Deep-Sea Pycnogonids and Crustaceans of the Americas Michel E

Deep-Sea Pycnogonids and Crustaceans of the Americas Michel E. Hendrickx Editor

Deep-Sea Pycnogonids and Crustaceans of the Americas Editor Michel E. Hendrickx Inst Ciencias del Mar y Limnología National Autonomous University of Mexico Mazatlán, Mexico

ISBN 978-3-030-58409-2 ISBN 978-3-030-58410-8 (eBook) https://doi.org/10.1007/978-3-030-58410-8

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This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland This book is the result of a large effort by many American scientists and some of their overseas colleagues. Participation of academics from Latin America is particularly important as it throws new light on the deep-sea communities occurring off their countries’ coasts or, in some cases, in much larger areas within the geographic limits of the Americas. In this context, we wish to dedicate this book to two of our colleagues who actively participated in the early edition of this book but passed away while this process was taking place.

Our Chilean colleague, Marco Antonio Retamal, was certainly a founding architect for the investigation in marine biology in Chile. Local expert in crustaceans, Marco Antonio was involved in a wide series of projects in his homeland, successfully forming and advising many students and young scientists during his career.

Jorge López, from El Salvador, shared his interest between the fshery activities in Central America and a genuine, strong desire of studying the deep-water fauna in the region. An expert in fsheries and conservation issues, he often participated in regional meetings and local advisory panels. Marco Antonio and Jorge were not only distinguished academics with a profound interest in the biology and ecology of crustaceans and deep-water communities in Latin America, they were also dear friends to many of us. Their contribution to this book will remain forever a contribution to our knowledge of the deep-sea fauna of the Americas. Both true gentlemen, they will be remembered with respect, affection, and admiration. Preface

Biogeography is a synthesis science integrating all the knowledge from geography, climatology, paleontology, zoology, and phylogenetic. The frst step to obtaining all this knowledge is a good sampling. Taxonomy is also essential to compare species between geographies. Nowadays, the knowledge acquired from diverse sources is collected in computer databases, and the largest ones in zoology and marine ecology are OBIS and WoRMS. However, before it can be integrated inside these databases, regional revisions are necessary to clarify the data. During the last decades, knowledge about the Arthropoda from the deep sea, especially Pycnogonida and Crustacea, increased considerably. The increase in marine deep-sea cruises with a mixture of boats, manned submersibles, or even ROVs brought several new species to light and more information about the way of life of these animals. Surprisingly, some areas on the planet remained very poorly known. It is the case for the coast of the Americas, more precisely South America, where a large gradient of latitudes offers a great diversity of ecosystems. The present volume is a very successful attempt to fll the gap in the knowledge on America’s crustacea. Crustaceans are present in all ecosystems and at any depths, from the pelagic to the benthic zones, and play a critical role in the food web. There are scavengers such as amphipods or isopods, plankton eaters like copepods, sus- pension feeders like cirripeds, and even carnivores such as lobsters or hermit crabs. At any given place in the oceans, these species they are essential to the functions of the ecosystems. The American continent, which extends from the Arctic to the Antarctic waters, separates the fauna between the Atlantic Ocean and the Pacifc Ocean. Very few benthic species, if any, can be found on both sides of the Americas. Furthermore, it is still unclear why the west American coast fauna is so different from the Indo- Pacifc one. The closing of the Isthmus of Panama, only 2.8 million years ago, explains partially this point as well as the long distances and strong currents separat- ing the East from the West Pacifc Ocean. Surprisingly, only 120 deep-sea species of decapods are known from the pacifc coast of the Americas considering that we fnd more than ten times this number in the Western Pacifc. Perhaps, the hypoxic conditions found along a large part of America’s coastlines, with the oxygen

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minimum zone found within a large bathymetric range, could well be responsible for this poor fauna diversity. Besides the scientifc value, this kind of “regional” cataloguing of crustacea is important for the history of marine biology, drawing up a list of oceanographic vessels and expeditions undertaken along the American coasts. It is also an opportunity to acknowledge the dedicated work of community experts such as taxonomists or ecologists. Some articles are very original and precious considering that they describe very light and fragile animals diffcult to discover. They were collected in the deep sea, near the bottom or inside of the upper layer of the sediments, which required the use of sledge-dredges with micrometric meshes or box-corer. The catalogue of Pycnogonida from the Uruguayan continental slope has been enriched with several new deep-sea species. The groups of peracarid crustacea are insuffciently sampled anywhere in the world. It is well known that the harpacticoid copepods are very rich in species, present in all the benthic habitats and sometimes forming huge biomass. However, surprisingly, the review of the species from the Americas is only comprised of 22 families and 169 species. For the larger peraca- rids animals, the knowledge is a little bit better: 27 families and 118 species of amphipods for western Mexico only. The biology of the pelagic species living in the largest ecosystems of the world is still poorly known. Therefore, the observations about the reproduction and bathymetric repartition of Nematocarcinu species in the Gulf of California are very useful. This group is present in all the oceans, characterized by their very long legs. For the frst time, the sex ratio has also been evaluated. A large sampling in the Gulf of Mexico by numerous oceanographic cruises brings good data on the 4 families and 14 species of Penaeoid shrimps. On the other side of South America, different cruises have collected 78 species of pelagic shrimps, Dendrobranchiata and Pleocyemata, between Peru and Tierra del Fuego. A large part of these is associated with the Nazca and Salas y Gomez Ridge sea- mounts. The lobsters are also included with the Nephropidae and Polychelidae of the Gulf of Mexico. The very large group of Galathoidea, one of the best-studied group of crustaceans in the world, has been studied on the Brazilian coast with the help of molecular tools. Fisheries in the deep sea are problematic. Generally, the growth rate of the tar- geted species is unknown as well as the generations turnover. In the case of crustacea, there is another diffculty: the halieutic models are validated for fsh, with linear growth, and not for crustacea with several moults. Studies on Lithodids from the Peruvian coast are bringing new useful data on species of Parolomis and Lithodes. In several places, the deep-sea populations of crustacea are already being overexploited. Other chapters bring interesting comments on decapod fsheries concerning 79 species in Uruguay and 181 species along the Brazilian coast, mainly on shrimps and geryonid crabs. For each species, an updated synonymy is given. The sampling of deep-sea crustacea along the Chilean coast is showing a change in the diversity between the warm waters in the north and sub-Antarctic waters south of Chiloé Preface ix

Island brought by the Humbolt Current. The pacifc coast of Central America was also sampled from Guatemala to Panama. Conservation strategies are proposed to keep an equilibrium between healthy ecosystems and sustainable fsheries in the deep sea. In these environments, the biological conditions are different from the shallow-water fsheries, and the stocks are easily overexploited because the species have extreme longevity, slow growth rates, late maturity, and low fecundity. This volume highlights the ambitious exploration sampling programs in the deep sea to increase our knowledge of the fauna from North, Central, and South America. This knowledge could help to protect the deep-sea environment essential for the sustainability of our planet.

MNHN-Nouméa Bertrand Richer de Forges Paris, France

Prologue

Although investigation in the Latin America part of the Americas’ deep waters has signifcantly increased in the last decades, there is still a profound gap between what we know about this environment in most of the northern hemisphere and in the southern hemisphere. Exploratory cruises fnanced by local governments are few, and occasional bilateral projects, although often well fnanced and organized, have not been suffcient to signifcantly increase the rate at which deep-water communities are sampled and studied. On the other hand, the number of scientists who have found opportunities to get involved in this kind of research is extremely reduced due to a certain lack of priority for this category of science. Globally, the number of large, modern sea-going vessels available for research is also very reduced in several countries in the region. Operational costs are extremely high and, in some cases, fnding resources to support offshore surveys is near to impossible. Some research vessels are quite old and should ideally be replaced by more effcient vessels, but, again, there are often other priorities and critical investments that are needed and focused on more urgent situations. Equipment is another issue, and prices of modern sampling gear and recording equipment are most of the time too high for local budgets. In some countries, a strong interest has arisen for deep-water fshing and some of our colleagues have had the opportunity of taking advantage of this sampling effort to gather valuable information on deep-water communities that were previously mostly unknown. This, of course, raises the question of deciding whether the deep-water fshing industry should be given a “carte blanche” or be supervised in order to limit its impact on the poorly known, slow-growing species that might prove to be commercially exploitable on the short term but might be profoundly affected on the long term. Remote equipment have been intensively deployed in regions where institutions or government agencies can afford the acquisition of these expensive equipment and support their offshore operation. Long-term video recording from deep-water observatories is certainly a friendly way to study the local fauna, provided we do have a fair knowledge of what species inhabit the area. These low-impact techniques will certainly proliferate in the near future, but ele- vated cost to build and maintain these infrastructures obviously represent a huge limitation for many countries. One of the purposes of this book is to call the attention

xi xii Prologue to the extraordinary richness of the deep-sea fauna in our region. Simultaneously, it is a warning that what we know about this fauna, its distribution, its basic biology, and its ecology is far from being suffcient in order to understand and manage all the factors that are already impacting negatively on these communities. In fact, they could very well vanish before we are able to study all its components and their inter- action. The challenge is enormous and tantalizing. To be successful, it will take a gigantic effort, a great deal of investment, and, above all, comprehensive multina- tional collaboration programs aimed at sharing our present experience and our future goals. Contents

1 The Deep-Water Crustacean and Pycnogonid Fauna of the Americas in a Global Context �� 1 K. E. Schnabel, S. T. Ahyong, A. J. Gomez, M. E. Hendrickx, R. A. Peart, and J. N. J. Weston 2 Deep-Sea Pycnogonids from Uruguay: Every Deep Cruise Adds Valuable Information �� 25 A. Soler-Membrives, R. A. Lucena, J. B. Company, and G. Rotllant 3 The Deep-Water Colossendeis tenera Hilton, 1943 (Pycnogonida, Pantopoda, Colossendeidae) off Western Mexico �� 39 Michel E. Hendrickx 4 The Deep-Water Benthic Harpacticoida (Copepoda) of the Americas �� 47 S. Gómez and K. I. Rivera-Sánchez 5 Updated Checklist of Deep-Sea Amphipods (Amphilochidea and Senticaudata) from Western Mexico, NE Pacific Ocean �� 127 I. Winfeld, Michel E. Hendrickx, and M. Ortiz 6 Isopoda Epicaridea from Deep Water Around North and Central America �� 143 John C. Markham 7 Biodiversity of the Deep-Sea Isopods, Cumaceans, and Amphipods (Crustacea: Peracarida) Recorded off the Argentine Coast �� 157 Brenda L. Doti, Ignacio L. Chiesa, and Daniel Roccatagliata 8 Benthic Invertebrate Communities in the Continental Margin Sediments of the Monterey Bay Area �� 193 John S. Oliver, Kamille K. Hammerstrom, Linda A. Kuhnz, Peter N. Slattery, James M. Oakden, and Stacy L. Kim

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9 Deep-Water Penaeoid Shrimp of the Southern Gulf of Mexico Upper Slope: Distribution, Abundance, and Fishery Potential �� 237 Adolfo Gracia and Ana Rosa Vázquez-Bader 10 Sex Distribution and Reproductive Trends in the Deep-Water Species of Nematocarcinus (Crustacea: Decapoda: Nematocarcinidae) from Western Mexico �� 257 J. C. Hernández-Payán and M. E. Hendrickx 11 Influence of Environmental Variables on the Abundance and Distribution of the Deep-Water Shrimps Nematocarcinus faxoni Burukovsky, 2001 and N. agassizii Faxon, 1893 (Crustacea, Decapoda, Nematocarcinidae) off Western Mexico �� 273 Vanesa Papiol, José C. Hernández-Payán, and Michel E. Hendrickx 12 Pelagic Shrimps (Crustacea, Decapoda, Dendrobranchiata, and Caridea) in the Southeast Pacific �� 293 Guillermo L. Guzman and C. Nicole Olguin 13 Deep-Sea Lobsters (Polychelidae and Nephropidae) from the Continental Slope of the Southern Gulf of Mexico: Distribution and Morphometric Relationships �� 311 P. Briones-Fourzán, E. Lozano-Álvarez, A. R. Vázquez-Bader, and A. Gracia 14 New Molecular Data on Squat Lobster from the Coast of São Paulo State (Brazil) (Anomura: Munida and Agononida) and Insights on the Systematics of the Family Munididae �� 343 I. Miranda, P. A. Peres, M. D. S. Tavares, and F. L. Mantelatto 15 Biology and Distribution of Agononida longipes (Crustacea, Decapoda, Munididae) in the Colombian Caribbean Sea �� 357 Margarita Fierro-Rengifo, Néstor H. Campos-Campos, and Carlos Mario López-Orozco 16 King Crabs of Peruvian Waters During 2003–2004: New Insights �� 375 J. Arguelles, P. Larriviere, S. Thatje, and M. Pérez 17 Lower Slope and Abyssal Benthic Decapods of the Eastern Pacific �� 395 M. K. Wicksten 18 Conservation Strategies for Potential New Deep-Sea Crustacean Fisheries in the Colombian Caribbean under an Ecosystem Approach �� 421 J. Paramo, M. Grijalba-Bendeck, D. Pérez, A. Espinoza-Tenorio, and M. Wolff Contents xv

19 Diversity, Abundance, and Biomass of Deep-Sea Decapod Crustaceans of the Uruguayan Continental Slope in the Southwestern Atlantic Ocean �� 443 G. Rotllant, A. Verdi, R. Santos-Bethencourt, N. Bahamón, and J. B. Company 20 Deep-Sea Megacrustacean Biodiversity (Crustacea, Decapoda) in the South Gulf of Mexico �� 473 A. R. Vázquez-Bader and A. Gracia 21 Catalogue of Typical Deep-Sea Decapod Fauna from Brazilian Waters �� 499 I. Cardoso Azevedo, F. Alves-Junior Almeida, and T. Rodrigues Garcia Almeida 22 An Annotated Checklist and Bibliography of Deep-Water Isopods and Decapod Crustaceans from Chile, Including the Submarine Ridge Salas y Gomez and Nazca Plates �� 585 M. A. Retamal, G. Guzmán, and P. De los Ríos-Escalante 23 Deep-Water Stomatopod and Decapod Crustaceans Collected off Central America by the R/V “Miguel Oliver” �� 603 M. E. Hendrickx and J. López 24 Diversity and Biology of Deep-Water Crustaceans in Costa Rica �� 633 J. C. Azofeifa-Solano and J. Cortés 25 New Records of Crustaceans Collected Between 670 and 3400 m in the Colombian Caribbean �� 659 B. Martínez-Campos, R. A. Plazas-Gómez, H. F. Sáenz, L. F. Dueñas, J. León, and V. Puentes

Index �� 689