos lleva el triunfo de
vitoria.
Con este extremo tal que
desatina,
mi esperanza y mi vida van
buscando
el medio[1274] que tras él
siempre camina.
Y si grandes peligros van
pasando,
ninguno les empece ni fatiga;
de todos ellos salen
escapando.
El agua no les daña, porque
amiga
á mis lágrimas tristes se ha
mostrado,
pues que ellas dan camino en
que las siga.
El fuego no las quema, que
abrasado
de otro fuego mayor siempre
me siente,
y assí passan por él muy sin
cuidado.
También mi sospirar nunca
consiente
que el viento les fatigue ni dé
pena,
si aquel de mis sospiros no
está ausente.
Amor con mi ventura así lo
ordena,
para mostrar en mí su gran
potencia,
porque á perpetua pena me
condena.
Dada está contra mí cruel
sentencia,
que no pueda morir, ni yo
matarme

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