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Cephalopods of the Broad Caribbean

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Title:
Cephalopods of the Broad Caribbean distribution, abundance, and ecological importance
Physical Description:
Book
Language:
English
Creator:
Judkins, Heather L
Publisher:
University of South Florida
Place of Publication:
Tampa, Fla
Publication Date:

Subjects

Subjects / Keywords:
Mollusca
Caribbean diversity
Biogeography
Latitudinal gradients
Coastal Atlantic Ocean
Dissertations, Academic -- Marine Science -- Doctoral -- USF   ( lcsh )
Genre:
non-fiction   ( marcgt )

Notes

Summary:
ABSTRACT: The Broad Caribbean region is defined as the Gulf of Mexico, and the coastal and marine areas of the Caribbean Sea, including the chain of islands forming the Greater and Lesser Antilles, Turks and Caicos, the Bahamas, and the gulf coasts of the United States, Central and South America (Stanley, 1995). The cephalopods of the Broad Caribbean were examined in terms of distribution, abundance, and ecological importance. A suite of 5190 preserved cephalopod specimens were identified and catalogued to produce regional maps of cephalopod distribution within the Broad Caribbean. Eighteen range extensions were noted for known species. Regional species richness was examined with respect to Rapoport's Rule with an eye toward possible cephalopod hotspots in the region. Cephalopods of the Broad Caribbean within the latitudinal bands of 8°N and 30°N do not support Rapoport's Rule as they exhibit increasing species richness with increasing latitude.Eight subareas were chosen to compare species richness. Regionally, species richness is patchy, with the largest concentration of cephalopods off the eastern Florida coast. Areas of the southern Caribbean Sea are in need of more samples for accurate assemblage counts and more meaningful comparisons with other Caribbean regions. Rarefaction curves were used to normalize the variously sized samples throughout the Broad Caribbean. A checklist of the Gulf of Mexico based on literature developed a picture for the northern regions of the Broad Caribbean. This checklist provided an updated account of cephalopod species that were reported from smaller literature works. Lastly, the first observation in the North Atlantic Ocean of the deep-sea squid Asperoteuthis acanthoderma (family Chiroteuthidae) was described. The description is based on two nearly intact, but damaged, specimens that were found floating at the surface in the waters off Key West and Marathon, Florida in 2007.All previously known records are recorded from a few specimens scattered in the western Pacific Ocean. There is a need for increased sampling throughout the Broad Caribbean to explore the systematics, life histories, distribution patterns, and potential fisheries for this group of organisms.
Thesis:
Dissertation (Ph.D.)--University of South Florida, 2009.
Bibliography:
Includes bibliographical references.
System Details:
Mode of access: World Wide Web.
System Details:
System requirements: World Wide Web browser and PDF reader.
Statement of Responsibility:
by Heather L. Judkins.
General Note:
Title from PDF of title page.
General Note:
Document formatted into pages; contains 147 pages.
General Note:
Includes vita.

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University of South Florida Library
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University of South Florida
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All applicable rights reserved by the source institution and holding location.
Resource Identifier:
aleph - 002063932
oclc - 558634848
usfldc doi - E14-SFE0003021
usfldc handle - e14.3021
System ID:
SFS0027338:00001


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ABSTRACT: The Broad Caribbean region is defined as the Gulf of Mexico, and the coastal and marine areas of the Caribbean Sea, including the chain of islands forming the Greater and Lesser Antilles, Turks and Caicos, the Bahamas, and the gulf coasts of the United States, Central and South America (Stanley, 1995). The cephalopods of the Broad Caribbean were examined in terms of distribution, abundance, and ecological importance. A suite of 5190 preserved cephalopod specimens were identified and catalogued to produce regional maps of cephalopod distribution within the Broad Caribbean. Eighteen range extensions were noted for known species. Regional species richness was examined with respect to Rapoport's Rule with an eye toward possible cephalopod hotspots in the region. Cephalopods of the Broad Caribbean within the latitudinal bands of 8N and 30N do not support Rapoport's Rule as they exhibit increasing species richness with increasing latitude.Eight subareas were chosen to compare species richness. Regionally, species richness is patchy, with the largest concentration of cephalopods off the eastern Florida coast. Areas of the southern Caribbean Sea are in need of more samples for accurate assemblage counts and more meaningful comparisons with other Caribbean regions. Rarefaction curves were used to normalize the variously sized samples throughout the Broad Caribbean. A checklist of the Gulf of Mexico based on literature developed a picture for the northern regions of the Broad Caribbean. This checklist provided an updated account of cephalopod species that were reported from smaller literature works. Lastly, the first observation in the North Atlantic Ocean of the deep-sea squid Asperoteuthis acanthoderma (family Chiroteuthidae) was described. The description is based on two nearly intact, but damaged, specimens that were found floating at the surface in the waters off Key West and Marathon, Florida in 2007.All previously known records are recorded from a few specimens scattered in the western Pacific Ocean. There is a need for increased sampling throughout the Broad Caribbean to explore the systematics, life histories, distribution patterns, and potential fisheries for this group of organisms.
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Cephalopods of the Broad Caribbean: Di stribution, Abundance, and Ecological Importance by Heather L. Judkins A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy College of Marine Science University of South Florida Co-Major Professor: Joseph Torres, Ph.D. Co-Major Professor: Michael Vecchione, Ph.D. Susan Bell, Ph.D. David Mann, Ph.D. Mark Luther, Ph.D. Date of Approval: June 10, 2009 Keywords: mollusca, Caribbean diversity, biogeography, latitudinal gradients, coastal Atlantic Ocean Copyright 2009, Heather L. Judkins

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ACKNOWLEDGEMENTS I would like to acknowledge th e team of people who have helped me through this process. Dr. Joseph Torres, Dr. Michael Vecchione, and my entire committee, Dr. Susan Bell, Dr. David Mann, and Dr. Mark Luther, have been instrumental in terms of advice, pathways to follow, and excellent sounding boa rds for my many questions. I would like to acknowledge Dr. Clyde Roper for his foresigh t in seeing that I had potential to add a small part of research to the cephalopod scientific community and his words of wisdom and encouragement throughout this entire vent ure. I would like to thank Dr. Nancy Voss for her advice and guidance in understa nding of cephalopod species of the Broad Caribbean. Her assistance in my work is be yond compare. Dr. Ronald B. Toll and Dr. Steven C. Hess were instrumental in documenting cephalopod specimen information from the Hourglass Cruises that were conducted from 1965 through 1967, by the Marine Research Laboratory of the Florida Board of Conservation on the continental shelf of the Gulf of Mexico which was included in my di ssertation. I also thank Teresa Greely and Angela Lodge for their assistance thr oughout the two years of my GK12 Oceans Fellowship and the opportunity to coordinate the regional competition for the National Ocean Sciences Bowl that I have been so excite d to be a part of for the College of Marine Science at USF. Lastly, I wouldnt be where I am without the unfailing support and constant source of smiles from my family and friends who have been there through the ups and downs of my journey. Thank you.

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i TABLE OF CONTENTS TABLE OF CONTENTS i LIST OF TABLES iii LIST OF FIGURES iv ABSTRACT v CHAPTER 1. INTRODUCTION 1 1.1 Broad Caribbean 1 1.2 Cephalopod Biology 3 1.3 Distribution of Cephalopods 4 1.4 Abundance of Cephalopods 4 1.5 Cephalopods in Food Webs 5 1.6 Overall Significance 7 1.7 Study Goals 8 1.8. Study Area 9 1.9 Materials and Methods 9 1.10 References 13 CHAPTER 2. BIOGEOGRAPHY OF CEPHALOPODS OF THE BROAD CARIBBEAN 16 2.1 Introduction 16 Broad Caribbean 16 The Cephalopods 18 Biogeography of Cephalopods 18 Abundance of Cephalopods 21 Ecological Focus 21 Study Area 24 2.2 Materials and Methods 24 Statistical analysis 26 2.3 Results 27 Rapoports Rule and Species Richness 27 Cephalopod Biogeography 28 Distribution and Abundance 29 Range Extensions 29 New Species to Area 38 2.4 Discussion 38 Rapoports Rule and Species Richness 38 Cephalopod Biogeography 41

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ii Distribution and Abundance 43 Range Extensions 45 New Species 45 2.5 References 47 CHAPTER 3. CEPHALOPODA (MOLLUSCA: CEPH ALOPODA) OF THE GULF OF MEXICO 52 3.1 Introduction 52 3.2 Major Systematic Revisions 56 3.3 Comparative Assessment of the Gr oup in the Gulf of Mexico 56 3.4 Explanation of Checklist 57 3.5 Acknowledgements 60 3.6 References 61 3.7 Gulf of Mexico Checklist 66 3.8 Taxonomic Summary for Cephalopods of the Gulf of Mexico 73 CHAPTER 4. FIRST RECORDS OF Asperoteuthis acanthoderma (Lu, 1977) (CEPHALOPODA: OEGOPSIDA: CHIROTEUTHIDAE) FROM THE NORTH ATLANTIC OCEAN; STRAITS OF FLORIDA 74 4.1 Abstract 74 4.2 Introduction 75 4.3 Systematics 76 4.4 Description 76 4.5 Discussion 81 4.6 Acknowledgements 86 4.7 References 87 4.8 Figure Legends 88 CHAPTER 5. CONCLUSIONS 94 5.1 Summary 94 5.2 References 100 APPENDICES 102 Appendix A: Regional distri bution point coordinates 103 Appendix B: Species ric hness comparison quadrats 104 Appendix C: Cephalopod distribution maps 105 ABOUT THE AUTHOR End page

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iii LIST OF TABLES Table 2.1 Regional cephalopod locations 20 Table 2.2 Region, species number and sample number for hotspots 33 Table 2.3 Cephalopod range extensi ons to Broad Caribbean 35 Table 3.1 Checklist of the Cephalopods of the Gulf of Mexico 66 Table 3.2 Taxonomic summary for cephal opods of the Gulf of Mexico 73 Table 4.1 Measurements of Asperoteuthis acanthoderma specimens 89 Table 5.1 Gulf of Mexico checklist species not included in Ch. II 95

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iv LIST OF FIGURES Figure 1.1 Current flow through the Cari bbean Sea and Gulf of Mexico 2 Figure 1.2 A summary of the role of cephalopods in the worlds oceans 6 Figure 1.3 Study Area 9 Figure 2.1 Current flow through the Caribbean Sea and Gulf of Mexico 17 Figure 2.2 Study area 24 Figure 2.3 Species richness per 5 latitudinal band 30 Figure 2.4 Comparison of species richne ss and number of individuals 30 Figure 2.5 Rapoports RuleRarefaction curve 31 Figure 2.6 Rapoports RuleChao 1 estimator graph 31 Figure 2.7 Eight study sites for richne ss and diversity comparisons 32 Figure 2.8 Species richness regional comparison 32 Figure 2.9 Hotspot species observe dRarefaction curve 33 Figure 2.10 Hotspot Chao 1 estimator of regional species richness 34 Figure 2.11 Broad Caribbean sample effort map 34 Figure 4.1 Key West Asperoteuthis acanthoderma specimen 91 Figure 4.2 Marathon Asperoteuthis acanthoderma specimen 92 Figure 4.3 Internal organs of th e Key West specimen 93

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v CEPHALOPODS OF THE BROAD CARIBBEAN SEA: DISTRIBUTION, ABUNDANCE, AND ECOLOGICAL IMPORTANCE Heather L. Judkins ABSTRACT The Broad Caribbean region is defined as the Gulf of Mexico, and the coastal and marine areas of the Caribbean Sea, including the chain of islands forming the Greater and Lesser Antilles, Turks and Caicos, the Bahamas, and the gulf coasts of the United States, Central and South America (Stanley, 1995). The cephalopods of the Broad Caribbean were examined in terms of distribution, a bundance, and ecological importance. A suite of 5190 preserved cephalopod specimens were identified and catalogued to produce regional maps of cephalopod distribution with in the Broad Caribbean. Eighteen range extensions were noted for known species. Re gional species richness was examined with respect to Rapoports Rule with an eye toward possible cephalopod hotspots in the region. Cephalopods of the Broad Caribbean within the latitu dinal bands of 8 N and 30 N do not support Rapoports Rule as they ex hibit increasing sp ecies richness with increasing latitude. Eight subareas were chosen to compare species richness. Regionally, species richness is patchy, w ith the largest concentration of cephalopods off the eastern Florida coast. Areas of the southern Cari bbean Sea are in need of more samples for accurate assemblage counts and more meaningful comparisons with other Caribbean regions. Rarefaction curves were used to normalize the variously sized samples throughout the Broad Caribbean. A checklist of the Gulf of Mexico based on literature developed a picture for the nor thern regions of the Broad Caribbean. This checklist

PAGE 8

vi provided an updated account of cephalopod sp ecies that were reported from smaller literature works. Lastly, the first observation in the North At lantic Ocean of the deep-sea squid Asperoteuthis acanthoderma (family Chiroteuthidae) was described. The description is based on two nearly intact, but damaged, specimens that were found floating at the surface in the waters off Key West and Marathon, Florida in 2007. All previously known records are recorded from a few specimens scattered in the western Pacific Ocean. There is a n eed for increased sampling thr oughout the Broad Caribbean to explore the systematics, life hi stories, distribution patterns, an d potential fisheries for this group of organisms.

PAGE 9

1 CHAPTER 1 INTRODUCTION Cephalopods of the Broad Caribbean have not been reviewed in recent years. The unique features, currents, and coastlines of th e Gulf of Mexico, Caribbean and the Straits of Florida combine as a potentially un ique ecotone for cephalopod distribution, abundance, and diversity. Work conducted by G. Voss (1956), C.F. Roper (1984), N. Voss (1998), M. Vecchione (2002), and ot hers provide background for further exploration of cephalopods in this ar ea. The present study focuses on over 5000 specimens collected from the area to extend cephalopod research. 1.1 The Broad Caribbean The Broad Caribbean region is defined as the Gulf of Mexico, and the coastal and marine areas of the Caribbean Sea, including the chain of islands forming the Greater and Lesser Antilles, Turks and Caicos, the Bahama s, and the Gulf coasts of North, Central and South America (Stanley, 1995). The region is influenced by waters that flow through the lower Caribbean islands, originating from the Guiana Current that m oves north along Brazils coast. The Guiana Current is joined by the North Equatori al Current, which flows through the lower Caribbean, veering north around western Cuba and into the Gulf of Mexico. Some upwelling occurs along the southern region of the Caribbean Sea (Longhurst, 1998). The majority of the flow moves around the Gulf coasts of the United States, flowing down along the west Florida coast before moving thro ugh the Straits of Florida to become the

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2 Gulf Stream which moves northward through the Bahamas and along the eastern coast of Florida. The general movement of currents in the Broad Cari bbean is from east to west with gyres often spinning off th e main water flow (Stanley, 1995). Fig 1.1: Major currents of Broad Cari bbean (from Carpenter et al. 2002) The surface temperature of the ocean in th e tropical half of the Broad Caribbean region is averages 27 C with little variati on throughout the year. Temperatures in the southern portion of the Gulf of Mexico also average near 27 C but the northern Gulf of Mexico has larger temperature fluctua tions due to seasona l changes from 16 C in winter to 28 C in summer (Stanley, 1995). Salinity is relatively high in the su rface waters between January and May, 36.36, and lower between June and December in the region, 36.06 (Tomczak & Godfrey, 1994), mainly due to the inflow in late fall of lower-salinity waters from the Orinoco and Amazon Rivers as well as equatorial convergence coming through the region. Geologically, the Caribbean Sea consists of two main basins separated by a broad, submarine plateau. Two tectonic plates, the Caribbean plate and the North American

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3 plate, potentially influence hi storical biogeography in the area. The Cayman Trench, a trench between Cuba and Jamaica, contai ns the Caribbeans deepest point (7,535 m below sea level). 1.2 Cephalopod Biology The cephalopods are molluscs with many uni que features separating them from other molluscan groups including a closed circ ulatory system, the reduction and in most cases, absence of an external shell, a sophisticated ner vous system, jet propulsion for movement, many camouflage adaptations, a nd a predatory lifestyle (Brusca, 2003). Cephalopods possess a biting apparatus resembling an inverted parrot beak, and almost all have some form of radula, or rasping tongue. Food-capturing mechanisms vary from sucker-bearing arms and tentacles to hooks modified from suckers for catching prey. In particular, nektonic cepha lopods are impressive predators. Octopods are adept in stalking or hiding out to attack prey. Neritic octopods home in visually on prey and they approach by partially raising an ar m and gliding in that direction. The attack is a pounce during whic h many of the arms and the web are thrown over the prey to immobilize it. The octopods will then bite and poison the organism (Vecchione, 2002). Although research is scare in terms of the life history of many cephalopod species, the consensus is that they possess a fast growth rate and rela tively short lifespan. They grow exponentially until mature and then they spawn and die. Cephalopods have various spawning methods ranging from polyc yclic spawning and multiple spawning to intermittent terminal and continuous spawning. All spawning strategies end in the death of the cephalopods after the spawni ng period (Jereb & Roper, 2005).

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4 1.3 Distribution of Cephalopods Past cephalopod studies have focused on small pockets of the Broad Caribbean such as the Gulf of Mexico and Florida re gions by Voss (1956), the Straits of Florida by Cairns (1976), the eastern Gulf of Mexic o, Passerella (1991), and regions surrounding Colombia, by Gracia (2002). Cephalopods have various distributi on patterns ranging from coastal species to entirely pelagic. Coastal species include the economically important lologinids and octopods Offshore, the ommastrephids and histioteuthids are of greater importance. Cephalopod species exhibit considerable di versity in their depth distribution as well. Squid, such as Doryteuthis are surface dwellers living within 500m of the ocean surface while chiroteuthids are deep-sea or ganisms with ranges down to 4000m and beyond. There are temperature tolerance patt erns that exist among cephalopods; certain cephalopods thrive in the warmer wa ters of the Caribbean proper ( Octopus zonatus, Octopus maya ) while others have an affinity for the subtropical temperatures of the northern sections of the Gu lf of Mexico such as Ancistroteuthis lichensteinii Distribution patterns are importa nt for researchers and fisher y experts to understand as conservation measures and management prac tices are created fo r cephalopod groups. 1.4 Abundance of Cephalopods : Since their appearance in the Cambria n, cephalopods have evolved to include some of the largest past a nd present invertebrates (over 20m) and to become common predators in all shallow and deep seas. Fo r part of lifes history, over 200 million years, they were probably the top predators in the mari ne environment. After the Jurassic, it is believed that fishes superseded their impor tance as predators. While fishes and

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5 cephalopods both evolved to cope with the same physical marine environment, much stimulus to cephalopod evolution also came from their interaction with fish and later with other vertebrate predators, the reptiles, s eals, cetaceans, and birds (Clarke, 1996). Cephalopod species vary in abundance in the Broad Caribbean ranging from schools of thousands in Illex species to solitar y individuals of Discoteuthis discus and Asperoteuthis acanthoderma Cephalopods today are not as numerous as they once were due to the constant struggle be tween them and all other mari ne species, including other cephalopods. Their present success is appa rent in their morphological diversity, abundance, and major role they play in the ecology of the sea (Young et al. 1998). 1.5 Cephalopods in the food web Cephalopods are important predators on micronekton and macrozooplankton and a major food source for nektonic fish. Diets of cephalopods change as they mature. Most begin as paralarvae preying upon small crustacea ns, and as they grow, move onto larger crustaceans, fish, and other mollusca. Sm all, immature longfin inshore squid ( Doryteuthis pealeii ), for example, will feed on planktonic or ganisms while larger individuals feed on larger crustaceans and small fish. Studies ha ve also shown that the immature squid will feed on euphausiids and arrow worms while adu lts feed on small crabs, polychaetes, and shrimp (Cargnelli, 1999). Fish species prey ed on by the longfin inshore squid include silver hake, mackerel, herring, menhaden, sand lance, bay anchovy, weakfish, and silversides. As a food source for other orga nisms, cephalopods have been found in large numbers in the diets of seabirds, seals, whales, and fish (Cargnelli, 1999). Prey preference varies among species. The congeners, Doryteuthis opalescens (Pacific) and Doryteuthis pealeii (Atlantic) are important commercial squid with

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6 different adult sizes and prey preferences A study by Karpov and Cailliet (1978) found that Doryteuthis opalescens fed mainly on fish, ce phalopods, gastropods, and polychaetes. Doryteuthis pealeii were found to eat more equal proportions of fish and fellow squid (Amaratunga, 1983). Figure 1.2 A summary of the role of cephalopods in the worlds oceans and seas as expressed by their position in the en ergetic hierarchy. (Clarke, 1996) Cephalopods are born into the third trophic level and progress one to two stages through their life. Research has not shown them achieving to p-predator status because there always seems to be some vertebrate that preys upon them (Summers, 1983). Diving birds such as penguins and murres actively search for cephalopods as part of their diet as do toothed whales and seals. Sei and Fin whales are the major baleen predators on oceanic squid. Those whales seek copepods and euphausiids, which are probably also the prey of squids. Examples of seabirds dependent on cephalopods would be the thick-

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7 billed murres of the North Pacific and the emperor penguins of the Antarctic (Williams, 1995). Ogi lists squid as 72.6% of the diet of thick-billed murres (Summers, 1983). 1.6 Overall significance: Cephalopods play an important role in the Broad Caribbean region in terms of fisheries and overall biodiver sity. At present, cephalopods contribute only 3% to the tonnage of global fisheries, but their total value lies third, below only shrimps and tuna (Clarke, 1996). Cephalopods are important as a food source for humans and well over 3 million tons are caught yearly. Fishing has b ecome increasingly intense in the Orient and the eastern Atlantic Ocean. Most of the ce phalopods caught in the Broad Caribbean were landed by Mexican fishermen, bringing in between 19,000 and 31,000 tons (Vecchione, 2002). Ideally, a thorough knowledge of the system atics of a species is the required foundation upon which all other biological a nd resource management studies must be based. For example, in the Gulf of Campeche, Mexico, a traditional fishery was believed to be based on Octopus vulgaris Lamarck, 1798, a ubiquit ous octopus of broad distribution. In the absence of local studies, knowledge about the biology of O. vulgaris from other seas was applied to the Campeche octopus for fishery statistics and management purposes. The discovery th at the octopus was actually a new species, described as O. maya Voss and Solis, 1966, with a very different life hi story, explained the problems that had plagued biologists assigned to study the fishery and develop recommendations. This example underscore s the need for sound systematic knowledge of species and populations if we are to regulate fisheries for these forms (Roper, 1983).

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8 Biodiversity studies are m oving from a focus on indi vidual species to the ecosystem level. Similarly, fisheries manageme nt has progressed from single species to multiple target species to ecosystems (such as the large marine ecosystem approach of Sherman et. al. 1990). For an ecosystem ma nagement effort to have any chance of success, information is needed on all a bundant or ecologically important species (Vecchione and Collette, 1996). Fisheries manage ment is an integral part of ecosystem management practices. Changes in both ta rgeted and bycatch species need to be monitored. An example is the fishery fo r bottom fish on Georges Bank. Of the fish caught in 1963, 67% were the prized cod, hake, and flounders, whereas 24% was made up of unwanted species. By 1986 the dominant catch had shifted dr amatically, with 14% wanted species and 74% junk species. Such changes in populations of large predators cause profound effects throughout the f ood web (Vecchione and Collette, 1996). 1.7 Study Goals The goals of the present study include el ements of biogeography, distribution, and ecology of cephalopods. They are: 1. To determine the distribution and ab undance of cephalopod species in the Broad Caribbean region. 2. To discern general biogeographic pa tterns of the group in the Broad Caribbean region. 3. To examine the distribution of the gr oup for possible alignment to Rapoports Rule (RR) as studied by other au thors (Macpherson 2002, Fortes 2004, Rosa et al., 2008). 4. To create a comprehensive checklist of th e major species found in the Gulf of Mexico. 5. To describe new species to the region. This includes range extensions for currently known species in the area.

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9 1.8 Study Area The Broad Caribbean includes the Bahama Islands, Gulf of Mexico, and the Caribbean Sea. The range for the study was between 8 N -30 N and 58 W 97 W. Figure 1.3: Study Area 1.9 Materials and Methods The most helpful taxonomic information include examination of comparative material from a variety of locations, incl uding from the type locality when possible (Vecchione; Collette, 2000). Based on cu rrent cephalopod dichotomous keys, type specimens, literature, and expert opinions, the specimens analyzed were identified to the species level. Once the specimen was identi fied, it was plotted on a distribution map of the Broad Caribbean using the ArcView 9.2 mapping program. An acceptable biogeographic study includes many specimens of each species under consideration, including all the life-histor y stages, precise distribution within the

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10 study area, and a complete geographic range of each species. This study included 4190 specimens that had known locales and 1000 se condary regional specimens that were distributed throughout the Broad Caribbean for a total o 5190 specimens examined. Many of the regional specimens were from the Caribbean Sea and were important to include. The regional data was used in the analysis of the distributions of cephalopods but not included for the application of RR. A key component of biodiversity explora tion is the discovery of new species. Some factors that affect proper iden tification of cephalopods are the: Basis for identification: recent compre hensive reviews are more reliable. Experience level of identifiers: some taxonomic groups are more difficult than others to identify. Difficulty of characters necessary for confident identification. Life-history stages examined. Conditions of specimens: quality can be greatly affected by methods used for collection, fixation and preservation, pot entially limiting or eliminating the usefulness of important characteristics. Possible presence of similar, easily confused species: confidence in identification is limited. (Vecchione and Collette, 2000) The above six factors were taken in to consideration while the study was conducted. Identification guides (Voss 1956; Nesis 1975, 1987; Roper and Young, 1984; N. Voss 1998; Vecchione, 2002) were used to id entify each organism to species level. The majority of the specimens examined had been preserved well a nd were in excellent condition. The specimens date from 1898 to present. Identification of specimens was conducte d in the micronekton museum at the College of Marine Science at the University of South Flor ida, the Rosenstiel School for Marine and Atmospheric Science, University of Miami as well as the National Museum of Natural History at the Smithsonian Inst itution in Washington, D.C.. A dissecting

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11 microscope was used to identify small specim ens and specific features of the animals. Micrometer calipers were used for lengt h measurements. The bulk of preserved specimens analyzed were from two institutions, the Smithsonian Institutions National Museum of Natural History and the University of Miamis Rosenstiel School of Marine and Atmospheric Science. Smaller collec tions from the Florida Fish and Wildlife Research Institute were also analyzed. The methods used for identifying and catal oguing specimens were to identify the individual using the cephalopod dichotomous ke ys, and to record its mantle length, sex, number of gill lamellae (octopods), and species name into an excel file for statistical analysis. If the specimen was in a mixed lo t, individuals were separated according to species. All station and cruise informati on was copied and put into the additional collection jars as need ed. Once all the organisms were identified, known specimens from all institutions were also added to the excel file for analysis. Di stribution was calculated using the 9.2 ArcView software program. Chapter II of the dissertation focuses on the biogeography of cephalopods in the Broad Caribbean region. Species richness of cephalopods was analyzed for alignment to Rapoports Rule which was proposed by Stevens (1989): that species ri chness tends to be greater towards lower latitude s. Potential cephalopod hotsp ots are compared within 8 subareas of the Broad Caribbean and lastl y, distribution of all species examined are plotted on regional maps of the area. Ei ghteen range extensions are added to the cephalopod database for the region. The third chapter of this study utilizes cephalopod literature as the basis for a species checklist of the Gulf of Mexico. Comprehensive studies of large oceanic regions are not pr esent in current cephalopod research. The

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12 chapter updates Voss (1956) and utilizes over 40 cephal opod studies of the area to compile the checklist. Chapter IV describe s a range extension of the chiroteuthid, Asperoteuthis acanthoderma, a species newly found in the Broad Caribbean. It describes two specimens discovered floating dead at the surface off the coast of Key West and Marathon, Fl in 2007. Both specimens were examined by the author in collaboration with experts in the field. Unique defining characteristics were used for identification with emphasis on the Y-shaped funnel locking mechanism, sucker ring form and dentition, beak morphology, photophore patch configuration on ventral surface of eyeballs, and the numerous small cartilaginous tubercles that cover the mantle, head and aboral surface of the arms. The concluding chapter is an overall summary of the Broad Caribbean cephalopod assemblage.

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13 REFERENCES 1. Amaratunga, T.1983. The Role of Cephalopods in the Marine Ecosystem. Advances in assessment of world cephalopod resources; 231 : 379-415. 2. Brusa, Richard C., Gary J. Brusca. 2003. Invertebrates, Second edition; Sinauer Associates, Inc. Publishers; Sunderland, Ma; p. 714-760. 3. Cairns, S. 1976. Cephalopods collected in the Straits of Florida by the R/V Gerda. Bulletin of Marine Science; 26(2): 233-272. 4. Cargnelli, L., Sara Griesbach, Cathy Mc Bride,Christine A. Zetlin, and Wallace W. Morse, 1999. Longfin Inshore Squid, Loligo peal eii, Life History and Habitat Characteristics in NOAA Technical Memorandum NMFS-NE-146 NOAA: Northeast Region, Woods Hole, MA. 1-27. 5. Clarke, M.R. 1996. The role of cephalopods in the wo rlds oceans: an introduction. Phil.Trans. R. Society London. B.; 351: 979-983. 6. Fortes, R.R., R.S. Absalao, 2004. The applicability of Rapoports rule to the marine molluscs of the Americas. Journal of Biogeography; 31: 1909-1916. 7. Gracia, A. 2002. Cephalopods (Mollusca: Cephalopoda) of the Upper Colombian Caribbean Shelf. Biol. Invest. Mar. Cos. 31, 219-238. 8. Jereb, P. and Roper, C.F.E. 2005. Cepha lopods of the world. An annotated and illustrated catalogue of cephalopod species known to date. FAO species catalogue for fisheries purposes, N4, v.1, FAO, Rome. 9. Karpov K.A., & G.M. Calliet. 1978. Feed ing dynamics of Loligo opalescens. Calif. Dept. of Fish and Game Bulletin; 169: 45-65. 10. Longhurst, A.1998. Ecological geography of the sea. Academic Press; 146-150. 11. Macpherson, E. 2002. Large-scale species-ric hness gradients in the Atlantic Ocean. Proceedings of the Royal Society of London; B; 269: 1715-1720. 12. Nesis K.N. 1975. Cephalopods of the American Mediterranean Sea; English Translations of Selected Publica tions on Cephalopods; Editor M. Sweeney; Smithsonian Institutions Libraries; 1; 318358. -----1987. Cephalopods of the world. (Edited by L. Burgess) T.F.H. Publication, Inc. Ltd. New Jersey; 351 pp.

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14 13. Passarella, K.C., 1990. Oceanic Cephalopod Assemblage in the Eastern Gulf of Mexico in Department of Marine Science University of South Florida: St. Petersburg, Fl. 50pp. 14. Roper, C.F.E., 1983. An overview of cephalopod system atics: status, problems, and recommendations. Memoirs of the National Museum Victoria.; 44: 13-27. 15. Roper, C. F. E., M. J. Sweeney, and C. Nauen. 1984. Cephalopods of the World. An Annotated and Illustrated Ca talogue of Species of Inte rest to Fisheries. FAO Fisheries Synopsis No.125, v.3, Food and Agricultural Organization of the United Nations, Rome. 277 pp. 16. Rosa, Rui, Heidi M. Dierssen, Liliana G onzalez, and Brad Seibel. 2008. Ecological biogeography of cephalopod molluscs in th e Atlantic Ocean: historical and contemporary causes of coastal dive rsity patterns; Global Ecology and Biogeography; 17: 600-610. 17. Sherman K., L.M. Alexander & B.D. Go ld. Editors.1990. Large marine ecosystems: patterns, processes, and yields. Ameri can Association for the Advancement of Science, Washington D.C. 18. Stanley, Sonja (editor), 1995. A Global Re presentative System of Marine Protected Areas: Vol 2, Marine Region 7: Wider Cari bbean, Great Barrier Reef Marine Park Authority, The World Bank, The Worl d Conservation Union, (IUCN); 15-42. 19. Stevens, G.C., 1989. The Latitudinal Gradient in Geographical Range: How So Many Species Coexist in the Tropics. The American Naturalist; 133(2): 240-256. 20. Summers, W.C., 1983. Physiological and Trophic Ecology of Cephalopods The Mollusca. Ecology; 6: 261-279. 21. Tomczak, Mattias, J. Stuart Go dfrey. 1994. Regional Oceanogrpahy: An Introduction. Pergamon; 310-317. 22. Vecchione, M., Bruce B. Collette, 1996. Fisheries Agencies and Biodiversity. Ann. Missouri Bot. Gard; 83: 29-36. 23. Vecchione, M., M.F. Mickevich, K. Fauchald, B.B. Collette, A.B. Williams, T.A. Munroe, and R.E. Young, 2000. Importance of Assessing Taxonomic Adequacy in Determining Fishing Effects on Marine Diversity. ICES Journal of Marine Science; 57: 677-681.

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15 24. Vecchione, 2002. Cephalopods. Carpenter, K.E.(ed.) The living marine resources of the Western Central Atla ntic. Volume 1: Introducti on, molluscs, crustaceans, hagfishes, sharks, batoid fishes, and chimaeras. Cephalopods of Western Central Atlantic; FAO Species Identification Guide fo r Fisheries Purposes and American Society of Ichthyologists and Herpetologi sts Special Publication No. 5. Rome, FAO. 2002. 1-600. 25. Voss, G.L. 1956. A review of the cephal opods of the Gulf of Mexico. Bull Mar. Sci. Gulf and Caribbean; 6:85-178. 26. Voss, N.A., Vecchione, M., Toll, R.B. & Sweeney, M.J.(eds) 1998. Systematics and biogeography of cephalopods. Smithson, Contrib. Zool.; 586: 1-599. 27. Williams, T.D. 1995. The penguins: Spheniscid ae. Oxford University Press, Oxford. 28. Young, R. E, L. A. Burgess, C. F. E. Roper, M. J. Sweeney, and S. J. Stephen 1998. Classification of the Enopl oteuthidae, Pyroteuthidae, and Ancestrocheiridae. Pp. 239 in N. A. Voss, R. B. Toll, and M. Vecchione, eds. Systematics and Biogeography of Cephalopods. Smithsoni an Contributions to Zoology 586(1).

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16 CHAPTER 2 BIOGEOGRAPHY OF CEPHALOPODS OF THE BROAD CARIBBEAN REGION 2.1 Introduction Cephalopod studies in the Caribbean re gion have not provi ded a well-rounded, comprehensive view of distribution and a bundance in the group. Various island groups or individual cephalopod species have been addressed, (Diaz, 2000, 2004; Gracia, 2002) mainly in coastal waters, but to date, no study describes the Broad Caribbean species complex as a whole. Rosa et al. (2008) and Smith et al. (200 2) conducted literaturebased studies on latitudinal gr adients of species richness but none have utilized large numbers of specimens from the region to improve our understanding of cephalopod ecology. This paper fills that need an d based on 5190 specimens, reports on the distribution, abundance, and di versity of cephalopods in th e Broad Caribbean region. The Broad Caribbean The Broad Caribbean region is defined as the Gulf of Mexico and the coastal and marine areas of the Caribbean Sea, including the chain of islands forming the Greater and Lesser Antilles, Turks and Caicos, the Bahamas, and the gulf coasts of the United States, Central and South America (Stanley, 1995) The Caribbean Sea proper covers approximately 1,063,000 square miles while the Gu lf of Mexico is smaller, covering an area approximately 1,592,842 square kilometers (615,000 sq. miles).

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17 The region is influenced by waters that flow through the lower Caribbean islands, originating from the Guiana Current that m oves north along Brazils coast. The Guiana Current is joined by the North Equatori al Current, which flows through the lower Caribbean, veering north around western Cuba and into the Gulf of Mexico. Some upwelling occurs along the southern regi on of the Caribbean Sea (Longhurst, 1998). Loop Current water moves through the Gulf of Mexico, flowing down the west Florida coast before moving through the Straits of Flor ida. The water becomes the Gulf Stream, which moves northward through the Bahamas and eastern coast of Flor ida. The general movement of the Broad Caribbean is from eas t to west with gyres often spinning off the main water flow. (Stanley, 1995) (Fig. 2.1) Figure 2.1: current flow through Caribbe an Sea and Gulf of Mexico (from Carpenter et al. 2002) The surface temperature of the ocean in th e tropical half of the Broad Caribbean region averages 27 C with little variation throughout the year. Temperatures in the southern portion of the Gulf of Mexico also average near 27 C but the northern Gulf of Mexico has larger temperature fluctua tions due to seasona l changes: from 16 C in winter

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18 to 28 C in summer (Stanley, 1995). Salinity is relatively hi gh between January and May (36.39) and lower between June and December in the surface waters of the region (36.09) (Tomczak & Godfrey, 1994), due mainly to the inflow in fall of lower-salinity waters from the Orinoco and Amazon Rivers. Geologi cally, the Caribbean Se a consists of two main basins separated by a broad, submarine plateau. The Cayman Trench, a trench between Cuba and Jamaica, contains the Cari bbeans deepest point (7,535 m) and divides two tectonic plates (Stanley, 1995). The Cephalopods Fewer than 1000 species of living cephal opods have been described worldwide; over 720 are listed in the present catalogues (Jer eb et al., 2005). Cephalopods occur in all marine habitats, though none are found at sa linities less than 17.5. Their depth range extends from the intertidal to over 5,000 m. Due to their acc essibility, many of the near surface and coastal cephalopod sp ecies of the Greater Caribbean have been studied in detail (Voss 1956, 1973; LaRoe, 1967; Lipka, 1975; Passarella, 1990). Deep-sea species are more difficult to study because of ne t avoidance and other escape tactics by the cephalopods (Passarella, 1990). A diverse cephalopod fauna is associated with the bottom in both shallow and deep-seas. Biogeography of the Region Briggs (1995) divided the Broad Caribb ean into four distinct regions: the Brazilian Province, the Caribbean Province, the West Indian Province and the Carolina Region. The Brazilian Province occupies the area between the Orinoco delta and Cape Frio which has a distinct biota because of the habitat change. Almost the entire northeastern coast of South America is virtua lly devoid of coral reef s and there are vast

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19 stretches of mud bottom. Near the mouths of the rivers, salinity is greatly reduced (Briggs, 1995). The West Indian Province includes an extensive geographic area and consists entirely of islands. Bermuda is an isolated northern outpost while the main portion is an archipelago stretching from the Bahamas to Grenada. Recognition of a West Indian Province means that the Straits of Floridaonly 80 km wide, is an important barrier to the dispersal of the marine shore biota. It s eems clear that the barrier results not from distance, but from the fast-flowing Florida Current. The Yucata n Channel is only 180km wide and the passage between Grenada a nd Trinidad is about 100km wide (Briggs, 1995). The Caribbean Province extends sout hward from Cape Canaveral in eastern Florida, Cape Romano in western Florida, a nd Cape Rojo, Mexico. From those points, it follows the shore all the way to the northe rn edge of the Orinoco River delta in Venezuela. The west coast of Florida supports a complex biotic assemblage. From north to south, warm-temperate species reach their range limits and tropical species make their appearance at various points. There is also considerable faunal change with depth, the tropical species being more numerous on the outer edges of the shelf. The northern Gulf of Mexico is included as pa rt of the Carolina Region. Within this region, the warmtemperature biota occupies an area north of the tropical boundaries at Cape Romano, Fl, and Cape Rojo, Mexico (Briggs, 1995). Examples of Broad Caribbean cephalopod regional locations (Table 2.1) are as follows:

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20 Table 2.1: Examples of regional cephalopod locations (Clarke, 1996) Region: Species: The western central Atlantic (WCA) includes the world s greatest concentration of small countries including th e full range of the worlds majo r political systems. All of the Caribbean Sea and nearly a ll of the Gulf of Mexico are included within one or another of the regions 42 jurisdictiona l units, the largest number fou nd in any ocean area of this size. When the EEZs are compared to the geographic and ecological features of the same area, it becomes clear that the countri es of the region are faced with managing the biological outcomes of oceanic and ecological processes that ope rate on a scale that is far larger than any of the regions individual management units (Smith et al., 2002). Four commercially important squid sp ecies occur in the Caribbean: longfin squid ( Doryteuthis pealeii ), arrow squid ( Doryteuthis plei ), brief squid ( Lolliguncula brevis ), and southern shortfin squid ( Illex coindeti ). The sharptail shortfin squid ( Illex oxygonius ) is found as well in the Caribbean re gion but is often unrecognized in the catches. Octopus maya is a commercially important oct opod species. It is important to accurately identify and to determine the distri bution of commercially harvested species. Voss studied the seasonal distribution of th e commercially harvested species and found Inshore coastal region including continental shelf Octopodidaerocky or coral shores Sepiolidssand or mud Loliginidae SeasonallyOmmastrephidae Off slopes and island slopes spawning aggregations Surface water offshore Onychoteuthids, Argonautids, Ommastrephids Midwater Histioteuthids and Enoploteuthids

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21 that their distribution showed a relationship between squid occurrence and temperature, bottom topography, and areas of high pr oductivity (Voss & Brakoniecki, 1985). Abundance of Cephalopods Today, cephalopods are important in neri tic waters although numerically they only constitute a small part of the shelf fa una. In most nearshore regions they are outnumbered by fish of similar size, except during certain seasons and in some localities. In oceanic waters they are more diverse in size and play an important role in food webs (Clarke, 1996). Abundance of cephalopods varies (dependi ng on group, habitat, and season) from isolated territorial individuals (primarily benthic octopods) thr ough small schools with a few dozen individuals, to huge schools with millions of oceanic squids (Vecchione, 2002). Approximately 109 cephalopod species in 31 families occur in the Western Central Atlantic Ocean and ad jacent areas (Caribbean Sea an d Gulf of Mexico). Both decapods and octopods are common in those waters (Vecchione, 2002). Records of cephalopod species in the Gulf of Mexico da te back to Leseur (1821), but the modern comprehensive systematics of the group be gin with Voss, who reported 24 neritic and oceanic species in 1954, and 42 species in 1956. Since that time, many species have been added to the list (P assarella, 1990). Ecological Focus The ecological portion of this paper ex amines Rapoports Rule (RR) by focusing on small-scale patterns within a region that had been described as an ecotone. Stevens (1989) proposed that the greater biodiversity often seen in the tropi cs is explained by the fact that tropical species ha ve very narrow ranges while at higher latitudes there is a

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22 higher proportion of species with wider ranges (Stevens, 1989). He explained the pattern on the basis of differing tolerances of tr opical and temperate species to climatic variations. Organisms inhabiti ng lower latitudes are subject to less variation in climate, and therefore their geographical distributions tend to be limited to a narrow climatic range. Higher latitude species would be adapted to more marked climate variation (Fortes, 2004). Biodiversity for the purpose of this study is defined as species richnessnumbers of species per area examined. The species ri chness correlation is found in all higher taxa whose geographical ranges ar e well known, both terrestrial and marine. Rapoport (1982) had noted that the latitudina l ranges of individual species became smaller in lower latitudes. Thus more species could be acco mmodated at lower latitudes because each required less space. Many studies have tested RR and the outco mes have been mixed. Some authors (Steele, 1988, Macpherson, 2002; Roy et al. 199 8) have found evidence in their studies supporting RR while others (Clarke, 1992; Mokievsky & Azovsky, 2002) failed to find such a relationship (Rosa, et al. 2008). Fo rtes (2004) examined se lected bivalves and gastropods along the Pacific and Atlantic co asts of the Americas and attempted to evaluate the applicab ility of Rapoports rule in thos e regions; he concluded that Rapoports Rule does apply in those cases (Fortes, 2004). Other studies, such as one conducted by Rohde (1992), suggests that Rapoports Rule does not apply to all taxonomic categorie s. The study focused on marine teleosts using data collected from the Indo-Pacific a nd Atlantic oceans. Rohde (1992) found that RR does not apply in all range areas within a taxons range and that it may be premature

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23 to explain greater species numbers by narro wer environmental tolerances of tropical species (Rohde, 1992). The biogeographical pattern proposed by Stevens (1989) has acquired increasing importance among research ers as an explanation for the biodiversity gradient related to latitude (F ortes, 2004). Many studies have utilized a large latitudinal range (ie. 80 N to70 S) for analysis while this study will examine a narrow range, 22 of latitude, for comparison purposes. Understand ing the application of Rapoports rule may be essential for conservation a nd management (Fortes, 2004). The systematics of cephalopods are rapidly changing, as research has increased in the past 25 years. Phylogenetic relationships among families within the major groups remain uncertain (Vecchione, 2002). Species th at have been collected in the Caribbean allow the opportunity to further describe th e cephalopod assemblage as a whole as well as addressing the ecological importance of the cephalopods within the region. The goals of this study were: 1. To examine Rapoports Rule within the latitudinal range of 8 N to 30 N in the Broad Caribbean, looking for an increase in cephalopod diversity at lower latitudes follo wing Stevens (1989) original predictions for RR. 2. To compare the cephalopod species ri chness to that of other studies conducted (Smith et al., 2002). There are species hot spots reported for portions of the Broad Caribbean using a wide range of vertebrate and invertebrate species. The present study will address how the cephalopods fit into that picture. 3. To determine the species composition, distribution and abundance of cephalopod species within the Broad Caribbean region. The specimens examined come from a variety of preserved materials from various institutions within the region. The study will expand on Vosss work (1956, 1973, 1985) to include all cephalopod species and their importance in terms of abundan ce, distribution, and ecology.

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24 Study Area: The Broad Caribbean region includes the Ba hama Islands, Gulf of Mexico, and the Caribbean Sea. The range for the present study is between 8 -30 N and 60 95 W (Figure 2.2). Figure 2.2: Study Area 2.2 Materials and Methods The present study used prev iously collected specimens, both identified and unidentified, to determine th e distribution of all cephal opods in the Broad Caribbean region. The most reliable taxonomic informa tion comes from examination of specimens, in conjunction with, but not limited to, data co mpiled from sources in the literature. The most helpful taxonomic studies include exam ination of comparative material from a variety of locations, including specimens from the original type locality when possible (Vecchione and Collette, 2000). Based on cu rrent cephalopod dichotomous keys, type specimens, literature, and expert opinions, the specimens analyzed were identified to the

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25 species level. Once a specimen was identifie d, it was plotted on a di stribution map of the Broad Caribbean region using the ArcView 9.2 mapping program. The present study included 4190 specimen s that were collected from known locations, and 1000 specimens collected within less-specific regional locations within the Broad Caribbean. Twenty coastal regions were arbitrarily assigned for inclusion into the dataset (Appendix A). Many of the regional specimens were from the Caribbean Sea and were important to include. The regional data were used in the analysis of the distributions of cephalopods but not included for the a pplication of RR or species richness analysis. Identification guides (Voss 1956; Roper and Young, 1984; Nesis 1987; Vecchione, 2002) were used to identify each or ganism accurately to the species level. The majority of specimens examined had b een preserved well and were in excellent condition. The specimens dated from 1898 to pr esent. The bulk of preserved specimens analyzed were from two institutions, the Sm ithsonian Institutions National Museum of Natural History and the University of Mi ami Rosenstiel School of Marine and Atmospheric Science. Smaller collections fr om the Florida Fish and Wildlife Research Institute were also analyzed. The Hour glass cruises were co nducted from 1965-1967 in the shelf waters off west central Florida where cephalopod species were identified and documented by Dr. Ronald Toll and Dr. Stev en Hess. Approximately 500 specimens were included in the present study from those cruises. Identification of specimens was conducte d in the micronekton museum at the College of Marine Science at the University of South Flor ida, the Rosenstiel School for Marine and Atmospheric Science, Univers ity of Miami as well as the Smithsonian

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26 Institutions National Museum of Natural History. A dissect ing microscope was used to identify small specimens and specific features of the animals. Micrometer calipers were used for length measurements. The methods used for identifying and cat aloguing specimens were to identify each specimen using the cephalopod dichotomous keys and then to record its mantle length, sex, number of gill lame llae (octopods), and species name into an excel file for statistical analysis. Once all the organism s were identified, known specimens from all institutions were also added to the excel file for analysis. Distribution was calculated using the 9.2 ArcView software program. Statistical Analysis Rapoports Rule (RR) was evaluated by us ing species richness, which is defined here as species number per 5 latitudinal bin within the study s scope (5 bands). This was plotted as species richness ve rsus latitude (Fig. 2.3). Fi gure 2.4 represents the two variables addressed when comparing the 5 degr ee latitudinal binsth e number of species, as well as total number of individuals used for calculations in each bin. Rarefaction curves were created for all 5 latitudinal bands using Primer 6.2 for estimates and graphic output (F ig. 2.5, 2.6). Rarefaction is a tool used to correct for unbalanced sampling structure. The rarefac tion curve is produced by repeatedly resampling the pool of N individuals or N samp les, at random, plotting the average number of species represented by 1,2,.N individuals or samples (Gotelli & Colwell, 2001). It is dependent on the shape of the species abundance curve rather than the absolute number of specimens per sample. This method is valid when the same groups of organisms are being compared and contrasted. Another requi site is that all the habitats sampled be

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27 similar, in this case, coastal habitats. Methods of capture should be similar and lastly, this method does not specify which species take n from the residue will be present, and it can only be used to interpolate (Sanders, 1968). The rarefaction curve (Fig. 2.5) is species observed (Sobs) compared to latitude. A second set of curves (Fig. 2.6) is the Chao 1 estimator curve which gives a most likely total species es timate for each region based on the actual sample provided. Anne Chao devised a non-parametric estimator used for species richness comparisons. It is based on the number of rare species in a sample and creates an estimate of total spec ies for a region (Magurran, 2004). The Sobs graph accounts for sample-size differences while the Chao 1 estimator gives an absolute number for species richness in a region. Hotspots were calculated using an excel spreadsheet for comparison of species numbers in 8 subareas (Figs. 2.7, 2.8). The 8 sites were chosen by location in the Broad Caribbean which also correlates with a sp ecies richness study conducted by Smith et al. (2002). The 8 subarea coordinates are in Appe ndix B. Rarefaction curves for the sites were created using Primer 6.2 for comparison of the species observed and the Chao 1 estimator (Fig. 2.9, 2.10). 2.3 Results Rapoports Rule and Species Richness Species richness in the Broad Caribbean showed an increase with increasing latitude (Fig. 2.3). The 8-10 band showed the lowest species richness, 34 species, gradually increasing up through the highest latitude s to the 26-30 band with 77 species represented. Figure 2.4 compares the number of individual cephalopods examined to the number of species found in each latitudinal band. There was an increase in species found

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28 per band while the number of individuals varied among bands. Latitudinal band 11-16 N had a decrease in individuals examined while species richness was increasing, which indicates that sampling effort was not the sole factor for species richness increasing northward. The species observed (Fig. 2.5) rarefaction curve shows a trend for all latitudinal bands headed towards asymptote but only the higher latitudes close in on approaching it. The Chao 1 estimator curve (Fig. 2.6) represents the expected number of species found in each band. Cephalopod Biogeography Eight subareas (Fig. 2.7) were chosen to compare species richness within the Broad Caribbean. Each of the regions in corporated features of biogeographical significance; e.g. important curren t patterns or seafloor features or exhibited potential as human management areas. These subar eas correspond to a biogeography study of various marine organisms compared by Smith et al. (2002) analyzi ng invertebrates and vertebrates for potential dive rsity hotspots in the region. Figure 2.8 compares the 8 subareas and the number of species per region. It shows that subarea 4 (Eastern central Florid a) has the highest sp ecies richness (n=32), followed by subarea 1, northern Gulf of Mexi co (n=27), subarea 3 in the Straits of Florida (n=22), subarea 8 in the southwestern Caribbean Se a (n=20), subarea 2 in the West central Florida waters (n=13), subarea 5 with 11 different species, subarea 6 with 4 species, and lastly, subarea 7 with only 3 species types co llected. Rarefaction curves were used to determine the expected number of species per sample site as a function of organisms sampled. Table 2.2 is the number of samples and species analyzed for the bar graph analysis.

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29 Figures 2.9 and 2.10 show the rarefacti on curves derived for each region. Subregion 6 was not included due to the low num ber of samples (n=4). The trendline is similar with differences between subareas in close proximity of one another. Subarea 7 and 8 are both in the lower Caribbean Sea yet show a larg e difference in both species observed and expected species. Another ex ample of difference in species richness is between subareas 2 and 4. Subarea 2, the eas tern central Gulf of Mexico has a much lower number of species observed and expected number of species than subarea 4 of approximately the same latitude. Distribution and abundance of Broad Caribbean Cephalopods Distribution maps for each cephalopod sp ecies found within the Broad Caribbean are in Appendix C. Species maps are in orde r by family groups in most cases. All 5190 specimens were located within the distribut ional effort map shown in Figure 2.11. The figure represents all of the known coordinate sampling sites for specimens as well as the 20 regions created to define areas throughout the Broad Caribbean, including the 1000 specimens that were collected with a regiona l location but lacked precise latitude and longitude coordinates (Appendix A). Cephalopod Range Extensions Range extensions for several species of cephalopods in the Broad Caribbean region were observed based on the dataset collected in the present study. Table 2.3 shows 18 extensions to previously know n ranges based on the work of Roper and Sweeney, 1984; Nesis, 1987; and Vecchione, 2002.

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30 Species Richness by Latitude 34 53 54 76 770 10 20 30 40 50 60 70 80 90 8-1011-1516-20 21-2526-305 degree latitudinal binsTotal # of species Figure 2.3: Species richness for all cephalopod species per 5 latitudinal bin Figure 2.4: Comparison of species richness and number of individuals RR total species0 10 20 30 40 50 60 70 80 90 8-1011-1516-20 21-2526-30Latitude (deg N)# species RR# individuals0 500 1000 1500 2000 2500 8-1011-1516-20 21-2526-30# individuals

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31 Species Observed (RR Analysis)0 10 20 30 40 50 60 70 80 115294357718599113127141155169183197211225239253267281295309323337351Number of SamplesNumber of Species Sobs 26-30 Sobs 21-25 Sobs 16-20 Sobs 11-15 Sobs 8-10 Figure 2.5: RR rarefaction curv es for species observed RR Chao 1 estimates0 10 20 30 40 50 60 70 80 90 100 115294357718599113127141155169183197211225239253267281295309323337351Number of samplesNumber of Species Chao1 26-30 Chao1 21-25 Chao1 16-20 Chao1 11-15 Chao1 8-10 Figure 2.6: RR samples; Chao 1 estimator curve

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32 Figure 2.7: Eight subareas for richness and diversity comparison; Subarea 1= Northern Gulf of Mexico; Subarea 2= West central Flor ida; Subarea 3= Straits of Florida; Subarea 4= East central Florida; Subarea 5= Mid-Isl and Caribbean group; Subarea 6= Southeast Caribbean Sea; Subarea 7= South central Cari bbean Sea; Subarea 8= Southwest Caribbean SeaColombia area Figure 2.8: Species Richness Regional Compar ison; Subarea 1= Northern Gulf of Mexico; Subarea 2= West central Florida; Subarea 3= Straits of Florida; Subarea 4= East central Florida; Subarea 5= Mid-Island Caribbean gr oup; Subarea 6= Southeast Caribbean Sea; Subarea 7= South central Caribbean Sea; Subarea 8= Southwest Caribbean SeaColombia area Species Richness Comparison0 5 10 15 20 25 30 35 1 2 3 4 5 6 7 8Broad Caribbean Regions Species Number 1 2 3 4 5 6 7 8

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33 Table 2.2: Region, Species # and Sample # for hotspot comparison Location Region Species # # samples North GOM 1 27 46 West Central Florida 2 13 151 Straits of Florida 3 22 32 Eastern Central Florida 4 32 68 Mid-Island Group 5 11 17 Southeast Caribbean 6 4 4 South Central Caribbean 7 3 10 Southwestern Caribbean 8 20 25 Hotspot Species ObservedRarefaction Curve0 5 10 15 20 25 123456789101112131415161718192021222324252627282930Sample numberCumulative # species Region1 Region2 Region3 Region4 Region5 Region7 Region8 Figure 2.9: rarefaction curve species ob served comparison among regions 1-8

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34 Chao 1 Estimate Curve0 5 10 15 20 25 30 123456789101112131415161718192021222324252627282930Sample numberCumulative number of species Region 1 Region 2 Region 3 Region 4 Region 5 Region 7 Region 8 Figure 2.10: Chao 1 estimated number of species for regions 1-8 Figure 2.11: Sample Effort Map of Study; x= sample site

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35 Table 2.3. Cephalopod distribution and range extensions to the Broad Caribbean Gulf of Mexico Caribbean Sea Atlantic Ocean Range Extension Architeuthidae Architeuthis sp X X X Bathyteuthidae Bathyteuthis abyssicola X X X X Bathyteuthis sp. X X X Chiroteuthidae Chiroteuthis sp. Asperoteuthis acanthoderma X X X Grimalditeuthis bonplandi X Planctoteuthis danae X Cycloteuthidae Discoteuthis discus X Cranchiidae Bathothauma lyromma X X Cranchia scabra X X X Egea inermis X Helicocranchia pfefferi X X Leachia atlantica X Leachia lemur X X Leachia spp. X X Liocranchia reinhardti X X Megalocranchia abyssicola X Megalocranchia sp. X X Sandalops melancholicus X Taonius pavo X X Teuthowenia megalops X Enoploteuthidae Abralia redfieldi X X Abralia veranyi X X X Abraliopsis atlantica X Abraliopsis pfefferi X Enoploteuthis leptura X X Enoploeuthis anapsis X X Histioteuthidae Histioteuthis corona X X X Histioteuthis dofleini X X Histioteuthis reversa X X Histioteuthis sp. X X Stigmatoteuthis arctura X X Joubiniteuthidae Joubiniteuthis portieri X

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36Loliginidae Doryteuthis brasilensis X Doryteuthis ocula X Doryteuthis pealeii X X X X Doryteuthis plei X X X X Doryteuthis roperi X X X X Doryteuthis sp. X X X Doryteuthis surinamensis X Lolliguncula brevis X X X X Pickfordiateuthis pulchella X X X Sepioteuthis sepioidea X Lycoteuthidae Lycoteuthis diadema X X Lycoteuthis sp. X Lycoteuthis springeri X Selenoteuthis scintillens X X X X Mastigoteuthidae Mastigoteuthis agassizi X X Mastigoteuthis hjorti X X Octopoteuthidae Octopoteuthis sp. X Taningia danae X Ommastrephidae Illex coindetii X X X X Illex illecebrosus X X X Illex oxygonius X Hyaloteuthis pelagica X Ommastrephes bartrami X X X Ornithoteuthes antillarum X X X Onychoteuthidae Ancistroteuthis lichensteinii X Onychoteuthis banksii X X X Onykia sp. X Moroteuthis aequatorialis X Pholidoteuthidae Pholidoteuthis adami X X X Pyroteuthidae Pterygioteuthis gemmata X X Pterygioteuthis giardi X Pterygioteuthis sp. X X X Pyroteuthis margaritifera X X X Sepiolidae Heteroteuthis dispar X X Nectoteuthis pourtalesi X Austrorossia antillensis X

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37Rossia bullisi X X X X Rossia tortugaensis X X Neorossia sp X X Semirossia equalis X X X Semirossia tenera X X X X Stoloteuthis leucoptera X Spirulidae Spirula spirula X X X Thysanoteuthidae Thysanoteuthis rhombus X X Alloposidae Haliphron atlanticus X Argonautidae Argonauta argo X X Argonauta sp. X Bolitaenidae Japetella diaphana X X Bolitana pygmaea X Octopodidae Bathypolypus bairdii X X X Benthoctopus januari X X Benthoctopus oregonae X X Danoctopus schmidti X X Euxaoctopus pillsburyae X Ocellate Octopods X X Octopus briareus X X X X Amphioctopus burryi X X X X Octopus carolinensis X X Macrotritopus defilippi X X X Octopus filosus (hummelincki) X X Octopus joubini X X X Callistoctopus macropus X X Octopus maya X X Octopus occidentalis X X Octopus americanus X X X Octopus zonatus X Pteroctopus tetracirrhus X X X Scaeurgus unicirrhus X X X X Tetracheledone spinicirrus X X Tremoctopodidae Tremoctopus gelatus X Tremoctopus violaceus X X X Opisthoteuthidae Grimpoteuthis megaptera X X Grimpoteuthis sp. X X

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38Opisthoteuthis agassizii X X X Vampyroteuthidae Vampyroteuthis infernalis X X X New Species to Area Two Asperoteuthis ancanthoderma specimens were examined demonstrating a very significant range extension to the speci es; the only other speci mens found previously were off the coasts of Japan. Two nearly in tact individuals were found floating dead on the surface of the ocean and collected: one specimen was found off the coast of Key West, FL and the other off the coast of Mara thon, FL. Both specimens were dissected and analyzed for identification. It was determined that they were both Asperoteuthis acanthoderma A description paper is in press w ith the Proceedings of the Biological Society of Washington (Judkins et al. 2009). 2.4 Discussion Rapoports Rule and Species Richness Rapoports Rule attributes the many observati ons of increasing diversity with decreasing latitude to a reduction in size of species distributional ra nges as you approach the equator (Rosa et al., 2008). Stevens ( 1989) supported his claim with studies of diverse taxa including North American tr ees, North American marine molluscs, freshwater and coastal fishes, reptiles and amphibians. Since that time, many scientists have studied the ecological patterns driving biological diversity. There have been numer ous hypotheses to explai n diversity patterns (Peet, 1974; Evans, 2005) and various groups of organisms have been examined to prove or disprove RR. Many studies of mari ne groups have supported RR (Steele, 1988;

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39 Stevens, 1996; Roy et. al. 1998, 2000; Rex et al. 2000, 2005; Macpherson 2002) while others have failed to find such a relati onship (Clarke, 1992; Lambshead et al. 2000; Mokievsky and Azovsky, 2002; Rosa et al., 200 8). The latter studies found a negative correlation to RR as discussed below. There have been a few molluscan species richness studies in the Atlantic Ocean to date (Rosa et al., 2008, Fortes and Absa lao, 2004, Macpherson, 2003). Fortes and Absalao (2004) examined gastropods and biva lves using literature -based studies from both the Pacific and Atlantic sides of the c ontinental North and Sout h American coasts. After analyzing 4067 species they determined that RR applied to these organisms on both coasts. They noted that regional features, su ch as the size of a bi ogeographical province, seemed to affect the pattern strongly. They also found support for RR when they incorporated depth into the study. Macpherson (2003) studied the variability in size of species ranges in terms of depth and latitude for various marine taxa, including cephalopods and fishes in the Atlantic Ocean. The results showed that RR could hold true for those organisms but was not solely responsible for latitu dinal patterns in range sizes. The research conducted by Rosa et al. ( 2008) examined cephalopod species of the coastal Atlantic Ocean using primary literat ure, reports, and online databases. Their results showed that latitudinal gradients of species richness were present along both Atlantic coasts, but were distinct from one a nother. When the median latitudinal ranges of the Western Atlantic neritic cephalopods were determined, it was evident that the size of the distributional ranges did not decline with decreasing latitude which means that RR may not explain the distributi on patterns. Stevens, (1996) proposed that RR could extend

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40 to elevation and water depth in terms of sp ecies richness. When species depths were taken into consideration for the organisms in the western Atlantic, RR was exhibited (Rosa et al. 2008). The present specimen-based study showed that the cephalopods of the Broad Caribbean do not exhibit the diversity pattern s described originally by Stevens (1989). Within the small latitudinal range of 8 to 30 N, cephalopods of the region increase in species richness with increasing latitude. The lack of concor dance with RR in the present study agrees with Rosa et al. (2008). It should be noted that the lowest latitude band, (8 10 ) includes only 3 latitude while the other 4 include 5 in each band. Therefore, there may be more than the 34 species in the regi on that the present study suggests. However, the species richness trend is still obvious as latitude increases. One of the reasons for an increase in speci es number at higher latitudes could be the convergence of the Florida current and the North Equatorial current in the middle to northern end of the study area. The two cu rrents converge to become the Gulf Stream and may be transporting cephalopods northwar d. The Florida Current becomes the Gulf Stream and then leaves the coast of the eastern United States at Cape Hatteras to head across the Atlantic Ocean. Th is current has a profound influence on the distribution of shore animals in the wester n Atlantic (Briggs, 1995). Another factor possibly cont ributing to the northward in crease in species richness is the larger number of studies in the northern Broad Caribbean region. Numerous studies have been conducted in Florida waters which may be contributing to the increased richness of cephalopods in the northern por tion of the study area. The RR rarefaction curve (Fig. 2.5) showed a similar richness tre nd for all regions and pointed out the need

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41 for more sampling in the lower latitudes, as the upper 2 bands were close to asymptote while the lower 3 bands were still rising. The Chao 1 estimator analysis showed differences among species richness between the bands. Note that the 8 10 latitude band approached 60 species expected and then decreased rapidly (F ig. 2.6). A possible reason for the decline is that there was a smaller sample size for that latitudinal band in conjunction with numerous singleton species counted, which caused a decrease in the species richness curve for that region. Cephalopod Biogeography Based on range map overlays, Smith et al. (2002) examined the distribution of 1172 vertebrate and invertebrate species and concluded that the area of highest species richness was located in the waters surroundi ng southern Florida, the eastern Bahamas, and northern Cuba. Secondary centers of dive rsity were located (in descending order of richness) on the continental sh elves of northern South America, Central America, and in the northern Gulf of Mexico. Those patterns are apparently r obust as they are repeated in composite distributions for fishes and for othe r invertebrates taken separately (Smith et al., 2002). Analyzing the present studys cephalopod species richness information, eastern central Florida has the highest species ric hness in the Broad Cari bbean (n=32), likely because the Gulf Stream acts as a large transporter of larvae from the southern waters feeding into it. Another possibl e, but less likely, factor for this subareas species richness is that it is a well-traveled path for seasonally migrating cephalopods. The cephalopods of the Broad Caribbean generally follow the pattern found by Smith et al.(2002) in that the two regions exhi biting the highest richne ss were the same in

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42 both studies. However, subarea 6, the south eastern Caribbean Sea, had too few samples to include in the analysis (n=4). Subarea 7, south central Caribbean Sea, had 10 samples and was included in the analysis. Smith et al suggested that the southern edge of South America was the second riches t in terms of species wher eas the present study does not show that trend. The two areas of low sampling effort (6 & 7) over 111 years of collecting indicate the need for furt her fieldwork in those regions. The subarea rarefaction curve displays curi ous trends within the Broad Caribbean. It was anticipated that there would be vari ations in species richness between the major basins of the Caribbean and the Gulf of Mexic o. This does not appear to be the case. For example, subareas 7 and 8 were both located in the lower Caribbean and yet showed significantly different trends in species richness (Fig. 2.9). Another example of a variation within regions can be seen between su bareas 2 and 4 in the northern sections of the Broad Caribbean. A reason for the variati on in this case could be the intense work that was conducted by the Hourglass cruise s in the mid 1960s by the Marine Research Laboratory of the Florida Board of Conserva tion, creating an artifact from sampling efforts. Two experts identified and cata logued cephalopod shelf species which were included in the present study. Although over 500 specimens were included in subarea 2, the species richness of the suba rea (n=13) was still lower th an subareas 3 (n=22) and 4 (n=32). Another explanation for the increas e in species richness in subarea 4 could be the influence of the water depth changes a nd flow of the Gulf Stream. These two examples of variation within regions (Gulf of Mexico and Caribbean basin) exceed the variations between the northern and s outhern regions of the studys scope.

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43 Distribution and abundance There were 110 cephalopod species from 27 families examined in the present study. This number is larger than that found in earlier studies (Voss, 195642), but quite similar to more recent work (Nesis 1975-100), (Vecchione, 2002-109). This study examined unidentified and identified specime ns that were primarily from two large institutions. Broad Caribbean cephalopods are widely distributed, many finding niches within coastal and island ecosystems. The abundance of organisms differs between species, with Doryteuthis plei (n=1205) and Doryteuthis pealeii (n=702) being the most numerous squid. Doryteuthis plei and D. pealeii move in large schools, are commercially harvested, used in medical studies and are distributed throughout the region. Octopus joubini (n=351) and Octopus americanus (vulgaris) (n=306) were the mo st abundant octopods and are also commercially harvested thr oughout the region. Over 30 species were represented by 3 or fewer specimens for the en tire region. An example of a lesser known cephalopod is Discoteuthis discus found in deep water with little known about its distribution or biol ogy (Vecchione, 2002). Euaxoctopus pillsburyae inhabit small niches surrounding a particular island group and only 1 specimen was recorded for this study. Few deep-sea cephalopods have been collecte d, indicating the need for increased research efforts in the area to uncover other unique organisms below 500m with regularity. Few studies of cephalopod distribution have been reported in context for the Broad Caribbean area. Two studies that focused on the region were completed by Barrientos and Garcia-Cubas (1997) and Vecchio ne et al. (2001). Barrientos focused on three loliginids, Doryteuthis ( Loligo ) plei, Doryteuthis ( Loligo ) pealeii and Lolliguncula

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44 brevis in the western Gulf of Mexico. The authors associated squid abundance and distribution with currents, warm layers, fronts, water masses, and climate changes (Barrientos and Garcia-Cubas, 1997). These re sults are concordant with the current findings as D. plei were most abundant (n=1205) followed by D. pealeii (n=702) with L. brevis (n=374) rounding out the top three most abundant cephalopods in this study. Another study (Vecchione et al., 2001) centered on paralarval cephalopod distribution and abundance in th e western North Atlantic Oc ean. The two most abundant and frequently collected speci es were the neritic squids Doryteuthis pealeii and Illex illecebrosus collected north of Cape Hatteras, both valuable fishery resources. The highest abundance and diversity of planktoni c cephalopods in the oceanic samples were consistently found in the vicinity of the Gulf Stream. According to Vecchione (2001) the most likely species other than D. pealeii to be present in the south is D. plei (Vecchione, 1981). Other Doryteuthis species such as D. ocula are restricted to Caribbean islands whereas Doryteuthis roperi once considered a Caribbean species, had a range extension into the Gulf of Mexico and Atlantic Ocean, demonstrated in the present study. Doryteuthis pealeii was abundant in the Broad Cari bbean in the present study as well as in Vecchiones work while Illex illecebrosus was only present in small numbers (n=6). Another Illex species, I. coindetii was more abundant in the Gulf of Mexico and the northern Caribbean (n=82) which sugge sts that it is bette r suited for tropical temperatures than Illex illecebrosus Findings here differ from Vecchiones(2001) work in that D. roperi were abundant (n=89) and found not only in the Caribbean but were distributed through the Straits of Florida a nd up the Gulf Stream to the northernmost region of the study. D. ocula is confined to the Caribbean as the 3 specimens revealed in

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45 this study. It should be noted that there se em to be fewer species found in the western regions of both the Gulf of Mexico and the Ca ribbean Sea. This could be attributed to low sampling effort (Fig. 2.11). Range Extensions The observed range extensions are extrem ely valuable. There have been recent studies conducted (Rosa et al ., 2008; Judkins et. al 2008) that compare organisms of the region based on literature s ources so to find 18 new extensions based on fresh identification is helpful in st udying the biology and habits of th e organisms. It is also important as the hunt for new fisher y resources increases worldwide. New Species to the Area It is not surprising to find a new speci es in the Broad Caribbean as research funding has decreased and cephalopod studies ha ve been limited in past years. The species described, Asperoteuthis acanthoderma was found prior to this only in the West Pacific Ocean. More studies targeting deep water species are needed to determine the role these large and important species play in the oceans f ood webs as well as learning their biology and habits to bett er understand cephalopod adaptations. To summarize, the present study found that 110 cephalopod species of the Broad Caribbean are dispersed thr oughout its waters, they occupy a myriad of niches along coastlines and deep water, and there is no support for RR. Hotspots are patchy with the most species richness occurring along the eastern edge of Fl orida in the Gulf Stream. The range extensions based on data important to note for future conservation and fishery ventures. Asperoteuthis acanthoderma is a fortunate discovery and begs the question

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46 what else is down there? Areas of the s outheastern Caribbean as well as the western Gulf of Mexico appear to be understudied. The present study is the firs t to examine the specimen-based species richness area with rarefaction curves to support it, atte mpting to determine species richness of the Broad Caribbean region. The study size range is unique in that it is not encompassing numerous latitudes but focusing on 22 of a tropical/subtropical area important to a variety of species. Studies surrounding all aspects of cephalopod lifediversity, biology, ecology, and capture methods would improve the world database for these important organisms.

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47 REFERENCES 1. Barrientos, Gerardo, Antonio Garcia-Cuba s. 1997. Disstribution and abundance of the Family Loliginidae (Mollusca: Cephalopod a) in Mexican waters of the Gulf of Mexico. Rev. Soc. Mex. Hist. Nat.; 47: 123-139. 2. Briggs, John C. 1995. Global Biogeography ; Elsevier Science B.V.; 454 pp. 3. Cairns, S. 1976. Cephalopods collected in the Straits of Florida by the R/V Gerda. Bulletin of Marine Science; 26(2): 233-272. 4. Clarke, A. 1992. Is there a latitudinal diversity cline in the sea? Trends in Ecology and Evolution; 7: 286-287. 5. Clarke, M.R.1996. The Role of Cephalopods in the worlds oceans: general conclusions and the future. Phil. Trans. R. Society London. B.; 351: 1105-1112. ------1996. The role of cephalopods in the worlds oceans: an introduction. Phil. Trans. R. Society London. B.; 351: 979-983. 6. Diaz, J.M. 2004. First Record of an oct opus species if the family Tremoctopodidae (Cephalopda: Octopoda) for the Colombian Caribbean. Biol. Invest. Mar. Cost. 33; 283-286. 7. Diaz, J.M. and Gracia, N. 2000. Squids and octopuses (Mollusca: Cephalopoda) of the Colombian Sea. Biota Colombiana; 1(2) 195-201. 8. Fortes, R.R., R.S. Absalao, 2004. The applicability of Rapoports rule to the marine molluscs of the Americas. Journal of Biogeography; 31: 1909-1916. 9. Gotelli, Nicholas, Robert K. Colwell. 2004. Quantifying biodiversity: procedures, and pitfalls in the measurement and comparison of species richness Ecology Letters; 4: 379-391. 10. Gracia, A. 2002. Cephalopods (Mollusca: Cephalopoda) of the Upper Colombian Caribbean Shelf. Biol. Invest. Mar. Cos. 31, 219-238. 11. Jereb, P. and Roper, C.F.E. 2005. Cepha lopods of the world. An annotated and illustrated catalogue of cephalopod species known to date. FAO species catalogue for fisheries purposes, N4, v.1, FAO, Rome. 12. Judkins, H, M. Veccione, & C.F.E. Roper. In press. Cephalopods (Mollusca: Cephalopoda) of the Gulf of Mexico. Gulf of Mexico Origin, Waters, and Biota Biodiversity, Volume 1 (Darryl L. Feld er and David A. Camp, editors, Texas A&M University Press, College Station, Texas).

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48 13. Judkins, H. Debra A. Ingrao, C.F.E. Roper. In press. First records of Asperoteuthis acanthoderma (Lu, 1977) (CEPHALOPODA: OEGOPSIDAE: CHIROTEUTHIDAE), from the North Atla ntic Ocean, Straits of Florida. Proceedings of the Biologi cal Society of Washington. 14. Lambshead, P.J.D., J. Tietjen, T. Ferraro, & P. Jensen. 2000. Latitudinal diversity gradients in the deep sea with special refernce to North Atlantic nematodes. Marine Ecology Progress Series; 194: 159-167. 15. LaRoe, E.T. 1967. A contribution to th e biology of the loliginidae (Cephalopoda: Myopsida) of the Tropical Western Atlantic. Master of Science thesis; University of Miami, Coral Gables, Fl. 16. Lipka, D.A. 1975. The systematics and zoog eography of cephalopods of the Gulf of Mexico. PhD. Dissertation, Texas A&M University; 204 p. 17. Longhurst, A.1998. Ecological geography of the sea. Academic Press; 146-150. 18. Macpherson, E. 2002. Large-scale species-ric hness gradients in the Atlantic Ocean. Proceedings of the Royal Society of London; B; 269: 1715-1720. -----2003. Species range size distributions for some marine taxa in the Atlantic Ocean. Effect of latitude and depth. Biological Journal of the Linnean Society; November; 80(3): 437-455. 19. Magurran, Anne E. 2004. Meas uring Biological Diversity Blackwell Publishing. 256p. 20. Mokievsky, V. and Azovsky, A.2002. Re-evalu ation of species diversity patterns of free living marine nematodes. Mari ne Ecology Progress Series; 238: 101-108. 21. Nesis K.N. 1975. Cephalopods of the American Mediterranean Sea EnglishTranslations of Selected P ublications on Cepha lopods; Editor M. Sweeney; Smithsonian Institutions Libraries; 1; 318358. -----1987. Cephalopods of the world. (Edited by L. Burgess) T.F.H. Publication, Inc. Ltd. New Jersey; 351 pp. 22. Passarella, K.C., 1990. Oceanic Cephalopod Assemblage in the Eastern Gulf of Mexico Department of Marine Science University of South Florida: St. Petersburg, Fl. 50pp. 23. Peet, R.K. 1974. The Measurement of Species Diversity. Annual Review of Ecological Systems; 5: 285307. 24. Rapoport, E.H. 1982. Areography:geogr aphical strategies of species; 1st English ed. B. Drausal, transl. Publ. Fundacion Bariloche. V.1; Pergamon, New York.

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49 25. Rex, M.A., J.A. Crame, C.T. Stuart, & G. Coyne. 2000. Latitudinal gradients of species richness in the deep-sea benthos of the North Atlantic; Proceedings of the National Academy of Sciences; USA; 97: 4082-4085. 26. Rex, M.A., J.A. Crame, C.T. Stuart, & A. Clarke. 2005. Larg e-scale biogeographic patterns in marine molluscs: a confluen ce of history and pr oductivity? Ecology; 86: 2288-2297. 27. Rohde, K.1992. Latitudinal gradients in spec ies diversity: the search for the primary cause. Oikos; 65: 514-527. 28. Roper, C.F.E., 1983. An overview of cephalopod system atics: status, problems, and recommendations. Memoirs of the National Museum Victoria.; 44: 13-27. 29. Roper C.F.E, M.J. Sweeney & C.E. Nauen, 1984. FAO species catalogue. v.3 Cephalopods of the world. An annotated a nd illustrated catalogue of species of interest to fisheries. FAO Fish. Synop. (125) v3: 277p. 30. Rosa, Rui, Heidi M. Dierssen, Liliana G onzalez, and Brad Seibel. 2008. Ecological biogeography of cephalopod molluscs in th e Atlantic Ocean: historical and contemporary causes of coastal dive rsity patterns; Global Ecology and Biogeography; 17: 600-610. 31. Roy, Kaustuv, David Jablonski, James W. Valentine, and Gary Rosenberg. 1998. Marine latitudinal diversity gradients: Test s of causal hypotheses. Proceedings of the National Academy of Science. USA. 95; March; 3699-3702. 32. Roy, K., D. Jablonski, J.W. Valentine. 2000. Dissecting latitudinal diversity gradients: functional groups and clades of marine bivalves; Proceeding of the Royal Society of London; B267: 293-299. 33. Sanders, H.L. 1968. Marine benthic di versity: a comparative study. American Naturalist. 102: 243-282. 34. Smith, M.L.,K.E. Carpenter, & R. W Waller; 2002. An Introduction to the Oceangraphy, Geology, Biogeography, and Fisheries of th e Tropical and Subtropical Western Central Atlantic. Carpenter, K.E.(ed.) The living marine resources of the Western Central Atla ntic. Volume 1: Introduction, molluscs, crustaceans, hagfishes, sharks, batoid fishes, and chimaeras. FAO Species Identification Guide for Fisheries Pu rposes and American Society of Ichthyologists and Herpetologists Speci al Publication No. 5. Rome, FAO. 2002. pp.1-600. 35. Stanley, Sonja (editor), 1995. A Global Re presentative System of Marine Protected Areas: Vol 2, Marine Region 7: Wider Cari bbean, Great Barrier Reef Marine Park Authority, The World Bank, The Worl d Conservation Union,(IUCN); 15-42.

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50 36. Steele, D.H.1988. Latitudinal variations in body size and species diversity in marine decapod crustaceans of the continental shelf. International Review of Hydrobiology; 73: 235-246. 37. Stevens, G.C., 1989. The Latitudinal Gradient in Geographical Range: How So Many Species Coexist in the Tropics. The American Naturalist; 133(2): 240-256. 38. Stevens, G.C. 1996. Exte nding Rapoports rule to the Pacifi c marine fishes. Journal of Biogeography; 23: 149-154. 39. Summers, W.C., 1983. Physiological and Trophic Ecology of Cephalopods The Mollusca. Ecology; 6: 261-279. 40. Sweeney, M.J., Clyde F.E. Roper, 1998. Classification, Type Lo calities, and Type Repositories of Recent Cephalopoda in Systematics and biogeography of Cephalopods M.V. Nancy A. Voss, Ronald B. Toll, and Michael J. Sweeney, Editor. Smithsonian Institution Press: Washington, D.C. 562-599. 41. Tomczak, Mattias, J. Stuart Go dfrey. 1994. Regional Oceanogrpahy: An Introduction. Pergamon; 310-317. 42. Vecchione, M. 1981. Aspects of the early life history of Loligo pealeii (Cephalopoda: Myopsida). Journal of Shellfish Resources; 1:171-180. 43. Vecchione, M., Bruce B. Collette, 1996. Fisheries Agencies and Biodiversity. Ann. Missouri Bot. Gard; 83: 29-36. 44. Vecchione, M., and B.B. Collette 1996b. The cen tral role of systematics in marine biodiversity issues. Oceanography; 9(1): 44. 45. Vecchione, M., M.F. Mickevich, K. Fauchald, B.B. Collette, A.B. Williams, T.A. Munroe, and R.E. Young, 2000. Importance of Assessing Taxonomic Adequacy in Determining Fishing Effects on Marine Diversity. ICES Journal of Marine Science; 57: 677-681. 46. Vecchione, M., C.F.E. Roper, Michael J. Sweeney, and C.C. Lu. 2001. Distribution, relative abundance, and developmental morphology of paralarval cephalopods in the western North Atlantic Ocean. U.S. Department of Commerce, NOAA Tech. Rep. NMFS 152, 54p. 47. Vecchione, 2002. Cephalopods. Carpenter, K.E.(ed.) The living marine resources of the Western Central Atla ntic. Volume 1: Introducti on, molluscs, crustaceans, hagfishes, sharks, batoid fishes, and chimaeras. Cephalopods of Western Central Atlantic; FAO Species Identification Guide fo r Fisheries Purposes and American Society of Ichthyologists and Herpetologi sts Special Publication No. 5. Rome, FAO. 2002. 1-600.

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51 48. Voss, G.L. 1956. A review of the cephal opods of the Gulf of Mexico. Bull Mar. Sci. Gulf and Caribbean; 6:85-178. ------1973. The potentially commercial species of octopus and squid of Florida, the Gulf of Mexico and the Caribbean Sea. Un iversity of Miami Sea Grant Program (NOAA Sea Grant No. 04-3-158-27) Miami, Fl. 49. Voss, G., Thomas F. Brakoniecki, 1985. Squid resources of the Gu lf of Mexico and Southeast Atlantic coasts of the United States. Science Council Studies; 9: 27-37. 50. Voss, N.A., Vecchione, M., Toll, R.B. & Sweeney, M.J.(eds) 1998. Systematics and biogeography of cephalopods. Smithson, Contrib. Zool.; 586: 1-599.

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52 CHAPTER 3 CEPHALOPODS (MOLLUSCA: CEPHALOP ODA) OF THE GULF OF MEXICO Heather L. Judkins, Michael Vecchione, and Clyde F.E. Roper (HJ) College of Marine Science, University of South Florida, St Petersburg, Fl, 33701; USA (email: Hjudkins@marine.usf.edu) (MV) NMFS National Systematics Laborator y, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012 USA (email: VecchioneM@si.edu) (CR) Invertebrate Zoology, MRC 118, National Museum of Natural History, Smithsonian Institution; Wash ington DC 20013-7012 USA (email: gsquidinc@verizon.net) 3.1 Introduction The cephalopods of the Gulf of Mexico have not been studied comprehensively since Voss (1956) monograph. Several cephalopod studies have included this region in broader studies (e.g., Vecchione 2002) or ha ve examined distribution based on limited geographic area (e.g. Nesi s 1975, Passarella and Hopki ns 1991). Collectively, approximately 109 species of cephalopods in 31 families occur in the Western Central Atlantic and adjacent areas, including the Ca ribbean Sea and the Gulf of Mexico. Both squids and octopods are common in these wa ters (Vecchione 2002). The present paper updates Voss works and summarizes the specie s found in the Gulf of Mexico to date. Two major groups of cephalopods exist toda y: the Nautiloidea, with a few species of the pearly nautilus confined to the IndoWest Pacific, and the Neocoleoidea, consisting

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53 of squids, cuttlefishes, octopods and their relatives. Neoc oleoidea comprise more than 700 species worldwide: (http://tolweb.org/tree?group= Cephalopoda&contgroup=Mollusca) The neocoleoids that exist today evolved from forms that first appeared in the upper Triassic and Lower Jurassic (between 200 and 150 million years ago). Although there are relatively few species of living cephalopods, they occupy a great variety of habitats in all of the worlds oceans. Individual species may be very abundant and are important in marine food webs. Some species are major targets for commercial fisheries. The Neocoleoidea contains two exta nt Superorders: the Octopodiformes (octopods and vampire squid) and the Decapodi formes (squid and cuttlefishes). These groups occupy all major habitats in the ocean s from intertidal to great depths (deepest record is 7279 m, Aldred et al. 1983, Voss 1988) and from southern to northern polar seas. No cephalopods are found in salinities less than about 17.5 pr actical salinity units (psu). Many species of oceanic cephalopods under go diel vertical migrations, wherein they occur at depths of about 400 m to 1000 m during the day, th en ascend to the uppermost 200 m or so during the night (e .g., Enoploteuthidae, Ommastrephidae). Abundance patterns vary (depending on group, habitat, and season) from isolated territorial individuals (primarily octopods and sepioids), through small schools with a few dozen individuals, to massive schools with millions of oceanic squids. Although many cephalopods reach large sizes, generally they have a very short life span. The life expectancy appears to be about 1 years in most cephalopods, but large species of squids and octopods and thos e in cold water habita ts may live longer.

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54 Conversely, some small oceanic squids such as pyroteuthids may complete their life cycles in less than six months. This is part of a life history strategy that seems designed for rapid increase in population size. It has been suggested that this lif e-strategy may guarantee survival against environmentally st ressful conditions, including those by heavy predation or over-fishing. However, as cephalopod fisheries experienced further extensive development, parallel concern de veloped regarding poten tial overexploitation (Jereb and Roper, 2005). Cephalopods are important in terms of food web relationships, commercial fisheries, and biomedical re search. Cephalopods are born in to the third trophic level and progress one or two stages th rough their life. Research ha s not shown them as achieving top-predator status because there always seems to be some vertebrate that preys upon them (Summers 1983). Cephalopod s are active predators that feed upon shrimps, crabs, fishes, and other cephalopods, and, in the case of octopods, on other molluscs. In turn, cephalopods are major food items in the diets of toothed whales, seal s, pelagic birds and both benthic and pelagic fishes as well as other cephalopods. Cephalopod fisheries provide an importa nt food source for humans across the globe. Over three million metric tons are ca ught each year worldwide (Jereb and Roper, 2005). Squid fisheries also have existed in North America, historically principally to provide bait for other fisheries, but recen t decades have seen the development of substantial fisheries for food production, as well. The total commercial catch of cephalopods in the Western Central Atlant ic varied during 1993 to 1998 between 19,000 tons and 31,000 tons, mostly landed in Me xico (Vecchione 2002). However, Voss reported in 1973 that of the numerous species kn own on the coasts of Florida, the Gulf of

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55 Mexico and the Caribbean Sea, only about 12 species seem to have actual or potential (fisheries) importance (Voss et al. 1973; p.1). Of those, sp ecies that are commercially important to the Gulf of Mexico include: Octopus maya Illex coindetii, Doryteuthis (Loligo) pealei and Doryteuthis (Loligo) plei (Voss et al. 1985). Squids also are important to biomedical research; for instance, much of what is known about human neurophysiology results fr om experiments with the giant axon of squids. Scientists culture squid in laborator ies in order to study the giant axon. Lee et. al. (1994) cultured the In do-West Pacific species Sepioteuthis lessoniana (Ferussac, 1830) for this purpose because of its large hatch lings, and the quality of its large-diameter axons for study (Lee 1994). LaRoe (1971) pr eviously had worked with a Caribbean species, S. sepioidea, for this purpose. Because of the highly developed brain and sensory organs, cephalopods are valuable in behavioral and compar ative neuroanatomical studies as well. The fauna of the Gulf of Mexico lacks th e nautiloids and true cuttlefishes but does include sepiolids, myopsid and oegopsid squids, octopods, and the vampyromorph, Vampyroteuthis infernalis Published records of cepha lopod species in the Gulf of Mexico date back to LeSueur (1821), but the modern, comprehensive systematic studies begin with G.L. Voss, who reported 42 neri tic and oceanic species in 1956(G. Voss 1956). Since then, many oceanic species have been added to the list (N. Voss and G. Voss 1962, Roper 1964, Voss 1964, Roper et al 1969, Lipka 1975, Passarella 1990). Although the composition of the cephalopod fauna off southern Florida is known almost exhaustively, the fauna of the rest of the Gulf of Mexico is less well studied. The cephalopods of the Mexican waters of the Gu lf were reviewed by Salcedo-Vargas (1991)

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56 using specimens and past lite rature, he reported some ques tionable identifications. The most recent compilation of the cephalopods in th e Gulf of Mexico is that of Vecchione (2002). 3.2 Major systematic revisions since 1954 The status of the systematics of cephal opods is rapidly changing, as research has increased substantially world-wide, during th e past 30 years. The families of living cephalopods are, for the most part, well reso lved and relatively we ll-accepted. Specieslevel taxa can usually be placed in welldefined families (Jereb and Roper, 2005). However, phylogenetic relationships among fa milies within the major groups remain uncertain (Vecchione 2002). Sweeney and Roper (1998, p.561) addressed the confusion, stating, We realize that numerous systematic problems exist at all taxonomic levels of the Cephalopoda. For example, several higher taxa have been proposed (i.e., superorder Pseudoctobrachia Guerra, 1992, and order Ci rroctopodida Young, 1989). The resolution of these problems requires considerable res earch and review, as new cephalopod research initiatives are being pursued globally. 3.3 Comparative assessment of group in GOM One of the elements absent in current cephalopod research is comprehensive studies of large oceanic or faunal regions. Numerous isolated island studies, studies from a fisheries perspective, or those for biomed ical advances do exis t, but the need for comprehensive systematics, abundance, di stribution, and ecology requires significant attention. The Broad Caribbean Realm, wh ich includes the Caribbean Sea proper, the Gulf of Mexico, and waters that extend through th e Bahamas, is an area that is in need of such comprehensive study. The first author of this study currently is researching this

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57 area. Species that have been collected in th e area allow the opportunity to further define phylogenetic relationships within the cephalopo ds as a whole as well as to address the trophic importance of cephalopods within the region, for example. Because of the great diversity in the fo rm, habitat, and behavior of cephalopods, no single method is adequate to capture and/or sample the complex cephalopod fauna (Rathjen 1991). The excellent vision and high mobility of most cephalopods traditionally means that they have been undersampled with standard trawling and collecting protocols. Despite their limitations, midwater trawls o ffer a starting point for population assessment of pelagic species and provide minimum estimates of oceanic cephalopod abundance (Passarella 1990). Numerous facilities around the Broad Caribbean house unidentified cephalopods which when identified, will add further insight to the diversity of the cephalopod families and ecology, and provide a better fisheries perspective about potentially viable future catches in the region. 3.4 Explanation of Checklist The classification and nomenclature used here follow that of Vecchione (2002), as it is the most recent compilation for the West ern Atlantic region. Orders and families are arranged phylogenetically, and genera a nd species are arranged alphabetically. Cephalopods are not exclusively benthic as are most other mollusca, and many are highly mobile, pelagic/oceanic species. This habita t niche requires the use of the term central in the Gulf of Mexico range column. Dept h data and overall range for organisms are in italics where they could not be determined ex clusively for the Gulf of Mexico. Depth ranges include paralarval thr ough adult stages, so they repr esent the total known vertical range for the species. However, most pela gic species exhibit several more specific

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58 ranges during different phases of their life cycl es: for example, paralarvae are epipelagic, restricted to the upper 100 m; many then undergo ontogenetic descent, moving into deeper waters with growth. Adults of many species then exhibit diel vertical migration from around 400 m to 1000 m during the day into the upper 200 m at night to feed. Very little information is available on biol ogy and lifestyle for ma ny of the deep sea cephalopods in the Gulf of Mexico; this explains the unknown notation in some columns. The abbreviations used in the HabitatBiology category are as follows: bat = bathypelagic (> 1000 m); ben = benthic; cep = coastal surface and epipelagic; crr = coral reef; cts = continental shelf; dps = deep sea; end = endemic; epi = epipelagic (0 m); ins = insular; mes = mesopela gic (200 m); ner = neritic; oce = oceanic; sft = mud, sands, clays; sgr = seagrass; shw = shallo w water; slp = continental slope. The abbreviations used in the Overall geographi c range category are as follows: AT = Atlantic Ocean; BE = Bermuda; BH = Baha mas; BR = Brazil; CH = Cape Hatteras, North Carolina; CT = Connecticut; CR = Caribbean; FL = Florida; IO = Indian Ocean; ME = Mediterranean; N = North; NE = New E ngland; PO = Pacific Ocean; S = South; SA = South America; ST = Subtropical; T = Tropical; TWA = Tropical Western Atlantic; UR = Uruguay; W = West; WA = Western Atlantic. The abbreviations used in the Gulf of Mexico Range category vary because of the high mobility of the species. In some cases, a specific region cannot be defined at this point. Therefore, the term, central (cen ) is indicative of the mid-Gulf of Mexico species. The term, entire (ent) is used wh ere the species is found throughout all regions of the Gulf of Mexico. Fo r those species that are found in more than one region, an

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59 overall region is use. For example, instead of se and ne being used, the term e (east) is used where appropriate.

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60 3.5 Acknowledgements (HJ) would like to acknowledge Dr. Jo seph Torres and the College of Marine Science at the University of South Florida fo r providing invaluable guidance, resources, and facilities for furthering her doctoral research.

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61 REFERENCES 1. Aldred, R. G., M. Nixon and J. Z. Young. 1983. Cirrothauma murrayi Chun, finned octopod. Philosophical Transa ctions of the Royal Society B 301: 1. 2. Clarke, M., and C. Lu. 1974. Vertical di stribution of cepha lopods at 30N 23W in the North Atlantic. Journal of the Marine Biological Association, U.K. 54(4): 969. 3. Clarke, M., and C. Lu. 1975. Vertical di stribution of cepha lopods at 18N 25W in the North Atlantic. Journal of the Marine Biological Association, U.K. 55(1): 165. 4. DeBose, J. L., and M. Vecchione. 2005. First documentation of the Roper inshore squid, Loligo roperi (Cohen 1976) in the Gulf of Mexico. Gulf of Mexico Science 23(1): 132. 5. Hochberg, F. G., M. Nixon and R. B. Toll. 1992. Order Octopoda Leach, 1818, Pp. 213 in Sweeney, M. J., C. F. E. Roper, K. M. Mangold, M. R. Clarke and S. v. Boletzky, eds. L arval and juvenile cephalopods: A manual for their identification., Sm ithsonian Contributions to Zoology 513: 1-282. 6. Jereb, P. and C.F.E. Roper, eds. 2005. Cephalopods of the World. An Annotated and Illustrated Catalogue of Sp ecies Known to Date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spir ulidae). FAO Species Catalogue for Fishery Purposes. No. 4, Vol. 1 Rome, Italy. 7. Kubodera, T., U. Piatkowski, T. Okutan i, and M. R. Clarke. 1998. Taxonomy and zoogeography of the family Onycho teuthidae (Cephalopoda: Oegopsida). Pp. 277 in N. A. Voss, R. B. Toll, and M. Vecchione, eds. Systematics and Biogeogra phy of Cephalopods. Smithsonian Contributions to Zoology 586. 8. LaRoe, E. T. 1971. The culture and Ma intenance of the Loliginid Squids Sepioteuthis sepioidea and Doryteuthis plei Marine Biology; v. 9, 9-25. 9. Lee, P. G., P. E. Turk, W. T. Yang, and R. T. Hanlon. 1994. Biological characteristics and biomedic al applications of the squid Sepioteuthis lessoniana cultured through multiple generations. Biological Bulletin 186: 328.

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62 10. Lesueur, C.A. 1821. Descriptions of Seve ral New Species of Cuttle-fish. Journal of the Academy of Natural Scie nces of Philadelp hia, 2(1):86-101. 11. Lipka, D. A. 1975. The Systematics and Zoogeography of Cephalopods from the Gulf of Mexico. Ph.D. dissertat ion, Texas A&M University, College Station, Texas. 351 pp. 12. Nesis, K. N. 1975. Cephalopods of the American Mediterranean Sea. English Translations of Selected Pub lications on Cephalopods. Smithsonian Institution Libraries, 2003, 1(1): 319. 13. Nesis, K. N. 1987. Cephalopods of the Wo rld. Squids, Cuttlefishes, Octopuses, and Allies. (trans. B. S. Levitov). T.F.H. Publications, Neptune City, New Jersey. 351 pp. 14. Nixon, M., and J. Z. Young. 2003. The Brains and Lives of Cephalopods. Oxford University Press, Oxford, U.K. 448 pp. 15. Okutani, T. 2005. Cuttlefishes and Squi ds of the World; 253 Pp. National Cooperative Association of S quid Processors, Tokyo, Japan. 254 pp. 16. Passarella, K. C. 1990. Oceanic Cephalopod A ssemblage in the Eastern Gulf of Mexico. Masters thesis, University of South Florida, St. Petersburg, Florida. 50 pp. 17. Passarella, K. C., and T. L. Hopkins. 1991. Species composition and food habits of the micronektonic cephalopod asse mblage in the Eastern Gulf of Mexico. Bulletin of Marine Science 49(1): 638. 18. Rathjen, W. F. 1991. Cephalopod Capture Me thods: An Overview. Bulletin of Marine Science. 49(1-2): 494. 19. Roper, C. F. E. 1964 Enoploteuthis anapsis a new species of enoploteuthid squid (Cephalopoda: Oegopsida) from the Atlantic Ocean. Bulle tin of Marine Science of the Gulf and Caribbean 14(1): 140. 20. Roper, C. F. E., C. C. Lu, and K. Mangold. 1969. A new species of Illex from the Western Atlantic and dist ributional aspects of other Illex Species (Cephalopoda: Oegopsidae). Procee dings of Biological Society of Washington 82: 295.

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63 21. Roper, C. F. E., C.C. Lu, and M. Vecchione. 1998. A revision of the systematics and distribution of Illex species. Pp. 405-423 in N.A. Voss, M. Vecchione R. B. Toll and M. J. Sweeney, eds. Systematics and Biogeography of Cephalopods. Smithsonian Contri butions to Zoology, 586(I-II). Tokai University Press. Tokyo, Japan. 22. Roper, C. F. E., M. J. Sweeney and M. Clarke. 1985. FAO Species Identification Sheets for Fishery Purposes. Pp. 117 in W. Fischer, and J. C. Hureau, eds. Southern Ocean. Food and Agricultural Organization of the United Nations, Rome, vol. 1. 23. Roper, C. F. E., M. J. Sweeney, and C. Nauen. 1984. Cephalopods of the World. An Annotated and Illustrated Catalogue of Species of Interest to Fisheries. FAO Fisheries Synopsis No.125, v.3, Food and Agricultural Organization of the United Nations, Rome. 277 pp. 24. Roper, C. F. E., and M. Vecchione. 1993. A geographic and taxonomic review of Taningia danae Joubin, 1931(Cephalopoda: Octopoteuthidae), with new records and observations on bioluminescence. Pp. 441 in T. Okutani, R. K. ODor and T. Kubodera, eds. The Recent Advances in Cephalopod Fishery Biology. 25. Roper, C. F. E., and R. E. Young. 1975. Vertical distribut ion of pelagic cephalopods. Smithsonian Contributions to Zoology 209: 1. 26. Salcedo-Vargas, M. A. 1991. Checklist of the cephalopods from the Gulf of Mexico. Bulletin of Marine Science 49(1-2): 216. 27. Summers, W. C. 1983. Physiological a nd trophic ecology of cephalopods. Pp. 261-279 in W. D. Russell-Hunter, ed. The Mollusca, Volume 6. Ecology, Academic Press, Orlando, Florida. 28. Sweeney, M. J., and C. F. E. Roper, 1998. Cl assification, type local ities, and type repositories of Recent Cephalopoda. Pp. 561 in N. A. Voss, M. Vecchione, R. B. Toll and M. J. Sweeney, eds. Systematics and Biogeography of Cephalopods. Smiths onian Contributions to Zoology 586(I-II). 29. Vecchione, M. 2002. Cephalopods. Pp. 150-244 in K. E. Carpenter, ed. The Living Marine Resources of the We stern Central Atlantic. Volume 1. Introduction, mollusks, crustaceans, ha gfishes, sharks, batoid fishes and chimaeras. FAO Identificatiion Guid e for Fisheries Purposes. The Food and Agriculture Organization of the United Nations, Rome, Italy.

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64 30. Vecchione, M. and C. F. E. Roper. 1991. Cephalopods observed from submersibles in the Western North Atlantic. Bulletin of Marine Science 49(1-2): 433. 31. Vecchione, M., C. F. E. Roper and M. J. Sweeney. 1989. Marine flora and fauna of the Eastern United States. Mollusca: Cephalopoda. NOAA Technical Report NMFS 73. U.S. Depa rtment of Commerce. 23 pp. 32. Vecchione, M., and R. E. Young. In press. The squid family Magnapinnidae (Mollusca: Cephalopoda) in the Atlan tic Ocean, with descriptions of new species. 33. Villanueva, R., M. A. Collins, P. Sanch ez and N. A. Voss. 2002. Systematics, distribution, and biology of the cirrate octopods of the genus Opisthoteuthis (Mollusca, Cephalopoda) in the Atlantic Ocean, with description of two ne w species. Bulletin of Marine Science 71(2): 933. 34. Voss, G. L. 1956. A review of the cephalopods of the Gulf of Mexico. Bulletin of Marine Science of the Gulf and Caribbean 6(2): 85. 35. Voss, G. L. 1964. A note on some cephalopods from Brazil with a description of a new species of octopod, Eledone massyae Bulletin of Marine Science of the Gulf and Caribbean 14(3): 511. 36. Voss, G. L. 1988. Evolution and phyloge netic relationship of deep-sea cephalopods (Cirrata and Incirrata). Pp. 253 in M. R. Clarke, and E. R. Trueman, eds. The Mollusca, Vol. 12. Paleontology and neontology of cephalopods. Academic Press, San Diego, California. 37. Voss, G. L., and T. F. Brakoniecki. 1985. S quid resources of the Gulf of Mexico and Southeast Atlantic coasts of the United States. Pp. 27-37 in V. M. Hodder, ed. Northwest Atlantic Fisher ies Organization: Sc ientific Council Studies, Special Session on Squids 9. 38. Voss, G. L., L. Opresko, and R. Thom as. 1973. The potentially commercial species of octopus and squid of Fl orida, the Gulf of Mexico and the Caribbean Sea. University of Miami Sea Grant Field Guide Series, No. 2: 1-33. 39. Voss, G. L., and R. R. Toll. 1998. The system atics and nomenclatural status of the octopodinae described from the Western Atlantic Ocean. Pp. 457 in N. A. Voss, R. B. Toll, and M. Vecchione, eds. Systematics and Biogeography of Cephalopods. Smiths onian Contributions to Zoology 586(I-II).

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65 40. Voss, N. A., and G. L. Voss. 1962. Two new species of squids of the genus Calliteuthis from the Western Atlantic with a redescription of Calliteuthis reversa Verrill. Bulletin of Marine Scie nce of the Gulf and Caribbean 12(2): 169. 41. Voss, N. A. 1980. A generic revision of the Cranchiidae (Cephalopoda; Oegopsida). Bulletin of Marine Science 30: 365. 42. Voss, N. A. 1988. Systematics and zooge ography of cephalopods. Malacologia 29(1): 209. 43. Voss, N. A., K. N. Nesis, P. Rodhouse. 1998. The cephalopod family Histioteuthidae (Oegopsida): system atics, biology, and biogeography. Pp. 293 in N. A. Voss, R. B. Toll, and M. Vecchione, eds. Systematics and Biogeography of Cephalopods. Smith sonian Contributions to Zoology 586(II). 44. Young, R. E., C. F. E. Roper. 1969. A monograph of the Cephalopoda of the North Atlantic: The family Cycloteu thidae. Smithsonian Contributions to Zoology 5: 1. 45. Young, R. E. 1975. Leachia pacifica (Cephalopoda, Teut hoidea): Spawning habitat and function of the brach ial photophores. Pacific Science 29: 19. 46. Young, R. E. 1978. Vertical distribution and photosensitive vesicles of pelagic cephalopods from Hawaiian Waters. Fishery Bulletin 76: 583. 47. Young, R. E., Vecchione, M., and Mangold K. 1995. Coleoidea Bather, 1888. Octopods, squids, cuttlefishes and th eir relatives. Version 01 January 1995 (under construction). http://tolweb.org/Col eoidea/19400/1995.01.01 in The Tree of Life Web Project, http://tolweb.org/ 48. Young, R. E, L. A. Burgess, C. F. E. Roper, M. J. Sweeney, and S. J. Stephen 1998. Classification of the Enoploteuthidae, Pyroteuthidae, and Ancestrocheiridae. Pp. 239 in N. A. Voss, R. B. Toll, and M. Vecchione, eds. Systematics and Biogeography of Cephalopods. Smithsonian Contribut ions to Zoology 586(1).

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66 Table 3.1: Checklist of cephalopods (Mo llusca: Cephalopoda) from the Gulf of Mexico Taxon Habitat-Biology Depth (m) Overall geographic range GOM range References/Endnotes Class: Cephalopoda Order: Spirulida Family: Spirulidae Spirula spirula (Linneaus, 1758) oce 550 worldwide T & ST ent 14, 29 Order: Sepioidea Family: Sepiolidae Austrorossia antillensis (Voss, 1955) oce 305 CR, N SA, GOM se 6, 12, 23, 34 Heteroteuthis dispar Ruppell, 1845 mes, oce 200 T, ST worldwide e 14, 16 Rossia bullisi Voss, 1956 end, oce ? T AT ent 12, 34 Semirossia equalis (Voss, 1956) sft 130 GOM, N SA ne 6, 12, 23, 34 Semirossia tenera (Verrill, 1880) sft 85 NE to GOM & CR ent 6, 12, 31 Order: Myopsida Family: Loliginidae Doryteuthis pealeii (LeSueur, 1821) cep, cts 1 WA; NS to VE; GOM and CR ent 12, 14, 29 Doryteuthis plei (Blainville, 1823) cep, cts, slp 1 WA, GOM, CR ent 12, 14, 29, 34 Doryteuthis roperi (Cohen 1976) cep, cts 1 WA, GOM, CR ent 4, 34

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67Lolliguncula brevis (Blainville, 1823) cts, shw 1 WA, GOM, CR ent 12, 29 Pickfordiateuthis pulchella Voss, 1956 sgr 1 FKBR se 14, 34 Sepioteuthis sepioidea (Blainville, 1823) crr, shw 1 BE, FL, BH, & CR se 12, 29, 34 Order: Oegopsida Family: Architeuthidae Architeuthis dux Steenstrup, 1860 oce, cts 400 AO ent 23 i Family: Brachioteuthidae Brachioteuthis sp. epimes 1 NA, PO, ME e 14, 16, 29 Family: Cranchiidae Bathothauma lyromma Chun, 1906 mes 1 T, ST e 16, 40 Cranchia scabra Leach, 1817 oce 2 T, ST worldwide ent 12, 14, 16 Egea inermis Joubin, 1895 oce 1 TA, WP, IO e 14, 16, 40 Helicocranchia pfefferi Massy, 1907 oce 1 T, ST worldwide e 14, 16 Leachia atlantica (Degner, 1925) oce 1 T, ST ent 16, 40, 44 Leachia cyclura LeSueur, 1821 oce 1-2000 T, ST worldwide ent 40 Leachia lemur (Berry, 1920) oce 1-2000 T, ST worldwide ent 40 Liocranchia reinhardti (Steenstrup, 1856) oce 1 circumglobal e 12, 14, 16 Megalocranchia spp. oce 1 T, ST worldwide e 16, 22 Family: Cycloteuthidae Cycloteuthis sirventi Joubin, 1919 oce 200-1000 T, ST AT ? 15, 29

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68Discoteuthis discus Young and Roper, 1969 oce 1 AT, S PO e 14, 16, 25 Discoteuthis laciniosa Young and Roper, 1969 epimes 400-1000 T, ST, AO, IO, PO ? 45, 48 Family: Enoploteuthidae Abralia redfieldi Voss, 1955 epi, ins 1 T, ST WA e 12, 14, 34 Abralia veranyi (Ruppell, 1844) oce, slp 20-800 T, ST AT, E AT ent 12, 14, 34 Abraliopsis atlantica Nesis, 1982 mes 1 T, ST AT e 16, 48 Abraliopsis pfefferi Joubin, 1896 mesbat 1 T, ST AT, IO, W PO e 2, 12, 14, 16 Enoploteuthis leptura (Leach, 1817) oce 200 T, ST AT, IO, W PO e 12, 16 Family: Magnapinnidae Magnapinna sp. bat, oce 1000 GOM, NA cen 32 ii Family: Pyroteuthidae Pterygioteuthis gemmata Chun, 1908 oce 1-600 T worldwide e 16, 23 Pterygioteuthis giardi Fischer, 1896 oce 1 T, ST worldwide e 12, 16, 23 Pyroteuthis margaritifera (Ruppell, 1884) oce 75 T, ST AIWPO e 12, 16, 25 Family: Ancistrocheiridae Ancistrocheirus lesueuri Orbigny, 1839 mes 1 T, ST worldwide e 12, 14, 16, 25, 46 Family: Histioteuthidae Histioteuthis bonnellii (Ferussac, 1834) oce 1 T, ST worldwide cen 14, 41 Histioteuthis corona (Voss & Voss, 1962) oce 200 T, ST worldwide e 12, 16 Histioteuthis reversa (Verrill, 1880) oce 1 ? cen 12, 29, 34, 42

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69Family: Bathyteuthidae Bathyteuthis abyssicola Hoyle, 1885 oce 200 T, ST worldwide e 12, 16, 34 Family: Chtenopterygidae Chtenopteryx sicula (Verany, 1851) mes, oce 200 AT, PO, ME e 14, 16 Family: Lepidoteuthidae Lepidoteuthis grimaldii Joubin, 1895 oce ? T worldwide ent 12, 14, 16 Family: Lycoteuthidae Lampadioteuthis megleia Berry, 1916 mes, oce 200 AT, SPO e 14, 16, 29 Lycoteuthis lorigera (Steenstrup, 1875 ) mes, oce 200 WA, GOM cen 14, 21, 29, 34 Lycoteuthis springeri (Voss, 1956) end, oce 200 T WA cen 12, 34 Selenoteuthis scintillans Chun, 1900 oce 200 WA, GOM e 14, 16 Family: Ommastrephidae Hyaloteuthis pelagica (Bosc, 1802) oce 1;1700 T, ST worldwide ent 12, 13, 23, 34 Illex coindetii (Verany, 1839) cts, ner, oce 1 AT ent 14, 22, 25, 29 Illex oxygonius Roper, Lu, and Mangold, 1969 ner, oce 50 WA se 12, 14, 29, 31 Ommastrephes bartramii (LeSueur, 1821) oce 1 worldwide ent 12, 29, 31 Ornithoteuthis antillarum Adam, 1957 ner, oce 1 PO, AT ent 14, 16, 29, 31 Sthenoteuthis pteropus (Steenstrup, 1855) oce 1 T, ST Pan-Atlantic ent 16, 29, 34 Family: Chiroteuthidae Chiroteuthis joubini Voss 1967 mes, oce 200 ST AO, IO e 16, 48

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70Chiroteuthis veranyi (Fersussac, 1834) mes, oce 200 T, ST W AT, E PO ent 12, 34 Chirotuethis mega (Joubin 1932) mes, oce 200 ? e 16, 48 Grimalditeuthis bonplandi (Verany, 1839) oce 200 AT, GOM cen 12, 29, 34 Planctoteuthis danae (Joubin, 1931) mes, oce 200 worldwide T, ST cen 29 Family: Pholidoteuthidae Pholidoteuthis adami Voss, 1956 oce 20 PO, GOM, AT cen 14, 34 Pholidoteuthis boschmai Adam, 1950 mes, oce 200 ST worldwide cen 29, 48 Family: Octopoteuthidae Octopoteuthis megaptera (Verrill, 1885) mes, oce 200 T, ST A se 12, 34 Octopoteuthis sicula Ruppell, 1844 mes, oce 200 T, ST A, IWP ? 15, 48 Tanangia danae Joubin, 1931 oce 25 T, ST worldwide e 12, 16, 24 Family: Onychoteuthidae Moroteuthis aequatorialis Theile, 1920 oce 100 T, ST AT, IO, W PO e 12, 48 Onychoteuthis banskii (Leach, 1817) oce 100 Worldwide se 7, 12, 14, 34, 38 Onykia carriboea LeSueur, 1821 epi, oce 1 T, ST worldwide e 12, 34 Family: Thysanoteuthidae Thysanoteuthis rhombus Troschel, 1857 oce 25 day nets AT, PO, IO, ME e 12, 14, 16, 25 Family: Mastigoteuthidae Mastigoteuthis agassizi Verrill, 1881 oce 200 T, ST A e 16, 29 Mastigoteuthis hjorti Chun, 1913 oce 200 T A cen 29

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71Mastigoteuthis magna Joubin, 1913 oce 200 N AT, GOM cen 29 Family: Joubiniteuthidae Joubiniteuthis portieri (Joubin, 1912) oce 1 AO, PO ent 2, 3, 12, 14, 16 Order: Octopoda Family: Opisthoteuthidae Opisthoteuthis agassizi Verrill 1883 ben, cts 100 AT, PO ent 12, 29, 33, 34, 36 Family: Octopodidae Benthoctopus januarii (Hoyle, 1885) dps 400 GOM, CR, TAT cen 12, 29 Octopus briareus Robson, 1929 shw ? W N AT, SE USA, BH, CR s 12, 29 Octopus burryi Voss, 1950 cts 10-200 GOM, CH to BR e 12, 29 Octopus defilippi group shw 6 > 60 WA, FL to BR e 12, 16, 29 Octopus hummelincki Adam, 1936 shw 1 TWA, FL to BR se 29 Octopus joubini Robson,1929 cts, shw 1 TWA ent 12, 29 Octopus macropus group shw ? TWA se 12, 29 Octopus maya Voss & Solis Ramirez, 1966 sgr, shw 1 S GOM s 12, 29 Octopus mercatoris Adam, 1937 ? ? GOM ? 29 Octopus cf. vulgaris group cts 1 WA, CT to BR ent 12, 29 Pteroctopus schmidti (Joubin, 1933) dps 300 Scattered se 29 Pteroctopus tetracirrhus (Chiaie, 1830) cts, sft 100 WA, CH to UR ent 12, 29 Scaeurgus unicirrhus (Chiaie, 1839-1841) sft 100 WA, N CH to S BR e 15, 29

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72Tetracheledone spinicirrus Voss, 1955 sft 200 GOM s 12, 29 Family: Alloposidae Haliphron atlanticus Steenstrup, 1861 oce 1 AT, PO, IO T, ST ent 12, 14, 16, 34 Family: Ocythoidae Ocythoe tuberculata Rafinesque, 1814 mes, oce 1 T, ST worldwide cen 48 Family: Bolitaenidae Bolitaena pygmaea (Verrill, 1884) oce >1000 T, ST WA ? Japetella diaphana Hoyle, 1885 dps 600 T, ST worldwide e 5, 12, 16, 45 Family: Tremoctopodidae Tremoctopus violaceus Chiaie, 1830 oce 1 T, ST worldwide se 12, 14, 25, 34 Family: Argonautidae Argonauta argo Linnaeus, 1758 oce 1 T, AT, PO, ME e 12, 16, 34 Argonauta hians Lightfoot, 1786 29 Order: Vampyromorpha Family: Vampyroteuthidae Vampyroteuthis infernalis Chun, 1903 bat 100 T, ST worldwide ent 12, 14, 16, 30, 34

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73 Table 3.2: Taxonomic summary for Ce phalopods of the Gulf of Mexico. Component Subgroups Spirulida Sepiodea Myopsida Oegopsida Octopoda Vampyromorpha Total Species 1 5 6 58 22 1 Number Endemic Not enough study yet to give good account Number Nonindigenous species Only 2 known at this time 1 1

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74 CHAPTER 4 FIRST RECORDS OF Asperoteuthis acanthoderma (Lu, 1977) (CEPHALOPODA: OEGOPSIDAE: CHIROTEUTHIDAE), FR OM THE NORTH ATLANTIC OCEAN, STRAITS OF FLORIDA Heather Judkins, Debra A. Ingrao and Clyde F. E. Roper (HJ) College of Marine Science, University of South Florida, 140 7th Ave South, St.Petersburg, FL 33701; Hjudkins@marine.usf.edu (DI) Mote Marine Laboratory, 1600 Ken Thompson Parkway, Sarasota, FL 34236; dingrao@gmail.com (CR) Musuem of Natural History, Smith sonian Institution, Washington D.C. 200137012 USA; gsquidinc@verizon.net 4.1 Abstract The first observation in the North Atla ntic Ocean of the deep sea squid Asperoteuthis acanthoderma (family Chiroteuthidae) is repo rted here from off the coast of Key West, Fl in the Straits of Florida. We describe the morphology of the two nearly complete, but damaged, specimens. A third reco rd is based on photographs of a specimen from off Grand Cayman Island; this specimen was not available for examination. The multiple occurrences of this species, hereto fore unknown in the North Atlantic Ocean, within a 10-month period are so unusual that we attempt to hypothesize an explanation for these events. All previously known records are recorded from a few specimens

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75 scattered from Hawaii to the Philippines. The present specimens were identified by the following characteristics unique to the species: Y-shaped funnel locking apparatus, sucker ring form and dentition, beak morphology, photophore patch configuration on ventral surface of eyeballs, and numerous, sm all cartilaginous tubercles that cover the mantle, head and the aboral surface of the arms. 4.2 Introduction A mature female Asperoteuthis acanthoderma was discovered approximately 90 miles southwest of Key West, Florida on Fe bruary 20, 2007. It was found floating at the surface above a bottom depth of approximately 259 meters. The specimen was retrieved by a charter boat captain, Clint Moore, who recognized the uniquene ss of the squid and donated it to Mote Marine Laboratory in Saraso ta, Florida. This sp ecimen is now located in the cephalopod collection of the Smithsonian Institutions National Museum of Natural History in Washington D.C. A second speci men was found by Captain Jack Carlson in the Straits of Florida, 16 miles east of Ma rathon, Florida in June 2007, floating at the surface above a water depth of approximately 355m (24 43N 81 06W). This specimen is deposited in the Marine Invertebrate Mu seum at the Rosenstiel School of Marine and Atmospheric Sciences, University of Mi ami, in Miami, Florida (UMML 31.3212). The posterior portion of the primary fin is missing in the Key West specimen, so the mantle length (ML) is measured from the anterior tip of the mantle, along the dorsal mid-line, to the posterior end of the existing portion of the fins. Also missing are both tentacular clubs and stalks, with the exception of a very short section of the left tentacle. Internal structures are in relatively good condition, includ ing the female reproductive

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76 organs, so that the principal specific characters are pres ent except for those found on the tentacular clubs. The Marathon specimen compliments the Key West specimen by having more complete tentacles that show the photophore pa tches along their lengths, and it retains the majority of the pair of flaps known as the sec ondary fin or tail flap along the sides of the tail. The tentacular clubs and eyes ar e missing. Although the internal organs are damaged, sufficient evidence remains to indicat e that the specimen is a spent female. 4.3 Systematics Asperoteuthis acanthoderma (Lu, 1977) Diagnosis .---A chiroteuthid with mantle, head an d arms covered with numerous, minute cartilaginous tubercles; photophore patch pres ent on ventral surface of eyes; 3-4 broad, rounded teeth on arm-sucker rings; funnel lock ing apparatus an inverted Y-shape. (Lu, 1977) Material examined .---Specimen 1. (Fig. 1) Female, 620+ ML, 90 miles southwest of Key West, FL. Found floating dead at surface. The posterior tip of the mantle, primary, and secondary fins are missing. When a pproximate measures are used, the TL is estimated to have been 1250 mm. Specimen 2. (Fig.2) Female, 1630 mm ML, and 3420 TL; measured to tip of tail posterior to t ail flap.. Found floating dead at surface, 16 miles off Marathon, FL. 4.4 Description External anatomy .---The description is based primarily on the Key West specimen with additional details included fr om the Marathon specimen as appropriate. The mantle is long, narrow, semigelatinous, with dark purple pigmentation. Numerous,

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77 minute cartilaginous tubercles cover the mantle head and arms. The mantle is tapered posteriorly towards the fins. The fins together form an elongate oval shape and are estimated to have been 220 mm long (incl uding estimated length of missing portion). The fin length is therefore approxima tely 50% of ML. The width of the fins is 340 mm at their widest part. The tail with its posterior extens ion of the gladius and lateral structures is missing altogether. The Marathon specimen had a nearly complete fin assemblage with only the very tip of the tail missing. The fi n length measures 460 mm, the tail is 610 mm long. The fin width measures 330 mm and the secondary fin width is 235 mm. The fins and tail flaps are each elongate oval-shape d. The fins become proportionately narrower posteriorly. The tail flaps are very thi n, while the fins are noticeably thicker. The funnel is large in comparison to th e head. It measures 173 mm from the anterior funnel opening to the posterior border along the ventral midline. The diameter of the anterior funnel aperture measured (flatt ened) is 86 mm. The funnel locking apparatus is ovoid and is in an inverted Y-shape; the mantle component matches the funnel counterpart. The dorsal component of the f unnel organ is roughly diamond shaped with one triangular flap on each side. The ventral pads are present and oval shaped. The flaps form a skirt-like sheath that is attached to the dorsal surface of the funnel organ suggesting that this organ hangs ventrally ope n while the animal is alive. The funnel valve is very large. The head is elongate, narrow ; it is deep dorsoventra lly, narrow laterally. Olfactory papillae are present on each side of the postero-lateral surface of the head in the form of a slender stalk with a bulbous termin al head. The eyes are large with a ventral photophore patch that extends toward the posteri or surface of each eye. Diameter of right

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78 eye, 28mm; horizontal openi ng of the eyes: left, 44 mm and right, 37 mm. The nuchal cartilage is long and narrow; the central groove is distinct. The arms are very long and attenuate. L ittle difference is apparent among the lengths of arms, but only two are completely in tact. The intact AL of right arm I is 880 mm and of intact left arm IV, 870 mm. All arms have approximately the same semigelatinous consistency and appear to be closely equal in diam eter at their bases. Right arm III has a distinct keel distally, but no other keels are omm. able. All arms have trabeculae along the oral bases from which the sucker stalks arise. The arm suckers are biserial and evenly spaced. The diameters of the suckers range from 4 mm to 12 mm on each arm, varying slightly between arms but c onsistently patterned smallenlargedsmall as they progress distally. On the arms that are intact, the tips have minute suckers on small stalks, clustered together much more cl osely than along the rest of arms. The arm sucker rings have 3 to 4 broad, rounded teeth around the rim. The remnants of the tentacles of the Key West specimen have only short, extremely thin stalks that extend from the arm crown. The longest remaining portion is on the right side and it measures 100 mm with no visible photophore patches. The Marathon specimen has larger tentacle rema nants measuring 1560 mm on the left side and 780 mm on the right side. No tentacular clubs are present. The tentacles have photophore patches scattered along their length in no discerna ble pattern. Forty such patches occur on the most intact (left) tentacle of the Marathon specimen. The gladius was not extracted from the Key West specimen because it is broken in many places. The gladius of the Marathon specimen is in much better condition. The gladius is approximately 1630 mm long and 74 mm wide at its widest point. The most

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79 anterior portion of the gladius is flattene d and becomes rounded as it extends to the posterior end. The gladius could not be co mpletely reconstructed because pieces are missing. The buccal membrane has 7 connectives th at attach on the dorsal sides of the bases of arms I and II and on the ventral sides of arms III and IV. The lappets are too indistinct to be counted. The beaks were extracted from the Key West specimen for examination and preservation. The anterior tip of the upper ma ndible is narrow, sharply pointed. It has a deep, dark brown pigmentation. The hood is normal in size and has a brownish coloration that lightens toward the edges, which are translucent in appearance. The curve of the dorsal half of the hood margin of the beak is irregularly sh aped, not smooth. The anterior tip of the lower beak also is pointed with its inner edges curved. It is somewhat broader than the upper mandible, with dark brown pigmentation and the edges of the wings are a lighter brown in tone. The radula has 7 teeth per transverse row. The width at the dorsal side of the radula measures 4 mm, while the ventral porti on of the set measures 3.5 mm. The length of the radula is 14 mm. The rhachidian tooth ha s a long central cusp with shorter, lateral cusps. The first lateral toot h is bicusped, with a long, point ed cusp and a shorter, blunt lateral cusp. The second lateral tooth is point ed with a broad base but no lateral cusp. The third lateral tooth is longer than the ot her lateral teeth and is pointed and curved. Most of the surface of the s quid is covered with cartilagi nous tubercles. They are very numerous and thickly-concentrated on the head, mantle and aboral surfaces of arms. There appear to be fewer on the oral surfaces of arms. Each tubercle has a wide base that

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80 ends with a pointed cartilag inous tip. Numerous dark pur ple chromatophores cover the squid, as well. Histological sections of the t ubercles of the mantle wall indicate that they are similar to a published desc ription (Roper and Lu, 1990), as well as in the original description of the mantle by Lu (1977) which states that there is an outer vacuolated dermal layer followed by a longitudinal muscle layer, an exceptionally thick vacuolated medullary layer, an inner circular muscle layer and the lining of the mantle cavity. Internal anatomy .---The internal organs of the Key West specimen are in relatively good condition, while the organs from the Marathon squid are significantly deteriorated. Key West specimen (Fig. 3): The gladius extends the length of the mantle on the internal dorsal side. Th e gills extend approximately half the length of the mantle cavity on both sides, lateral to the digestive gl and. The ink sac is small and appears to be empty; it was difficult to locate during dissecti on. The nidamental gl ands lay medial to the gills. They are paired, ova l, enlarged and milky white in appearance; the right gland is damaged by a tear. The branchial hearts are small, white, and round; they are located posterior to the kidney at the base of each gill with the systemic heart lying along the central midline, dorsal to the nidamental gl ands. The digestive or gans are intact; the stomach, thin walled, and collapsed, leads to th e caecum, which is creamy white in color, and thick with an ovoid spongy, reticulated mass. The ovary is posterior to the digestive organs, translucent and appears empty of oocytes and eggs. All organs appear proportional in size compared with mantle le ngth, with the exception of the substantially enlarged nidamental glands.

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81 4.5 Discussion The first species of Asperoteuthis was originally desribed as Chiroteuthis acanthoderma by Lu (1977) from the tropical wester n Pacific. Nesis (1980) moved this species to the new genus Asperoteuthis He considered C. acanthoderma Lu, 1977 to be a junior synonym of Aspreoteuthis famelica (Berry 1909), described from a damaged specimen caught near Hawaii. Subsquent s tudies found that A. famelica was actually Mastigotuthis famelica (Young, 1978), so the valid name for Lus specimens is Asperoteuthis acanthoderma (see Young et al. 2007, Arkhipkin et. al, 2008). We identify the present specimens as Asperoteuthis acanthoderma because of the unique characteristics that exist based on Lus description of the species. The genus Asperoteuthis is distinguished from the other genera of the family Chiroteuthidae by the presence of a terminal photophore on the tentacular clubs and an elongate oval photophoric patch on the ventral surface of each eyeball; the absence of intestinal photophores and of a tragus/antitragus in the f unnel component of the locking apparatus. Other features that distinguish Asperoteuthis from other chiroteuthids include 1) the structure of arm IV is similar to that of arms I-III, not significantly enlarged in advanced subadults; 2) absen ce of characteristic pairs of adjacent club suckers drawn together with an intermediate widening on th e stalk and absence of an enlarged, central tooth on the club suckers as occurs in other chiroteuthids; and 3) the forked (Y-shaped) funnel cartilage Asperoteuthis acanthoderma (Lu, 1977) is characterized by the possession of numerous, minute, sharply pointed, conical cartilaginous tubercles distributed over the

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82 entire surface of the mantle head, and arms. The tubercles measure 1 mm in diameter and 0.4 mm in height from a specimen of 144 mm ML. The tubercles are discrete structures, and their bases are embedded in the dermis of the mantle (Roper & Lu 1990, as C. acanthoderma). The small number of rounded teeth on the arm suckers (3-4) and the prominence if the funnel valve (Tsuchiy a & Okutani 1993) are additional specific characteristics. The morphological characteristics of our sp ecimens, that are about 4 times larger than the holotype and paratype, conform clos ely with the original description, which was based on females of 188 mm and 144.5 mm ML. Exceptions are minor, such as the inconspicuousness of the dermal tubercles on the smaller specimens, and the number of knobs on the tentacular stalks. We concur w ith Tsuchiya and Okuntani, (1993) that these and other minor differences are attributable to variation due to the size and stages of maturity of specimens. Distribution .---Asperoteuthis acanthoderma has been recorded in the central and western Pacific and Indo-Pacific waters as follows: in the southern part of Philippine Sea and in the Celebes Sea to the west, southeast, and east of the Philippine Islands by Lu (1977) and Nesis (1980), (as Chiroteuthis acanthoderma and Mastigoteuthis famelica, respectively). A specimen of A. acanthoderma was recorded near the bottom by a SERPENT-Project ROV off western Australia (18 30S, 115 30E) at 580 m depth in April, 2005 (Vecchione pers. omm..). A recent su rvey of literature of the cepha lopods of the Gulf of Mexico revealed that only three genera of Chiroteuthids previously have been recorded there: Chiroteuthis joubini, Chiroteuthis veranyi and Chiroteuthis capensis, as well as Grimalditeuthis

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83 bonplandi (Verany, 1839) and Plancoteuthis danae (Joubin, 1931) (Judkins et. al, in press). The present report of A. acanthoderma is therefore a dramatic range extension into the North Atlantic. Information on the vertical distribution of Asperoteuthis acanthoderma is limited, but the following capture data indicate that this is a deep-sea species (as also demonstrated by its morphology and rarity of capture): Holotype .---914-0 m (Lu 1977). Paralarvae .---Daytime 600-1100 m and night 700-925 m (Roper & Young 1975). Night on horizon of 200-300 m, (Lu 1977, Nesis 1980). Based on the limited information, we believe that the vertical movements of A. acanthoderma may not be ontogenetic descent but diel vertical migrati ons as Nesis (1980) suggested. This is similar to other chirot euthids (Roper & Young 1975). The state of knowledge about the chiroteuthids is e xpanding as teuthologists discover and report new findings of genera and species. For example, Young et. al. (2007) published information of a new Asperoteuthis species found off the coast of Hawaii. Asperoteuthis mangoldae (Young, Vecchione, & Roper, 2007) shares some A. acanthoderma characters but differs in the absence of tubercles in the skin; a more gelatinous consistency of tissues; the absence of an anti-tragus in the funnel locking apparatus; 8-10 slender, truncated teet h on the arm sucker rings; club and club photophore structure and fin lengt h characteristics. Arkhipk in and Laptikhovsky (2008) introduced a fourth species to the Asperoteut his genus, Aspreoteuthis nesisi which shares many characteristics of A. acanthoderma such as the funnel locking apparatus, the numerous tubercles on head and man tle, and the photophore patches on eyes.

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84 Asperoteuthis acanthoderma may be a circumglobal species whose broad distribution has just recently b een recognized because of fort uitous findings of specimens and intensified research in previously under-sampled study areas. The global understanding of the oceanic environment and its inhabitants is enhanced as those who participate in maritime activities, deep s ea fishermen, for example, become more cognizant of their surroundings and the organism s that inhabit their specific regions. Our knowledge is advanced when they report th eir observations and findings to marine scientists. The two specimens we examined from of f Florida and the one record via photos of A. acanthoderma in Grand Cayman suggest that ce rtain physical parameters of the region may be different from earlier years. All three specimens were found within a 10month period from mid-2006 to May 2007, rare di scoveries that seem to be more than coincidental. Even though they appear to be spent females that could have drifted to the surface, a typical event for many deep sea s quids and octopods, the total absence of specimens in the North Atlantic Ocean unt il now could suggest additional causative effects. Based on our detailed examination of the tw o large, nearly intact specimens, and our comparisons with descripti ons in the literature, we conc lude that our specimens are indeed A. acanthoderma Consequently, this species is now shown to occur not only in the tropical western Pacific O cean, but also in the Caribbean /Gulf of Mexico/Straits of Florida region.

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85 4.6 Acknowledgments Photos were taken by Chuck Stevenson at Mo te Marine Laboratory, for which we are grateful. We acknowledge the enormous help of the faithful Mote Marine lab volunteers as the dissections were conducted. We also thank the tw o captains from the fishing vessels that made the original finds and c ontacted us. Captain Clint Moore found the Key West specimen on February 20, 2007, while Captain Jack Carls on discovered the specimen off Marathon, FL on May 17, 2007; He coordinated with Gabe Delgado of the Florida Wildlife Research Institute to ensure retrieval of the specimen. We appreciate the help of Mike Vecchione, NMFS Systematic s Laboratory, Natural Museum of Natural History, Washington D.C., and Nancy Voss, Rosenstiel School of Marine and Atmospheric Science, and their expertise dur ing the examination of material and the preparation of this manuscript. Heather Judkins acknowledges the College of Marine Science at the University of South Florida for the opportunity to pursue her scientific goals. Clyde Roper acknowledge s with appreciation the asso ciation with and support of Mote Marine Laboratory as an adjunct scientist.

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86 REFERENCES 1. Judkins, H, M. Vecchione, & C. F. E Roper.In press. Cephalopods (Mollusca: Cephalopoda) of the Gulf of Mexico. Gulf of Mexico Origin, Waters and Biota Biodiversity, Volume 1 (Darryl L. Feld er and David A. Camp, editors, Texas A&M University Press, College Station, Texas). 2. Lu, C. C. 1977. A new species of squid, Chiroteuthis acanthoderma, from the Southwest Pacific (Cephalopoda: Chirot euthidae). Steenstrupia, Zoological Museum of University of Copenhagen; 4:179-188. 3. Nesis, K. N. 1980. Taxonomic Position of Chiroteuthis famelica Berry (Cephalopods, Oegopsida). Byulleten Moskovskogo Obsestva Ispytatelei Prirody (Otdel Biologicheskii); 85 (4): 59-66. ------. 1982. Cephalopods of the World, Squids, Cuttlefishes, Octopuses, and Allies. T.F.H. Publications, Inc. 385 pp. 4. Roper, C. F. E. and R. E. Young, 1975. Vertical Distribution of Pelagic Cephalopods. Smithsonian Contributions to Zoology, no. 209: 51pp. -----& C. C. Lu, 1990. Comparative Morphology and Function of Dermal Structures in Oceanic Squids (Cephalopoda). Smith sonian Contribution to Zoology, no. 493: 41 pp. 5. Tsuchiya, K. and T. Okutani. 1993. Rare and interesting squids in Japan, recent occurrences of big squids from Okinawa. Venus, 52 (4): 299-311. 6. Young, R. E., 1991. Chiroteuthid and rela ted paralarvae from Hawaiian waters. Bulletin of Marine Science, 49(1-2): 162-185. -----& C. F. E. Roper. 1999. Asperoteuthis acanthoderma (Lu, 1977). Version 01 January 1999. (under construction). http://tolweb.org/Asperoteuthisacanthoderma/19466/1999.01.01 in the Tree of Life Web Project, http://tolweb.org/ ------, M. Vecchione, & C. F. E. Roper. 2007. A new genus and three new species of decapodiform cephalopods (Mollusca: Cephalopoda). Review of Fish Biology and Fisheries, 17:353-365.

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87 4.8 FIGURE LEGENDS Fig. 1. Key West Asperoteuthis acanthoderma specimen. Fig. 2. Marathon Asperoteuthis acanthoderma specimen. Fig 3. Internal organs of Key West specimen: A. digestive gland B. gill; C. nidamental glands; D. branchial heart; E. stomach; F caecum; G. ovary. Manuscript Note.---After the manuscript was completed, we received information that an additional specimen of Asperoteuthis had been found floating at the surface by a fisherman five miles south off Little Cayman Island in the northern Caribbean Sea on 18 May 2008. It was donated to the Little Caym an Research Center for identification by station manager, Jon Clamp, who in turn, contacted us for positive identification via photographs. The damaged specimen had a mantle length of approximately 152.4 cm when collected. The specimen will be deposit ed at the Smithsonian Institution along with the Key West specimen once additional stud ies have been conducted. We thank Jon Clamp, Judie Clee, and the fisherman who di scovered the specimen for their efforts to insure that the specimen was made av ailable for study and permanent deposition.

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88 Table 4.1. Measurements of specimens (mm). Key West Marathon Sex Female Female Mantle Length: 620 1630 Total Length (existing portions): 1817 3420 Mantle width: 190 210 Head length : 230 Head width: 35 50 (eye to eye) Funnel valve length: 55 n/a (base-tip) Funnel valve-base width: 60 n/a Fin length (primary): estimated 220 460 Total fin length: n/a 610 Fin width primary: 340 330 Secondary fin width: n/a Tentacle length: R: 100 + L: n/a R: 1560+ L: 780 Club length: n/a n/a Arm length I: R L R L 880(ti) 655 845 810 Arm length II: 895 680 1030 970 Arm length III: 1000 897 760+ 1100

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89 Arm length IV: 560 870(ti) 520 825 Sucker diameter I: 1st 30cm tip 1st 30cm tip (right) 4 6 2 5 6 0.7 Sucker diameter IV: 4 4 1 4.5 4.5 4.5 (left) Eye diameter: 28 n/a Ti= to tip, complete; + = feature incomplete/broken

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90 Figure 4.1. Key West Asperoteuthis acanthoderma specimen.

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91 Figure 4.2. Marathon Asperoteuthis acanthoderma specimen.

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92 Figure 4.3. Internal organs of Key West specimen: A. digestive gland B. gill; C. nidamental glands; D. bran chial heart; E. stomach; F caecum; G. ovary.

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93 CHAPTER 5 CONCLUSIONS Summary The cephalopods of the Broad Caribbean are widely distributed with concentrations along the continen tal shelves. There are gaps in the collections from the western Gulf of Mexico and the Caribbean S ea, both nearshore and offshore, which are important to investigate further. Distribu tion maps (Appendix C) display a wide variety of localities that the cephalopods occ upy with island nations hosting many octopod species. According to the present study, Ra poports Rule does not apply to the cephalopods of the Broad Caribbean. This fi nding is supported by a literature survey conducted by Rosa in 2008 (Rosa et al., 2008) de scribing species richness of cephalopods along the Atlantic coast. Cephalopod species richness increased with increasing latitude contrary to Stevens (Stevens, 1989) belief that species richness increases towards the equator. The search for diversity hotspots of the Broad Caribbean was not conclusive in most instances as the sample sizes were too small to allow the rarefaction curves to reach asymptote. According to the analysis, region 4, the eastern coast of Florida exhibited the highest species richness (n=32). Species richness may be higher here because of increased nutrient mixing in the Gulf Stream, patterns of current transport in the region and presence of both shallow a nd deep water habitats existing along the eastern Florida coast Collection intensity could have b een an attributing factor to

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94 increased richness although the majority of th e rarefaction curves were headed towards an asymptote. The low sample size in the so utheastern Caribbean S ea (n=4) displays the need for increased study to enrich the ce phalopod knowledge base in that region. The Gulf of Mexico species checklist exposes the advantages and pitfalls of creating species checklist studies today based on literature alone. Th e literature provided an excellent collection of species that ar e found within the regi on, dating back to the 1800s. However, when it is compared to the actual data collected in chapter 2 of this study, discrepancies were discovered. Table 5.1: Gulf of Mexico Checkli st Species not included in Ch II: 1. Brachioteuthis sp. 10. Chiroteuthis veranyi 2. Leachia cyclura 11. Chiroteuthis mega 3. Cycloteuthis sirventyi 12. Pholidoteuthis boschmai 4. Discoteuthis laciniosa 13. Octopoteuthis sicula 5. Histioteuthis bonnelleii 14. Mastigoteuthis magna 6. Chtenopteryx sicula 15. Octopus mercatoris 7. Lepidoteuthis grimaldii 16. Pteroctopus schmidti 8. Lampadioteuthis megaleia 17. Ocychoe tuberculata 9. Chiroteuthis joubini Seventeen species (Table 5.1) were found in the Gulf of Mexi co checklist were not found in the biogeography study described in chapter 2. If the biogeographical results are combined with the checklist, the result is a total of 127 cephalopod species that are found in the Broad Caribbean region. When the species from Vecchiones work (2002) is merged in, a total of 131 species exist in the Broad Caribbean. Although the study presented a few discrepancies in the Gu lf of Mexico species totals, the overall picture is one of large species diversity throughout the region. A factor to consider is that some of the checklist species are based on one or two specimens which were badly

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95 damaged animals. The paucity of species samples demonstrates the need to investigate the Broad Caribbean thoroughly for additi onal samples to ascertain the validity, abundance, and range limits of species di stribution. The 18 range extensions reported here are of value for understanding how cephalop ods fit into the ecosystems of this area. Many facets are needed to create the comple te picture in terms of marine ecosystems and the role that the cephalopods play in them. Neither data nor literature alone will complete the picture for any diverse marine group. The deep-dwelling squid, Asperoteuthis acanthoderma was an exciting new find for the Broad Caribbean and illustrates how lit tle is known about the distribution of deepwater cephalopods. The species was only found off the coast of Japan prior to this discovery. Further research may reveal that A. acanthoderma s distribution is circumglobal. To find and describe a new sp ecies of large squid af ter extensive studies have been conducted in the region (G. Vo ss 1956, Lipka 1975, Nesis 1975, Roper et al. 1984, N. Voss et al. 1988, Passarella, 1991, V ecchione 2002) reveals the likelihood of finding new cephalopods in the Broad Caribbean. A recent paper (Norman & Hochberg, 2005) examined the status of octopods worldwide. There have been new discoveries in many regions of the world including New Zealand, New Caledonia, South Africa, and Hawaii. There is an obvious absence of octopod findings in the Broad Caribbean regi on. G. Voss (1977) calculated the projected amount of octopod species that would be found on an annual basis, and suggested that an average of 6.7 species being described annually worldwide. He also stated that it seems clear that we are still in the descriptive stag e of systematics. These statements seem to hold true over 25 years later (Norman & Hoc hberg, 2005). The study went on to describe

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96 the progress and impediments to taxonomic studies today. Progress is visible in advanced electronic communica tion, better technology, and bette r international links. Areas that are in need of improvement: poor retention of materials, few replicates, poor curation, little support for taxonomic resear ch, very few understudies to continue progress, and museum curators are not being replaced as they retire, collections being maintained but not active or growing, and lastly fewer primary field studies are being undertaken (Norman & Hochberg, 2005). Proper identification and collection will also provide better understanding of the potential for cephalopod fisheries. Over 50% of the global cephal opod catch recorded by the Food and Agricultural Organization (FAO) is not segregated into single species categories, and this significantly reduces any value the data ma y have for population assessment (Boyle & Boletzky, 1996) Collabor ation between scien tists throughout the Caribbean would further quantify and identi fy important and scarcely known species. Improved collection methods and new species specific research cruises are sorely needed to discover what else lies in the deep er waters of the Caribbean Sea and Gulf of Mexico. There is a demand for the information as finfish harvests collapse, the fisheries will turn to cephalopods for commercial expl oitation. An immediate priority for the ocotopods is production of detaile d and accurate descriptions of all species as the octopod taxonomy needs major revision and stab ilization (Norman & Hochberg 2005) Bottom depth and vertical migration can be important elements in the cephalopod lifestyle as many (e.g. Selenoteuthis scintillans, Spirula spirula ) ascend nightly for feeding (Roper and Young, 1975). Deep-sea cepha lopod collection has be en traditionally difficult (Wormuth & Roper, 1983). Also, connections in food webs are sometimes

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97 difficult to determine as the cephalopods need to break down their pr ey into small parts prior to digestion and stomach contents ar e therefore difficult to identify. Vertical migration of cephalopods and the role they pl ay in the transfer of energy between the layers are not well known. Less than half of the specimens with known locales of the 5190 examined had depth of collection recorded. The depth information available was not adequate to describe specific depth ranges for the cephalo pods in the region. Another challenge in this study was the la ck of detail on capture locations for many specimens. Many had only regional information associated with the sample and over 200 specimens were not used due to a complete absence of information (e.g. collected in the Caribbean Sea). A universal standard for recording sample information throughout the Broad Caribbean could impr ove sample details and would glean consistent accuracy for research studies. The sample informati on should include the following: latitude/longitude, date, time, de pth, water temperature, identification (if possible) and collection method used for capture. A technique that has been successful in solidifying relationships between cephalopod species is DNA analysis. An exam ple of an organism to examine in the Broad Caribbean for genetic relatedness could be Doryteuthis plei and determine the genetic linkages between the northern and southern populations. Also, DNA comparisons with cephalopod species from the eastern tropical Paci fic could lead to insights concerning the origins of the fauna. Past geologi cal occurrences provide an excellent backdrop to examine the or igins for cephalopods in the region. There have not been any previous comp rehensive cephalopod studies of the Broad Caribbean since the early work of Voss (1956) and there is little l iterature from other

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98 similar-sized regions for comparison. There are many smaller scaled studies, and it may be because cephalopod taxonomy is in flux that large biogeography studies have been neglected. Cephalopods have been the focu s of many studies in the Broad Caribbean region and the results from the present rese arch paper build and highlight additional advances in the comprehensive overview of the group. Much work is still needed to attain a better understa nding of this complex cephalopod assemblage.

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99 REFERENCES 1. Boyle, P.R. & Boletzky, S.V. 1996. Cephalopod populations, definition, and dynamics. Philosophical Transactions of the Royal Society, London. Series B. Biological Sciences; B351:985-1002. 2. Lipka, Douglas. 1975. The systematics and zoogeography of cephalopods from the Gulf of Mexico. Dissertation, Texas A&M. August 1975; 1-347. 3. Nesis K.N. 1975. Cephalopods of the American Mediterranean Sea; English Translations of Selected Publica tions on Cephalopods; Editor M. Sweeney; Smithsonian Institutions Libraries; 1; 318358. 4. ODor, R.K., J.A. Hoar, D.M. Webber, F.G. Carey, S. Tanaka, H.R. Martins and F.M. Porteiro. 1994. Squid ( Loligo forbesi ) performance and metabolic rates in nature. Physiology of Cephalopod MolluscsLife style and Performance Adaptations. Gordon and Breach Publishers. 163-177. 5. Passarella, K.C., 1990. Oceanic Cephalopod Assemblage in the Eastern Gulf of Mexico Department of Marine Science University of South Florida: St. Petersburg, Fl. 50pp. 6. Roper, Clyde F. E. and Richard E. Y oung. 1975. Vertical distri bution of pelagic cephalopods. Smithsonian Contri bution to Zoology; 209; 1-51. 7. Roper C.F.E, M.J. Sweeney & C.E. Nauen, 1984. FAO species catalogue. v.3 Cephalopods of the world. An annotated a nd illustrated catalogue of species of interest to fisheries. FAO Fish. Synop. (125) v3: 277p. 8. Rosa, Rui, Heidi M. Dierssen, Liliana G onzalez, and Brad Seibel. 2008. Ecological biogeography of cephalopod molluscs in the Atlantic Ocean: historical and contemporary causes of coastal dive rsity patterns; Global Ecology and Biogeography; 17: 600-610. 9. Semmens, Jayson M., Gretta T. Peel, Br onwyn M. Gillanders, Claire M. Walunda, Elizabeth K. Shea, Didier, Jouffre, Taro Ichii, Kartsen Zumhotz, Oleg N. Katugin, Stephen C. Leporati and Paul W. Sh aw. 2007. Approaches to resolving cephalopod movement and migration pattern s. Review of Fish and Biological Fisheries; 17: 401-423. 10. Stevens, G.C., 1989. The Latitudinal Gradient in Geographical Range: How So Many Species Coexist in the Tropics. The American Naturalist; 133(2): 240-256.

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100 11. Vecchione, 2002. Cephalopods. Carpenter, K.E.(ed.) The living marine resources of the Western Central Atla ntic. Volume 1: Introduction, molluscs, crustaceans, hagfishes, sharks, batoid fishes, and chimaeras. Cephalopods of Western Central Atlantic; FAO Species Identification Guide for Fisheries Purposes and American Society of Ichthyologists and Herpetologi sts Special Publication No. 5. Rome, FAO. 2002. 1-600. 12. Voss, G.L. 1956. A review of the cephalopo ds of the Gulf of Mexico. Bulletin of Marine Science: Gulf and Caribbean; 6:85-178. 13. Wormuth, John H. and Clyde F.E. Roper. 1983. Quantitative sampling of oceanic cephalopods by nets: problems and recomm endations. Biological Oceanography; 2 (2-4) 357377.

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101 APPENDICES

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102 Appendix A: Regional Dist ribution Point Reference Regional Distribution Points Regional Number Region Lat (N) Long (W) 1 NE Florida 29.26 -80.76 2 SE Florida 26.41 -79.65 3 Florida Keys 24.81 -80.58 4 SW Florida 26.29 -82.31 5 Bahamas 24.5 -76.19 6 West Cuba 22.77 -82.06 7 SE Cuba 20.42 -76.06 8 Dominican Republic 19 -70.94 9 Jamaica 18.09 -78.19 10 Puerto Rico--> Antigua 18.35 -64.92 11 Guadeloupe->Martinique 15.38 -61.24 12 St. Lucia--> Grenada 13.32 -61.24 13 Trinidad-->East Venezuela 11.31 -61.83 14 Columbia 9.47 -76.22 15 Panama-->Costa Rica 9.28 -81.39 16 MexicoYucatan area 21.47 -86.46 17 Belize 17.25 -88.43 18 Texas/ North Mexico 26.35 -96.51 19 Louisiana 29.57 -88.02 20 NW Florida 27 -83.63

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103 Appendix B: Species Richness Comparison Quadrats Species Richness Comparison Quadrats Area Region Latitude Longitude Area 1 Mississippi River Delta 28.3830.28N 86.4490.8W Area 2 West Central Florida 25.5628.38N 82.1284.12W Area 3 Straits of Florida 23.14-25.2N 79.982.17W Area 4 Eastern Central Florida 25.5628.33N 78.4880.32W Area 5 Mid Caribbean Islands 14.4218.04N 61.1264.35W Area 6 Southeast Caribbean Sea 10.4112.26N 61.1264.35W Area 7 South Central Caribbean Sea 11.1213.01N 68.571.44W Area 8 Southern Caribbean-Colombia region 8.30-11.28N 75.1978.27W

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146 ABOUT THE AUTHOR Heather Judkins received her Bachelors Degree in Marine Affairs from the University of Rhode Island in 1993. She moved to Florid a and received her Masters Degree in Secondary Science Education while teaching hi gh school marine science classes. In 2003, she entered the Marine Science graduate program through the University of South Florida pursuing a biologi cal track focusing on cephal opods. While enrolled, she received two fellowships through the GK12 Oceans Program which was sponsored by the National Science Foundation. She also c oordinated the Regional National Ocean Sciences Bowl, the Spoonbill Bowl, as a result of her fellowship project from 2004present. She has continued teaching full time while tackling her degree. She has coauthored two publications regarding cepha lopods of the Broad Caribbean to date.

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