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Red Snapper Biology in a
Changing World
CRC Marine Biology Series
The late Peter L. Lutz, Founding Editor, David H. Evans and
Stephen Bortone, Series Editors

Artificial Reefs in Fisheries Management


edited by Stephen A. Bortone, Frederico Pereira Brandini, Gianna Fabi, and
Shinya Otake
Biology of Sharks and Their Relatives, Second Edition
edited by Jeffrey C. Carrier, John A. Musick, and Michael R. Heithaus
The Biology of Sea Turtles, Volume III
edited by Jeanette Wyneken, Kenneth J. Lohmann, and John A. Musick
The Physiology of Fishes, Fourth Edition
edited by David H. Evans, James B. Claiborne, and Suzanne Currie
Interrelationships Between Coral Reefs and Fisheries
edited by Stephen A. Bortone
Impacts of Oil Spill Disasters on Marine Habitats and Fisheries in North
America
edited by J. Brian Alford, PhD, Mark S. Peterson, and Christopher C. Green
Hagfish Biology
edited by Susan L. Edwards and Gregory G. Goss
Marine Mammal Physiology
Requisites for Ocean Living
edited by Michael A. Castellini and Jo-Ann Mellish
Shark Research
Emerging Technologies and Applications for the Field and Laboratory
edited by Jeffrey C. Carrier, Michael R. Heithaus, and Colin A. Simpfendorfer
Red Snapper Biology in a Changing World
edited by Stephen T. Szedlmayer and Stephen A. Bortone

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Red Snapper Biology in a
Changing World

Edited by
Stephen T. Szedlmayer and Stephen A. Bortone
CRC Press
Taylor & Francis Group
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Boca Raton, FL 33487-2742

© 2020 by Taylor & Francis Group, LLC


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Contents
Preface.......................................................................................................................ix
Acknowledgments......................................................................................................xi
Editors..................................................................................................................... xiii
Contributors.............................................................................................................. xv

Chapter 1
Introduction................................................................................................................. 1
Stephen A. Bortone and Stephen T. Szedlmayer

Early Life History

Chapter 2
Red Snapper, Lutjanus campechanus, Larval Dispersal in the Gulf of Mexico........ 7
Donald R. Johnson and Harriet M. Perry

Chapter 3
Juvenile Red Snapper, Lutjanus campechanus, Densities on Small Artificial
Reefs to Estimate Year-Class Strength..................................................................... 27
Peter A. Mudrak and Stephen T. Szedlmayer

Site Fidelity, Movements, and Mortality

Chapter 4
A Review of Red Snapper, Lutjanus campechanus, Acoustic Telemetry Studies.......... 49
Laura Jay Williams-Grove and Stephen T. Szedlmayer

Chapter 5
Fishing Mortality Estimates for Red Snapper, Lutjanus campechanus, Based
on Acoustic Telemetry and Conventional Mark-Recapture...................................... 75
Peter A. Mudrak and Stephen T. Szedlmayer

v
vi Contents

Environmental Effects

Chapter 6
Polycyclic Aromatic Hydrocarbons in Red Snapper, Lutjanus campechanus,
and Sediment Samples after the Deepwater Horizon Oil Spill................................97
Claire E. Roberts and Stephen T. Szedlmayer

Chapter 7
The Effects of Modeled Dispersed and Undispersed Hypothetical Oil Spills
on Red Snapper, Lutjanus campechanus, Stocks in the Gulf of Mexico............... 123
Benny J. Gallaway, Wolfgang J. Konkel, and John G. Cole

Diets and Age Determination

Chapter 8
Diet Analyses of Red Snapper, Lutjanus campechanus, Based on DNA
Barcoding from Artificial Reefs in the Northern Gulf of Mexico......................... 143
Stephen T. Szedlmayer and Rachel A. Brewton

Chapter 9
Timing of Opaque Band Formation and Validation of Annular Increments in
Otoliths of Red Snapper, Lutjanus campechanus.................................................. 167
Stephen T. Szedlmayer, Erin J. Fedewa, and Morgan L. Paris

Chapter 10
A Comparison of Age Determination between Whole and Sectioned Otoliths
in Red Snapper, Lutjanus campechanus................................................................ 185
Stephen T. Szedlmayer and Danielle J. Grundy

Management

Chapter 11
Improving the Historical Baseline of the Gulf of Mexico Red Snapper,
Lutjanus campechanus, Fishery............................................................................. 201
Gary R. Fitzhugh, William C. Kline, Clay E. Porch, and
Christopher L. Gardner
Contents vii

Chapter 12
Management Strategies Influencing Recreational Red Snapper, Lutjanus
campechanus, Effort in the Gulf of Mexico: Why Can’t We Agree?..................... 233
Carrie M. Simmons, R. Ryan Rindone, and Michael F. Larkin

Chapter 13
A Comparison of Two Fishery-Independent Surveys of Red Snapper,
Lutjanus campechanus, from 1999–2004 and 2011–2015..................................... 249
Stephen T. Szedlmayer, Peter A. Mudrak, and Jessica Jaxion-Harm

Chapter 14
Artificial Reefs in the Future Management of Red Snapper, Lutjanus
campechanus.......................................................................................................... 275
Stephen A. Bortone

Index....................................................................................................................... 285
Preface
Red Snapper, Lutjanus campechanus, is an important commercial and recre-
ational fish species throughout its range in warm temperate waters in near coastal
and continental shelf waters along the coastal western Atlantic Ocean. As part
of an early fishery in the United States, it continues to command attention for
commercial, recreational, ecological, and biological reasons for many communi-
ties in the New World. Due to its importance, there has been much interest in
maintaining its status among a variety of scientific, social, and economic levels.
Concomitantly, since its early development as a fishery in the late 1800s, times
have changed. These changes are many and include increased commercial fishing
pressure due to increasing demand from a growing number of human consum-
ers. There has also been a tremendous increase in recreational fishing pressure,
principally the result of increasing access to the fishery through sport angling.
Moreover, technology has developed, thus increasing fishing efficiency and per-
haps exacerbating the effect of increased fishing participation. Simultaneously,
environmental changes resulting from an ever-increasing human population and
coastal development have led to environmental stress (habitat loss and degrada-
tion), potential climate change, and altered ecosystems, with additional pressures
on the fish and its fishery.
Appropriately, colleagues at federal, state, and local institutions (both public and
private) have shown concern as to how Red Snapper have been responding to the pres-
sures indicated above. Likewise, aquatic resource scientists have conducted studies
on Red Snapper throughout its long history as a fishery. Awareness has grown of the
stresses and pressures the fishery faces, coupled with the acknowledgment that while
efforts are underway to alleviate the effect of these pressures and stressors, there is a
need to be aware of the current status and potential future trends of the fundamental
biological features of the Red Snapper.
To help all concerned with an understanding of these biological aspects, a sym-
posium was held in September 2017 at the 147th Annual Meeting of the American
Fisheries Society in Tampa, Florida, entitled “Red Snapper Biology in a Changing
World.” This volume is a compendium of many papers presented at the symposium.
It includes additional research completed as a result of the symposium, which sought
to explore the aforementioned changing conditions surrounding the fishery and their
potential effects on Red Snapper, chiefly in the Gulf of Mexico and along the south-
eastern United States. The symposium explored the responses Red Snapper have
made, and are making, relative to their life history attributes, such as early life his-
tory and adult ecology, especially attributes associated with population distribution
and abundance, movement patterns, diets, age and growth, and fish health issues.
These are examined here in relation to ever-changing environmental responses Red
Snapper make to long-term changes in habitat, potential climate change, short-term
environmental stressors (e.g., oil spills and hypoxia), and fishery effects (changing
regulations). The overall goal of the symposium was to provide a better understanding
of Red Snapper biology in relation to population fluctuations. This information will

ix
x Preface

subsequently allow better-informed user groups and ultimately more circumspect


management decisions in an effort to maintain sustainable Red Snapper fisheries.

Stephen T. Szedlmayer
Stephen A. Bortone
Acknowledgments
We thank numerous colleagues who lent their expertise to conduct professional
peer reviews of the manuscripts. We also thank John Sulzycki and especially Alice
Oven and her staff at CRC Press (Taylor and Francis Group) for their help in seeing
this volume through to completion. Lastly, we thank the organizers of the American
Fisheries Society (Jeff Hill, University of Florida) and the participants who afforded
us the opportunity to bring together the scientific expertise essential to meeting the
objective of bringing focus to the future of Red Snapper as a sustainable resource.
We thank Diane Rome Peeples for permission to use her illustration of a Red Snapper
for the cover of this volume.

xi
Editors
Stephen T. Szedlmayer, PhD, at an early age became interested in marine biology
and marine science along the wetlands of southeastern Pennsylvania and the near-
shore coasts of southern New Jersey. Collecting various species of grass shrimp,
bivalve clams, polychaete worms, sticklebacks, killifish, and toadfish at a young age
instilled a life-long interest and endeavor to study marine biology and associated
fishes. He attended Millersville University, Lancaster, Pennsylvania, for undergradu-
ate studies. Due to Millersville’s newly established marine biology program and his
guidance from Professor Samuel J. Ha, he further advanced his interests and studies
of marine fish ecology. He continued his studies at the University of South Florida,
earning a Master’s degree in the Marine Science program under the supervision of
Professor John C. Briggs. After graduating from the University of South Florida,
he worked for a short period at Mote Marine Laboratory, Sarasota, Florida before
entering the graduate program at the Virginia Institute of Marine Science, College
of William and Mary, Gloucester Point, Virginia. At the Virginia Institute, he was
greatly influenced by the guidance of Professor John A. Musick and by an exciting
group of fellow graduate students. After completing his PhD, he accepted a post-
doctoral position at the Rutgers University Marine Field Station, Tuckerton, New
Jersey and collaborated with Professor Kenneth W. Able. He was subsequently hired
by the School of Fisheries, Aquaculture and Aquatic Sciences, Auburn University,
Alabama, where he rose through the ranks to professor and mentored numerous
graduate students on studies of various aspects of marine fish ecology. He continued
his studies of marine fish ecology and especially focused on habitats and life histo-
ries in Red Snapper in the northern Gulf of Mexico.

Recently retired as executive director of the Gulf of Mexico Fishery Management


Council, Stephen A. Bortone now is an environmental consultant with Osprey
Aquatic Sciences, LLC, based in Windham, New Hampshire (United States). As
a consultant, he specializes in fisheries and is noted especially as an authority on
artificial reefs. Currently, he is also Marine Biology series editor with CRC Press.
Previously, Dr. Bortone served as director of the Minnesota Sea Grant College
Program with an appointment as professor of biology at the University of Minnesota
Duluth. Earlier in his career, he was the founding director of the Marine Laboratory
at the Sanibel-Captiva Conservation Foundation in Sanibel, Florida, director
of environmental science at the Conservancy of Southwest Florida, and direc-
tor of the Institute for Coastal and Estuarine Research while professor of biology
at the University of West Florida. Dr. Bortone received the BS degree (Biology)
from Albright College in Reading, Pennsylvania (1968); the MS degree (Biological
Sciences) from Florida State University, Tallahassee (1970); and the PhD (Marine
Science) from the University of North Carolina, Chapel Hill (1973).
He has authored more than 180 scientific articles on the broadest aspects of the
aquatic sciences over the past 50 years. In addition, he has edited six books on the
aquatic sciences: Sea Grasses; Biology of the Spotted Sea Trout; Estuarine Indicators;

xiii
xiv Editors

Artificial Reefs in Fisheries Management; and Interrelationships between Corals


and Fisheries (all with CRC Press) in addition to Marine Artificial Reef Research
and Development: Integrating Fisheries Management Objectives with the American
Fisheries Society.
Internationally, he was a visiting scientist at The Johannes Gutenberg University
(Mainz, Germany) and conducted underwater fish surveys with colleagues from La
Laguna University in the Canary Islands for several years. He was also a Mary Ball
Washington scholar while conducting research and teaching at University College
Dublin, Ireland. Notable awards and honors include “Fellow” from the American
Institute of Fishery Research Biologists, “Certified Fisheries Professional” from the
American Fisheries Society, and “Certified Senior Ecologist” from the Ecological
Society of America.
Contributors
Stephen A. Bortone Jessica Jaxion-Harm
Osprey Aquatic Sciences, LLC School of Fisheries, Aquaculture and
Windham, Hampshire Aquatic Sciences
Auburn University
Rachel A. Brewton Fairhope, Alabama
Harbor Branch Oceanographic
Institute Donald R. Johnson
Florida Atlantic University Center for Fisheries Research and
Fort Pierce, Florida Development, Gulf Coast Research
Laboratory
John G. Cole University of Southern Mississippi
LGL Ecological Research Associates Ocean Springs, Mississippi
Bryan, Texas
William C. Kline
Erin J. Fedewa Riverside Technology
NOAA/AFSC Shellfish Assessment (Contracting for the National Marine
Program Fisheries Service)
Kodiak Fisheries Research Center Panama City Laboratory
Kodiak, Alaska Panama City, Florida

Gary R. Fitzhugh Wolfgang J. Konkel


National Marine Fisheries Service ExxonMobil Biomedical Sciences
Panama City Laboratory Annandale, New Jersey
Panama City, Florida
Michael F. Larkin
Benny J. Gallaway National Marine Fisheries Service
LGL Ecological Research Associates Southeast Regional Office
Bryan, Texas St. Petersburg, Florida

Christopher L. Gardner Peter A. Mudrak


Riverside Technology School of Fisheries, Aquaculture and
(Contracting for the National Marine Aquatic Sciences
Fisheries Service) Auburn University
Panama City Laboratory Fairhope, Alabama
Panama City, Florida
Morgan L. Paris
Danielle J. Grundy School of Fisheries, Aquaculture and
Washington Department of Fish and Aquatic Sciences
Wildlife Auburn University
Twisp, Washington Fairhope, Alabama

xv
xvi Contributors

Harriet M. Perry Carrie M. Simmons


Center for Fisheries Research and Gulf of Mexico Fishery Management
Development, Gulf Coast Research Council
Laboratory Tampa, Florida
University of Southern Mississippi
Ocean Springs, Mississippi Stephen T. Szedlmayer
School of Fisheries, Aquaculture and
Clay E. Porch Aquatic Sciences
National Marine Fisheries Service Auburn University
Southeast Fisheries Science Center Fairhope, Alabama
Sustainable Fisheries Division
Miami, Florida Laura Jay Williams-Grove
Southeast Fisheries Science Center
R. Ryan Rindone NOAA, National Marine Fisheries
Gulf of Mexico Fishery Management Service
Council U.S. Department of Commerce
Tampa, Florida Miami, Florida

Claire E. Roberts
School of Fisheries, Aquaculture and
Aquatic Sciences
Auburn University
Fairhope, Alabama
Chapter 1

Introduction

Stephen A. Bortone and Stephen T. Szedlmayer

The introductions to each of the chapters included herein, in total, provide the
necessary background information to fully understand the past and current situation
regarding the status of Red Snapper as an ecological component of the hard-bottom
biotope in the warm temperate waters off the southeastern United States and in the
Gulf of Mexico. Additionally, the chapters provide sufficient information to obtain
a fluent grasp of the current status of the Red Snapper fishery. Each chapter also
provides a summary of its findings as well as a sense of future directions of research
for each chapter’s respective subject. Consequently, this introduction will not reiter-
ate those features of Red Snapper and its associated fisheries (both commercial and
recreational) here.
As indicated in the Preface to this volume, the compilation of chapters offered is
largely the result of papers presented during a symposium held at the 147th annual
meeting of the American Fisheries Society in Tampa, Florida (United States) entitled
“Red Snapper Biology in a Changing World.” The chief purpose of this sympo-
sium was to bring together our current state of knowledge of the biology of the Red
Snapper, Lutjanus campechanus, as the species has responded to, and is likely to
respond to, the pressures of intense commercial and recreational fishing (and their
concomitant associated regulations) as well as potential changes wrought upon the
fishery by seemingly exacerbating environmental pressures associated with possible
climate change.
The chapters offered are grouped into sub-disciplines of the species’ life history.
The initial grouping, “Early Life History,” includes a chapter by Johnson and Perry
on the dispersal of Red Snapper larvae in the Gulf of Mexico. Here, the authors indi-
cate that western Gulf of Mexico populations (west of the Mississippi River Delta) do
not substantially interact with eastern Gulf populations, largely due to strong natal
retention of each population to its respective area along the northern Gulf of Mexico.
Consequently, these populations are potentially only weakly connected. This sce-
nario suggests a future with greater biological differentiation of Red Snapper stocks
in the northern Gulf of Mexico. Hence, fisheries managers should be mindful of this

1
2 Red Snapper Biology in a Changing World

life history feature when considering the application of Red Snapper stock param-
eters across the northern Gulf of Mexico.
The second chapter, by Mudrak and Szedlmayer, under the general heading of
“Early Life History” examines the utility of determining the relative abundance of
Red Snapper on artificial reefs. This study demonstrates the advantage that future
researchers may have in estimating population sizes of Red Snapper (especially in the
juvenile life stage) on relatively new, but controllable, habitats such as artificial reefs.
A second grouping of papers offered is organized under the heading of “Site
Fidelity, Movements, and Mortality” and is significant in that much of the future
management effort for Red Snapper will involve having a clear understanding of the
effect that changes in the environment may have on these life history features. The
chapter by Williams-Grove and Szedlmayer reviews our present understanding of
acoustic telemetry studies. This overview not only highlights the recent advances in
understanding the movements of Red Snapper but also indicates an ever-improving
future with regard to an even better understanding of this life history feature with the
advent of modern advances.
An additional chapter in this grouping, by Mudrak and Szedlmayer, points to a
future where sophisticated measures of fishing mortality may be obtainable through
acoustic telemetry in concert with traditional mark-recapture technology. The effi-
ciency gained through increasing accuracy with reduced effort points to a future
where management decisions can be made much more responsibly.
Two chapters are grouped under a heading of “Environmental Effects.” The
first of these, by Roberts and Szedlmayer, examines the presence of polycyclic aro-
matic hydrocarbons (PAHs) in Red Snapper, expected to be present when an oil
spill occurs. Based on past experience, no one would doubt the inevitability of a
future with more oil spills in near coastal areas off the northern Gulf of Mexico.
Interestingly, the data presented here indicate little evidence of presence of PAHs
in the tissues of Red Snapper that were likely to have been exposed to oil as part of
the Deepwater Horizon oil spill, which occurred in northern Gulf of Mexico waters.
This information portends a brighter future for Red Snapper stocks, at least based on
the early findings of tissue loading this study offers.
Additionally in the subject of “Environmental Effects” (especially as they relate
to oil spills), Gallaway et al. offer a chapter that models the effects of the dispersal of
a hypothetical oil spill on Red Snapper in the northern Gulf of Mexico. These study
results are especially important, as they incorporate effects of Red Snapper exposure
during the spawning season. This modeling effort indicates little significant effects
on the Red Snapper population, even with the addition of dispersants as an attempt
to reduce the oil spill effects. Using dispersants during oil spill events could also pro-
vide minor additional environmental benefits to surface-associated organisms (e.g.,
seabirds, sea turtles, marine mammals, etc.) as well as reducing threats to sensitive
shorelines.
“Diets and Age Determination” as a category includes three chapters, each
providing an improvement to our understanding of a future with well-understood
Red Snapper stocks. Szedlmayer and Brewton investigate employing sophisticated
DNA-barcoding techniques to help determine diets and food habits of Red Snapper
Introduction 3

associated with artificial reefs. Previous to this study, there was significant difficulty
in identifying stomach contents to the species level. Here, this obstacle is overcome,
and the study results indicate that food items consumed by Red Snapper occurred
on both near-reef and on-reef biotopes. These results re-emphasize the importance
of artificial reefs in providing a trophic base for artificial reef–associated fish preda-
tors and give managers additional insight into the role artificial reefs may play in the
future management of Red Snapper.
Szedlmayer et al. validate the timing of annulus formation in the otoliths of Red
Snapper from a mark-recapture study. Previous studies presented varied results in
the interpretation of annulus (age) formation on hard body parts of fishes, and espe-
cially in Red Snapper. The accuracy of these highly valued recapture data is signifi-
cant in determining the age and growth of Red Snapper in a warm temperate habitat.
Consequently, these study results will be valuable to future studies that examine Red
Snapper responses to changing growing conditions.
In addition to validating annular age-ring formation in Red Snapper, there is
a need to reduce costs and improve efficiency in the age determination process.
Herein, Szedlmayer and Grundy compare the repeatability of age determination
using high-effort/high-cost methods such as otolith sectioning with the low-effort/
low-cost gross inspection of otolith rings on whole, unsectioned otoliths. Their dem-
onstration of comparable age determination will lead to a more data-rich future with
regard to studies involving age determination in Red Snapper.
“Management” as a group heading for chapters presented here is the final, but
most important, feature of a volume that seeks to provide a direction for a sustainable
future with Red Snapper. Fitzhugh et al. significantly add to our understanding of
the past status and trends in the Red Snapper fishery. They accomplish this through
an examination of historical literature that was previously difficult to find, and in
some cases, unobtainable. Through their inspection of historical journals and popu-
lar records that were temporally topical, they are able to glean a different and better
understanding of the Red Snapper fishery in a “pre-digital” world. Their considerate
contribution not only pertains to the historical Red Snapper fishery but gives guid-
ance on how other fisheries should be investigated for historical records in the future.
Simmons et al. pose (and answer) a reasonable, but important, question with
regard to the Red Snapper fishery in the northern Gulf of Mexico. The historical
contentiousness of the fishery is examined to give a perspective as to the present
situation of Red Snapper management. While the management structure is not yet
finalized, their perspective gives us hope for a better future as we understand the
mistakes of the past.
In an additional chapter on “Management,” Szedlmayer et al. compare two
fishery-independent survey methods for estimating Red Snapper stocks. Evidence
offered by the application of these two methods (i.e., catch per unit effort [CPUE]
and absolute density based on side-scan sonar and hydroacoustic survey component)
provides future researchers with comparable stock assessment methodology that
will allow better future stock estimates and greater overall accuracy of our present
fish stock assessment techniques on fish species associated with structure and hard-
bottom biotypes.
4 Red Snapper Biology in a Changing World

Bortone considers the future management of Red Snapper with a more directed
effort, which will probably include artificial reefs. Because of insufficient and var-
ied research results to date, few fisheries (including Red Snapper) formally consider
artificial reefs as part of their respective management plans. The lack of predictable
results from artificial reef deployment efforts is chief among the reasons for the lack
of inclusion of artificial reefs in these plans. For a variety of reasons, Red Snapper
may be the ideal species to incorporate artificial reefs in a management portfolio
of possible options. Barriers to the inclusion of artificial reefs still exist, but future
efforts that include the results of directed research on life history limitations (i.e.,
bottlenecks) may provide the keys to the inclusion of artificial reefs in a sustainable
Red Snapper future.
In total, this volume represents a snapshot of the Red Snapper fishery and biology
at a time when many economic, environmental, and social pressures are accumulat-
ing. It is our expectation that this subject of the future management of Red Snapper
will be revisited, but with an altered direction, principally due to the perspectives
offered here.
Early Life History
Chapter 2

Red Snapper, Lutjanus campechanus,


Larval Dispersal in the Gulf of Mexico

Donald R. Johnson and Harriet M. Perry

CONTENTS

Abstract.......................................................................................................................7
Introduction.................................................................................................................8
Methods..................................................................................................................... 14
Shelf Dispersal..................................................................................................... 14
Dispersal into the Deep Basin.............................................................................. 14
Cross-Basin Connections..................................................................................... 15
Results and Discussion............................................................................................. 15
Shelf Dispersal..................................................................................................... 15
Dispersal into the Deep Basin.............................................................................. 16
Cross-Basin Connections..................................................................................... 17
Conclusions.......................................................................................................... 19
Acknowledgments..................................................................................................... 21
References................................................................................................................. 21

ABSTRACT

Red Snapper, Lutjanus campechanus, is a demersal, nonmigratory, continen-


tal-shelf species with strong site fidelity yet an extensive latitudinal habitat range.
Directly coincident with this range is the western boundary current system in the
warm temperate North Atlantic Ocean, suggesting that Red Snapper range may be
linked by larval dispersal. Information regarding the balances between natal reten-
tion of pelagic larvae and broad dispersal along the boundary current is fundamental
to understanding variation in population abundance by natural causes. The Gulf of
Mexico has a large population of Red Snapper and a long history of oceanographic
observations and oceanic current modeling. In the present study, this history of

7
8 Red Snapper Biology in a Changing World

current modeling was reviewed to examine possible connections of Red Snapper


populations through larval drift within the distributional limits of Red Snapper.
Ocean current dynamics, driven by climatological processes, may affect larval dis-
persal and regional differences that occur in the Gulf of Mexico. For Red Snapper,
connections were relatively weak across the Gulf of Mexico basin but sufficient to
allow genetic mixing. In the southern Gulf of Mexico, the relatively large produc-
tion of Red Snapper larvae on Campeche Bank was predominantly retained on the
Bank, with a small portion of these larvae transported by the Loop Current both to
the west Florida shelf and into the Florida Current. In the northern Gulf of Mexico,
topographic barriers and degree of penetration of the Loop Current with spin-off
eddies were important factors for the dispersal of larvae. Topographic features
reduced connections between the eastern and western Gulf of Mexico. Eastward
larval transport during high spawning months was directed from settlement areas
toward deep water. Westward transport facilitates larval dispersal, but this occurred
during months when spawning was reduced. Energetic Loop Current spin-off eddies
negatively affected highly fecund Red Snapper populations on the outer shelf west
of the DeSoto Canyon. Dispersal to the deep basin in the Gulf of Mexico was high
and variable, while larval redistribution to suitable settlement habitat was weak.
Ocean current modeling indicated that the more abundant Red Snapper population
in the western Gulf of Mexico did not contribute substantially to stocks east of the
Mississippi Delta. Also, there were areas of strong natal retention despite wide larval
dispersal, but substantial loss to the deep basin over much of the Gulf of Mexico. The
overall picture in the Gulf of Mexico was one of weakly connected populations with
scales of roughly 100–200 km set by hydrography.

INTRODUCTION

Red Snapper, Lutjanus campechanus, reside near structured continental shelf


habitat at depths between 10 and 190 m for most of their life span (Szedlmayer and
Shipp 1994; Schroepfer and Szedlmayer 2006; Gallaway et al. 2009; Topping and
Szedlmayer 2011; Piraino and Szedlmayer 2014; Williams-Grove and Szedlmayer
2016). Juveniles first recruit to low-relief, inner-shelf habitats, and as they age and
grow, they disperse to increasingly larger structures over the continental shelf
with no discernible migratory patterns (Szedlmayer and Conti 1999; Rooker et al.
2004; Szedlmayer and Lee 2004; Gallaway et al. 2009; Szedlmayer 2011; Syc and
Szedlmayer 2012). Red Snapper, while demersal and generally nonmigratory, has a
native range that extends from the equator off northeast Brazil, through the Gulf of
Mexico and along the eastern seaboard of the continental United States, to past Cape
Hatteras along the Mid-Atlantic Bight (Anderson et al. 2015). Records of Red Snapper
from the Caribbean and South America have been attributed to the Caribbean Red
Snapper, Lutjanus purpureus (Anderson et al. 2015). However, genetic studies indi-
cated that the Caribbean Red Snapper was the same species as Red Snapper (Gomes
et al. 2008, 2012). In addition, a lack of phylogeographic structuring in Red Snapper
(Cervigón et al. 1993), morphological similarities and shared biological traits (Rivas
Larval Dispersal in the Gulf of Mexico 9

1966; Wilson and Nieland 2001), and geographic distributional data (Anderson et al.
2015) all support the synonymy of the two species.
The large latitudinal range of Red Snapper and its western boundary location
coincide with the warm temperate western North Atlantic current system (Figure 2.1).
Concurrence of the Red Snapper range with this large-scale current suggests a fun-
damental linkage that occurs naturally through planktonic larval dispersal (Rindone
et al. 2015). The persistent western boundary current system in the North Atlantic
includes the North Brazil Current, the Guiana Current, the Caribbean Current, the
Yucatan Current, the Loop Current, the Florida Current, and the Gulf Stream (http://
oceancurrents.rsmas.miami.edu). Although these currents flow over deep water
along the continental shelf edge, interactions with shelf water and shelf-spawned
ichthyoplankton occur through highly variable spatial and temporal turbulent eddy

Figure 2.1 Western boundary system of surface currents in the North Atlantic, coincident
with the habitat range of Red Snapper. Blue vectors: speed 0.25 to 0.50 m/s. Red
vectors: speed >0.50 m/s. Currents are gridded averages from 1979 to 2017, com-
puted from satellite-tracked mixed layer drifters from the Global Drifter Program
(NOAA/Atlantic Oceanographic and Meteorological Laboratory [AOML]).
10 Red Snapper Biology in a Changing World

exchanges (Hanisko and Lyczkowski-Shultz 2003; Johnson et al. 2017). The dichot-
omy between an extensive geographic range and a relatively resident lifestyle sug-
gested that larval drift may be associated with a persistent transport pathway for
broad dispersal as an important component of Red Snapper life history strategy.
Red Snapper has a planktonic larval duration of ~30 days (Szedlmayer and Conti
1999; Rooker et al. 2004). This pelagic life-stage is sufficiently protracted to allow pas-
sive transport and entrainment into the offshore currents for broad dispersal, but is suf-
ficiently brief to allow some degree of retention to natal areas. Both broad dispersal and
natal retention are also ensured by a relatively long and intensive spawning period (May–
October) that can be affected by seasonal changes in ocean flow patterns (Collins et al.
2001). In addition, the interaction of strong offshore currents with waters of the conti-
nental shelf varies greatly along the current pathway, as do regional differences in dis-
persal mechanisms (Hare et al. 2002). Changes in the western boundary current system
(Toggweller and Key 2001) and local ocean transport conditions associated with climate
change (Taylor et al. 2012) may influence future distribution of Red Snapper, although it
is unclear how the specific balance of natal retention and broad dispersal will be affected.
Continental shelves (5 to 200 m deep) account for ~31% of the Gulf of Mexico area,
with Campeche Bank off the north coast of the Yucatan Peninsula, Mexico, account-
ing for ~25% of the total Gulf of Mexico shelf area (Figure 2.2). The Campeche Bank
is bounded on the east by the Yucatan Channel, where the intense Yucatan Current
and northward-intruding Loop Current create a transport linkage between larvae on
the Bank and dispersal to other parts of the Gulf of Mexico and the southeast U.S.
Atlantic coast. A large population of Red Snapper resides over the Campeche Bank, and
spawning occurs from February to November, with a peak in the early fall (Brulé et al.
2010). Although harvest of Red Snapper on Campeche Bank has decreased from earlier
high values (Brulé et al. 2010), its location adjacent to the Loop Current in the Gulf of
Mexico makes it a potential cross-basin source to other areas in the Gulf of Mexico.
Major drivers for currents on the shelves are river and estuary outflow, wind stress,
and energetic exchanges with the deep basin winds (Sturges et al. 2005). Circulation
in nearshore waters tends to flow in a counter-clockwise direction around the Gulf
basin (Ohlmann and Niiler 2005). Freshwater outflow from surrounding land sur-
faces produces a buoyancy layer that is constrained along the coastline in width and
direction by Coriolis effects (Gill 1982). In offshore waters, wind stress (direction
of forcing) has a major influence on circulation. For much of the year, wind stress
is westward and equatorward, producing a pattern that is stronger in the southern
Gulf and weaker in the north (Figure 2.3). This westward flow results in a negative
wind stress curl (Zavala-Hidalgo et al. 2014), causing clockwise circulation along the
outer shelf and slope. This clockwise flow was recorded in the earliest current charts
derived from ship logs and originally reported as the principal circulation in the Gulf
of Mexico (Maury 1855; Sturges and Blaha 1976). During mid-summer, when Red
Snapper spawning is at its peak, wind stress responds to continental heating with a
light shoreward component in the northern Gulf of Mexico (Figure 2.3). Ekman drift
from wind stress along coastlines counters the anti-clockwise nearshore buoyancy-
driven flows, complicating larval directional tendencies (Whitney and Garvine 2005)
and dispersing them into nearshore waters over the shelf (Johnson et al. 2001).
Larval Dispersal in the Gulf of Mexico 11

Figure 2.2 Gulf of Mexico with 200 and 1000 m isobaths. The 200 m isobath is defined as the
shelf edge, and the area between the 200 and 1000 m isobaths is defined as the
upper continental slope. TX/LA is the Texas–Louisiana continental shelf. MS/AL
is the Mississippi–Alabama continental shelf. Major topographic barriers to along-
shelf flow are the Mississippi River Delta, DeSoto Canyon, and the Apalachicola
Peninsula. The Flower Gardens National Marine Sanctuary and Florida Middle
Grounds Habitat of Particular Concern are noted by black diamonds.

The general tendency for surface currents to flow counter-clockwise near-


shore and clockwise offshore in the northern Gulf of Mexico becomes more
complex when factors such as wind stress variation, turbulent exchanges at the
shelf break, and continental shelf topography are considered. Across the north-
ern Gulf of Mexico, topographic barriers such as the Mississippi River Delta,
the DeSoto Canyon, and Apalachicola Peninsula interrupt the basic current pat-
terns, breaking the shelf into circulation regions (Ohlmann and Niiler 2005;
Johnson et al. 2009; Cardona et al. 2016). The Mississippi River Delta extends
close to the continental shelf edge, separating the Louisiana–Texas shelf from
the Mississippi Bight (Apalachicola, Florida to the Mississippi River; Perry et al.
2003). The Mississippi Bight is separated from the peninsular west Florida shelf
by the Apalachicola Peninsula and is itself separated into a western and an east-
ern region by the DeSoto Canyon. These regional separations create challenges
for management of the Red Snapper stock, since both adults and larvae are more
prevalent in the western Gulf of Mexico (Lyczkowski-Shultz and Hanisko 2007;
Johnson et al. 2009; Karnauskas et al. 2017).
12 Red Snapper Biology in a Changing World

Figure 2.3 Gulf of Mexico surface wind vectors for (A) January and (B) July. Vector direction
is direction of force (stress) on the ocean (oceanographic convention). Wind data
from NOAA/ (NCEI) monthly blended climatology.

A natural means of bypassing hydrographic separation on the shelf can be


through exchange of water with the deep basin and return to the shelf elsewhere
(Hare et al. 2002). However, this deep water exchange is difficult, as the dominant
flow of currents over the upper continental slope is along isobaths due to the law
of conservation of angular momentum (vorticity), and buoyancy differences in the
upper layer between the shelf and basin tend to be maintained. This results in density
fronts at the shelf edge that inhibit cross-isobath exchange. Transport of planktonic
larvae between the shelf and deep basin requires a driving mechanism in the form
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condensed from the sea, and fresh water always brings a big price.
There are no streams anywhere for miles around. The town is
situated in the crater of an extinct volcano, and there is one great
depression near by in which some famous stone tanks were made a
thousand or so years ago. These tanks are so big that if they were
cleaned out they might hold thirty million gallons of water. The water
is caught when it rains, and is sometimes auctioned off to the highest
bidder. The receipts go to the British Government, to which a good
rain may bring in fifteen or twenty thousand dollars or more.
This is my second visit to Aden. My first was sixteen years ago
when I stopped here on my way around the world. I do not see that
the town has changed and I doubt whether it has any more people
than it had then. The population is made up of all the nations and
tribes common to the Indian Ocean. It contains Arabs, Africans,
Jews, Portuguese, and East Indians. There are about four thousand
Europeans, including merchants, officials, and soldiers. The majority
of the people are Arabs and the prevailing colour is black. There are
tall, lean, skinny black Bedouins from interior Arabia, who believe in
the Prophet, and go through their prayers five times a day. There are
black Mohammedans from Somaliland and black Christians from
Abyssinia. In addition there are Parsees, Hindus, and East Indian
Mohammedans of various shades of yellow and brown. A few of the
Africans are woolly-headed, but more of them have wavy hair. The
hair of the women hangs down in corkscrew curls on both sides of
their faces. Of these people neither sex wears much clothing. The
men have rags around the waist, while the women’s sole garments
are skirts which reach to the feet.
The East Indians, who are everywhere, do most of the retail
business and trading. They are found peddling on every street
corner. They dress according to their caste and religion. The
Parsees, who are fire-worshippers, wear black preacher-like coats
and tall hats of the style of an inverted coal scuttle. The East Indian
Mohammedans wear turbans and the Hindus wrap themselves up in
great sheets of white cotton. There are besides many Greeks and
Italians, and not a few Persians. The English dress in white and wear
big helmets to keep off the sun.
This is the land of the camel. Caravans are coming in and going
out of the city every day bringing in bags of Mocha coffee and gums
and taking out European goods and other supplies to the various
oases. There is a considerable trade with Yemen as well as with the
tribes of southeastern Arabia. There are always camels lying in the
market places, and one sees them blubbering and crying as they are
loaded and unloaded. They are the most discontented beasts upon
earth, and are as mean as they look. One bit at me this afternoon as
I passed it, and I am told that they never become reconciled to their
masters. Nevertheless, they are the freight animals of this part of the
world, and the desert could not get along without them. They furnish
the greater part of the milk for the various Arab settlements, and the
people make their tents of camel’s hair. They are, in fact, the cows of
the desert. They are of many different breeds, varying as much in
character as horses. There are some breeds that correspond to the
Percheron, and the best among them can carry half a ton at a load.
There are others fitted solely for riding and passenger travel. The
ordinary freight camel makes only about three miles an hour and
eighteen miles is a good day’s work. The best racing camels will
travel twenty hours at a stretch, and will cover one hundred miles in
a day. Seventy-five miles in ten hours is not an uncommon journey
for an Arabian racer, and much better speed has been made. As to
prices, an ordinary freight camel brings about thirty dollars, but a
good riding camel costs one hundred dollars and upward.
Have you ever heard how the camel was created? Here is the
story of its origin as told by the Arabs. They say that God first formed
the horse by taking up a handful of the swift south wind and blowing
upon it. The horse, however, was not satisfied with his making. He
complained to God that his neck was too short for easy grazing and
that his hoofs were so hard that they sank in the sand. Moreover, he
said there was no hump on his back to steady the saddle.
Thereupon, to satisfy the horse, God created the camel, making him
according to the equine’s suggestions. And when the horse saw his
ideal in flesh and blood he was frightened at its ugliness and
galloped away. Since then there is no horse that is not scared when
it first sees a camel.
This story makes me think of the Arab tradition as to how God first
made the water buffalo, which, as you know, is about the ugliest
beast that ever wore horns, hair, and skin. God’s first creation was
the beautiful cow. When He had finished it the devil happened that
way, and as he saw it he laughed at the job, and sneered out that he
could make a better beast with his eyes shut. Thereupon the Lord
gave him some material such as He had put into the cow and told
him to go to work. The devil wrought all day and all night, and the
result was the water buffalo.
I have made inquiries here and elsewhere as to the Arabian horse.
He is a comparatively scarce animal and he does not run wild in the
desert, as some people suppose. Indeed, comparatively few of the
Arabian tribes have horses, and the best are kept on the plateau of
Najd, in the centre of the peninsula. They belong to the Anazah tribe,
which is one of the oldest of all, and which claims to date back to the
Flood. It is a wealthy tribe, and it has been breeding horses for many
generations. The best stock has pedigrees going back to the time of
Mohammed, and the very choicest come from five mares which were
owned by the Prophet and blessed by him. These horses seldom go
out of Arabia. They are owned by the chiefs, and are not sold, except
in times of the direst necessity. Now and then a few get into Egypt
and other parts of North Africa, and the Sultan of Turkey has usually
had some for his stables.
It is only occasionally that a pure-bred Arabian goes to Europe or
the United States. Two of the best stallions we ever imported were
those which General Grant brought from Constantinople. This was, I
think, during his tour around the world. While in Turkey he and the
Sultan visited the royal stables together. As they looked over the
horses the Sultan told Grant to pick out the one he liked best, and he
designated a dapple gray called the Leopard. “It is yours,” said the
Sultan, “and this also,” pointing to a four-year-old colt called Linden
Tree. In due time these two horses arrived in the United States and
were put on General Ed Beale’s farm near Washington. They were
used for breeding, and they produced about fifty fine colts.
CHAPTER XXIX
IN MOMBASA

Mombasa is the terminus of the Uganda Railway as it comes down


from Lake Victoria. It is the port of entry for all the sea-borne trade of
the seven provinces of British East Africa, or Kenya Colony, as it is
now called, Uganda, and adjacent territory. It is on an island halfway
down the coast of East Africa and just below the Equator, where old
Mother Earth is widest and thickest. If I should stick a peg down
under the chair in which I am writing into the old lady’s waist, and
then travel westward in a straight line I would soon reach the upper
end of Lake Tanganyika, and a little later come out on the Atlantic
Ocean just above the mouth of the Congo. Crossing that great sea, I
should make my next landing in South America, at the mouth of the
Amazon, and, going up the Amazon valley, I should pass Quito, in
Ecuador, and then drop down to the Pacific. From there on the trip to
the peg stuck in at Mombasa would comprise sixteen or more
thousand miles of water travel. I should cross the Pacific and Indian
oceans, and the only solid ground on the way would be the islands of
New Guinea, Borneo, and Sumatra.
Three thousand miles from Port Said and more than six thousand
miles from London, Mombasa is far below the latitude of the
Philippines. It is just about a day by ship north of Zanzibar and thirty
days’ sailing from New York.
So far, most of my travels in Africa have been in the sands, with
only a patch of green now and then. I was close to the Sahara in
Morocco, and I travelled many hundreds of miles over it while in
Algeria and Tunisia. In Tripoli my eyes were made sore by the glare
of the Libyan wastes and their dust blew across the Nile valley
during my stay in Egypt and the British Sudan. The Arabian desert
was on both sides of us as we came down the Red Sea and its
sands several times covered the ship. We had the rockiest of all
deserts in southern Arabia while that of Italian Somaliland was not
any better.
Here at Mombasa we are in the luxuriant tropics where the
surroundings remind me of Solomon’s song. All nature seems joyful.
The rain has conquered the sun and there are mosses, vines, and
trees everywhere, The shores of the mainland are bordered with
coconuts, we have mighty baobab trees loaded with green scattered
over the island, and even its cliffs are moss grown.
A jungle of green on a foundation of coral, Mombasa is only a mile
or so wide and three miles in length, but it rises well up out of the
sea and is so close to the continent that one can almost hear the
wind blow through the coconut groves over the way. On the island
itself the jungle has been cut up into wide roads. There is a lively
town with a polyglot population at one end of it, and the hills are
spotted with the homes of the British officials. The island has two
good harbours, a little one and a big one. The little one, which is in
the main part of the town, is frequented by small craft. The big one
could hold all the ships that sail the East Coast, and the people here
say it is to be the great port of this side of the continent. The larger
harbour is called Kilindini, a word that means “the deep place,” It has
only a few warehouse sheds and a pier above it, the main
settlements being across the island four miles away.
It was in Kilindini that I landed, and that under difficulties. Our ship
was anchored far out and our baggage was taken ashore in native
boats. Finding the main quay was crowded, I had my boatman go
direct to the custom house and let us out on the beach. The custom
house is a little shed about big enough for one cow situated so high
up above the water that our trunks had to be carried out upon the
heads of the Negroes. The water came up to their middles, but
nevertheless they waded through it and brought both us and our
baggage to the land. The customs examination was lenient. The
officers looked through our trunks for guns and ammunition and
warned us that we could not hunt elephants and hippopotami without
a two-hundred-and-fifty-dollar license. A little later the natives again
took our trunks and lugged them about a quarter of a mile to the top
of a hill, where we got the cars for Mombasa.
The word “cars” savours of electricity or steam. The cars I took
were run by men. Here in East Africa human muscle forms the
cheapest power. The wages of the natives run from five cents a day
upward, while in the interior there are many who will work eight or
nine hours for three cents. The result is that the trolley cars are
propelled by men. Each car consists of a platform about as big as a
kitchen table, with wheels underneath and an awning overhead. In
the middle of the platform there is a bench accommodating two to
four persons. The wheels run on a track about two feet in width, and
each car is pushed from behind by one or more bare-legged and
bare-headed men who run as they shove it up hill and down. There
are such car tracks all over the island, with switches to the homes of
the various officials. There are private cars as well as public ones,
and everyone who is anybody has his own private car with his
coolies to push him to and from work. At the beginning and closing of
his office hours, which here are from eight until twelve and from two
until four, the tracks are filled with these little cars, each having one
or more officials riding in state to or from the government buildings.
Kilindini harbour, or “the deep place”, is connected with the town of Mombasa by
a mile-long tramway, the cars of which are pushed by native runners. Mombasa is
the chief port of Kenya Colony.
In this African village there are 25,000 natives, representing perhaps a hundred
tribes, each with its own dress and customs. All, however, are eager buyers of the
gaudy print cloths in the bazaars of the Hindu merchants.

In Kenya Colony the East Indians complicate matters for the British government.
They practically control the retail trade and, having grown rich and prosperous,
have begun to raise embarrassing political issues.
I wish I could show you this old town of Mombasa. It began before
Columbus discovered America, and the citizens can show you the
very spot where Vasco da Gama landed when he came here from
India shortly after he discovered the new route to Asia by the Cape
of Good Hope. He landed here in 1498 at just about the time that
Columbus was making his third voyage to America. Even then
Mombasa was a city and Da Gama describes it. A little later it
became the property of the Portuguese. The most prominent building
in the town is the great red Fort of Jesus, built by the Portuguese in
1593, when the city was made the capital of their East African
possessions. It was later the scene of massacres and bloody fights
between Portuguese and Arabs. To-day the red flag of the Sultan of
Zanzibar flies over the old fort, now used as a prison, admission to
which is forbidden.
After the Portuguese were driven out the Arabs held Mombasa for
many years, and it was an Arab ruler, the Sultan of Zanzibar, who
owned it when the British came in. It still belongs to him in a nominal
way. He has leased it to the British for so much a year, but his flag
floats above John Bull’s ensign everywhere on the island.
Most of the population of Mombasa is African. Of the twenty-six
thousand inhabitants, only about three hundred and fifty are white.
There are people here from all parts of the interior, some of them as
black as jet, and a scattering few who are chocolate brown or yellow.
These natives live in huts off by themselves in a large village
adjoining the European and Asiatic quarters. Their houses are of
mud plastered upon a framework of poles and thatched with straw.
The poles are put together without nails. There is not a piece of
metal in any of them, except on the roof, where here and there a
hole has been patched up with a rusty Standard Oil can. Very few of
the huts are more than eight feet high, while some are so low that
one has to stoop to enter them. They are so small that the beds are
usually left outside the house during the daytime, and the majority of
each family sleep on the floor.
I find this African village the most interesting part of Mombasa. Its
inhabitants number over twenty-five thousand and comprise natives
of perhaps one hundred tribes, each of which has its own dress and
its own customs. Most of the women are bare-headed and bare-
legged; and some of the men are clad in little more than breech
cloths. Now and then one sees a girl bare to the waist, and the little
ones wear only jewellery. On the mainland all go more or less naked.
It is amazing how these people mutilate themselves so as to be
what they consider beautiful. The ears of many of the women are
punched like sieves, in order that they may hold rings of various
kinds. At one place I saw a girl with a ring of corks, each about as
big around as my little finger, put through holes in the rims of her
ears. She had a great cork in each lobe and three above that in each
ear. There was a man beside her who had two long sticks in his
ears; and in another place I saw one who had so stretched the lobe
holes that a good-sized tumbler could have been passed through
them. Indeed, I have a photograph of a man carrying a jam pot in his
ear.
The most numerous of the natives here in Mombasa are the
Swahilis. These are of a mixed breed found all along the central
coast of East Africa. They are said to have some Arab blood and for
this reason, perhaps, are brighter and more businesslike than the
ordinary native. The Swahilis are found everywhere. They have little
settlements in the interior in the midst of other tribes, and the Swahili
language will carry one through the greater part of Central and East
Africa. The British officials are required to learn it, and one can buy
Swahili dictionaries and phrase books. During most of my journey I
shall take a Swahili guide with me, or rather a black Swahili boy, who
will act as a servant as well as guide.
Let me give you a picture of the Swahili women as I see them
here. Their skins are of a rich chocolate brown and shine as though
oiled. They have woolly hair, but they comb it in a most extraordinary
way, using a razor to shave out partings between the rows of plaited
locks so that when the hair is properly dressed the woman seems to
have on a hood of black wool. I took a snapshot of two girls who
were undergoing the process of hairdressing yesterday, fearing the
while that their calico gowns, which were fastened by a single twist
under the armpits, might slip. A little farther on Jack took a
photograph of another giddy maiden clad in two strips of bright-
coloured calico and numerous earrings, while I gave her a few
coppers to pose for the picture. At the same time on the opposite
side of the street stood a black girl gorgeous with jewellery. In her
nose she had a brass ring as big around as the bottom of a dinner
bucket, and her ears had holes in their lobes so big that a hen’s egg
could be put through them without trouble. Not only the lobes, but
the rims also were punctured, each ear having around the edges five
little holes of about the size of my little finger. These holes were filled
with rolls of bright-coloured paper cut off so smoothly that they
seemed almost a part of the ear. The paper was of red, green, and
blue and looked very quaint.
The coast Negroes of East Africa are often Swahilis, descendants of Arab
traders and their native wives. They have a dialect of their own and pride
themselves on being more intelligent than the pure-bred Africans.
The Uganda Railroad plunges the traveller into the blackest of the Black
Continent, where the natives seem people of another world. The few clothes they
wear are a recent acquisition from the white men.
CHAPTER XXX
THE UGANDA RAILWAY

Travelling by railway through the wilds of East Africa! Steaming for


hundreds of miles among zebras, gnus, ostriches, and giraffes!
Rolling along through jungles which are the haunts of the rhinoceros
and where the lion and the leopard wait for their prey! These were
some of my experiences during my trip over the Uganda Railway
from Mombasa to Nairobi.
Only a few years ago it took a month to cover the distance
between these two points. To-day I made it in less than twenty-four
hours, and that in a comfortable car. The railroad fare, travelling first-
class, was fifty-eight rupees, which at normal exchange would total
about thirty dollars, and I had good meals on the way. The distance
is over three hundred miles, just about half the length of the railroad.
Wood-burning locomotives of the American type are largely used.
The maximum scheduled speed is twenty-five miles an hour. Trains
leave Mombasa daily for Nairobi and three times a week for Kisumu
on Lake Victoria, which is five hundred and eighty-four miles from
Mombasa.
Leaving Mombasa, our train carried us across a great steel bridge
to the mainland, and we climbed through a jungle up to the plateau.
We passed baobab trees, with trunks like hogsheads, bursting out at
the top into branches. They made me think of the frog who tried to
blow himself to the size of a bull and exploded in the attempt. We
went through coconut groves, by mango trees loaded with fruit, and
across plantations of bananas, whose long green leaves quivered in
the breeze made by the train as it passed. Now we saw a
gingerbread palm, and now strange flowers and plants, the names of
which we did not know. As we went upward we could see the strait
that separates Mombasa from the mainland, and higher still caught a
view of the broad expanse of the Indian Ocean. For the first one
hundred miles the climb is almost steady, and we were about one
third of a mile above the sea when we reached the station at Voi.
Here the country is more open, and far off in the distance one can
see a patch of snow floating like a cloud. That patch is the mountain
of Kilimanjaro, the top of which is more than nineteen thousand feet
above the sea. It is the loftiest mountain on the continent yet is not
much higher than Mt. Kenya, that other giant of British East Africa,
which rises out of the plateau some distance north of Nairobi.
After the jungle of the coast line, the country becomes
comparatively open and soon begins to look like parts of America
where the woods have been cut away and the brush allowed to grow
up in the fields. Here the land is carpeted with grass about a foot or
so high. Thousands of square miles of such grass are going to
waste. I saw no stock to speak of, and at that place but little wild
game. Without knowing anything about the tsetse fly and other cattle
pests, I should say that the pastures just back of the coast might
feed many thousands of cattle and hogs. The soil seems rich. It is a
fat clay, the colour of well-burnt brick, which turns everything red.
The dust filled our car; it coated our faces, and crept through our
clothes. When we attempted to wash, the water soon became a
bright vermilion, and the towels upon which we dried were brick-red.
My pillow, after I had travelled all night through such dust, had
changed from white to terracotta, and there was a Venetian red spot
where my head had lain. The wisest travellers cleansed eyes and
nostrils several times a day with an antiseptic solution.
It is a strange thing to go to sleep in the woods and then awake to
find one’s self rolling over a high, treeless country with game by the
thousand gambolling along the car tracks. We awoke on the Kapiti
plains, which are about a mile above the sea and two hundred and
sixty-eight miles from Mombasa. These plains are of a black sandy
loam and covered with a thick grass. They look much as Iowa,
Kansas, or Nebraska did when the railroads were first built through
them and the buffaloes galloped along with the cars. The same
conditions prevail here save that the game is of a half-dozen big
kinds, and most of it is such as one can see only in our zoölogical
gardens at home. According to law no shooting may be done for a
mile on each side of the track, so that the road has become a great
game preserve two miles in width and about six hundred miles long.
The animals seem to know that they are safe when they are near the
railroad, for most of them are as quiet as our domestic beasts when
in the fields.
Let me give you some notes which I made with these wild animals
on all sides of me. I copy: “These Kapiti plains are flat and I am
riding through vast herds of antelopes and zebras. Some of them are
within pistol shot of the cars. There are fifty-odd zebras feeding on
the grass not one hundred feet away. Their black and white stripes
shine in the sunlight. Their bodies are round, plump, and beautiful.
They raise their heads as the train goes by and then continue their
grazing. Farther on we see antelopes, some as big as two-year-old
calves and others the size of goats. The little ones have horns
almost as long as their bodies. There is one variety which has a
white patch on its rump. This antelope looks as though it had a
baby’s bib tied to its stubby tail or had been splashed with a
whitewash brush. Many of the antelopes are yellow or fawn
coloured, and some of the smaller ones are beautifully striped.
“Among the most curious animals to be seen are the gnus, which
are sometimes called wilde-beeste. As I write this there are some
galloping along with the train. They are great beasts as big as a
moose, with the horns of a cow and the mane and tail of a horse.
Hunting them is good sport.
“But look, there are some ostriches! The flock contains a dozen or
more birds, which stand like interrogation points away off there on
the plain. They turn toward the cars as we approach, then spread
their wings and skim away at great speed. Giraffes are frequently
seen. They are more timid than the antelope, and by no means so
brave as the zebras.”
All the steel in the bridges on the Uganda Railway was made in the United
States and put in place under American direction, because the British bidders
wanted three times as long and double the price for the job.

Built primarily to break up the slave-trade in East Africa, the Uganda Railroad
has also proved that the natives, under proper direction, can become useful
workers. Thousands of them have been employed in the construction and
maintenance of the line.

The natives rob the railroad of quantities of wire, which to them is like jewellery.
Both men and women load themselves down with pounds of it coiled around their
arms, legs, and necks, and even through their ears.
The Uganda Railway begins at the Indian Ocean and climbs over
some of the roughest parts of the African continent before it ends at
Lake Victoria, one of the two greatest fresh-water lakes of the world.
Leaving the seacoast, the rise is almost continuous until it reaches
the high plains of Kenya Colony. Here at Nairobi, where this chapter
is written, I am more than a mile above the sea, and, about fifteen
miles farther on at the station of Kikuyu, the road reaches an altitude
seven hundred feet above the top of Mt. Washington. From there the
ascent is steady to a point a mile and a half above the sea. Then
there is a great drop into a wide, ditch-like valley two thousand feet
deep. Crossing this valley, the railway again rises until it is far higher
than any mountain in the United States east of the Rockies. It attains
an elevation of eighty-three hundred feet, and then falls down to
Lake Victoria, which is just about as high as the highest of the
Alleghanies. The line was built by the British Government in less
than five years and has cost altogether some thirty-five million
dollars. It has a gauge of forty inches, rails which weigh fifty pounds
to the yard, and tracks which are well laid and well ballasted. In an
average year almost two hundred and fifty thousand tons of goods
and five hundred thousand passengers are carried over it, and its
earnings are more than its operating expenses.
It does not yet pay any interest on the capital invested, but it is of
enormous value in the way of opening up, developing, and protecting
the country. It was not constructed as a commercial project but to
combat the slave trade which flourished beyond the reach of the
British warships. To-day the Uganda line is the dominant influence of
Kenya Colony.
Among the most interesting features are the American bridges,
which cross all the great ravines between Nairobi and Lake Victoria.
Every bit of steel and every bolt and rivet in them was made by
American workmen in American factories, and taken out here and
put up under the superintendence of Americans. This was because
of John Bull’s desire to have the work done quickly and cheaply and
at the same time substantially. While he had been laying the tracks
from here to the sea our bridge companies had surprised the English
by putting up the steel viaduct across the Atbara River in the Sudan
within a much shorter time and far more cheaply than the best British
builders could possibly do. Therefore, when the British Government
asked for bids for these Uganda bridges, they sent the plans and
specifications to the English and to some of our American firms as
well. The best British bids provided that the shops should have two
or three years to make the steel work, and longer still to erect it in
Africa. The American Bridge Company offered to complete the whole
job within seven months after the foundations were laid, and that at a
charge of ninety dollars per ton, to be paid when all was in place and
in working order. This price was about half that of the British
estimates and the time was less than one third that in which the eight
bridges already constructed had been built, so the American
company got the contract. It carried it out to the letter, and had the
government done its part, the work would have been completed in
the time specified. Owing to delays of one kind and another, it really
consumed five months longer, but it was all done within the space of
one year, which was just about half the time that the British
contractors asked to get their goods ready for shipment.
The English were surprised at how easily and quickly the
Americans carried out their contract and how little they seemed to
make of it. A. B. Lueder, the civil engineer who was sent out to take
charge of the construction, was little more than a boy and had
graduated at Cornell University only a year or so before. There were
about twenty bridge builders and foremen from different parts of the
United States, and a Pennsylvania man named Jarrett who acted as
superintendent of construction. Arriving at Mombasa in December,
1900, these men had completed their work before the following
Christmas. They acted merely as superintendents and fancy
workmen. All the rough labour was done by East Indians and native
Africans, furnished by the British. When the road was started, the
government planned to use only Africans, but finding this impossible,
they imported twenty thousand coolies from India. The coolies came
on contracts of from two to five years, at wages of from four to fifteen
dollars a month and rations. The native labourers were paid about
ten cents a day.

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