Smithsonian at the Poles: Contributions to International Polar

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PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 163 TABLE 8. Endemic species of Paraeuchaeta and Heterorhabdidae found in four different areas of interest. A “�” indicates presence. Area of interest Species Southern Ocean Arctic-boreal Eastern Pacifi c Indo-West Pacifi c Paraeuchaeta antarctica � P. austrina � P. biloba � P. dactylifera � P. eltaninae � P. erebi � P. parvula � P. rasa � P. similis � P. tycodesma � P. birostrata � P. brevirostris � P. elongata � P. glacialis � P. norvegica � P. polaris � P. rubra � P. californica � P. copleyae � P. grandiremis � P. papilliger � P. eminens � P. investigatoris � P. russelli � P. simplex � Heterorhabdus austrinus � H. pustulifer � H. spinosus � H. paraspinosus � Heterostylites nigrotinctus � Paraheterorhabdus farrani � Heterorhabdus fi stulosus � H. norvegicus � H. tanneri � Paraheterorhabdus longispinus � Heterorhabdus abyssalis � H. americanus � H. prolixus � H. quadrilobus � Heterostylites echinatus � tribution within the Southern Ocean, but only P. antarctica has been collected in large numbers. The endemic species of the Arctic Ocean, including adjacent boreal waters, and the endemics of the eastern Pacifi c have also been found in large numbers. These species are all believed to inhabit waters of high primary and secondary productivity, where endemism may have developed as an adaptation to these eutrophic habitats (Park, 1994). Of the four endemics of the Indo-West Pacifi c, Paraeuchaeta russelli and P. simplex are neritic, inhabit- ing relatively shallow water. Paraeuchaeta eminens and P. investigatoris are deepwater species. All four species are common in waters of the Malay Archipelago, an area with relatively high primary and secondary productivity. High primary and secondary productivity, rather than a habitats abiological attributes, appears to have been the primary determinant for the evolution of endemicity among these species of Paraeuchaeta. Within the family Heterorhabdidae, six species are endemic to the Southern Ocean as compared to fi ve endemic
164 SMITHSONIAN AT THE POLES / PARK AND FERRARI species in the eastern Pacifi c. Four species are restricted to the Arctic-boreal area, including three species endemic to the boreal Pacifi c and one endemic species found in the boreal Atlantic. No endemic species of Heterorhabdidae is found in the Indo-West Pacifi c. All of the heterorhabdid species discussed here are assumed to be carnivores, with the exception of Heterostylites echinatus (see Ohtsuka et al., 1997), and carnivory is assumed to have arisen from suspension feeding within the Heterorhabdidae only once (Ohtsuka et al., 1997). The highest number of endemic heterorhabdid species, like the number of endemics of Paraeuchaeta, is found in the Southern Ocean. However, in contrast to species of Paraeuchaeta, of the six Southern Ocean endemic heterorhabdids, only Heterorhabdus spinosus is found in large numbers; it is common in coastal waters. Beyond the Southern Ocean, among the four endemic species of the Arctic and boreal seas, Heterorhabdus norvegicus is very common in the boreal Atlantic. Heterorhabdus fi stulosus, H. tanneri, and Paraheterorhabdus longispinus are very common along the coasts of the boreal Pacifi c, and all occur in large numbers in some localities. Among the fi ve endemic species of Heterorhabdidae in the eastern Pacifi c, all are limited in their distribution to waters close to the coasts of Americas, in areas of coastal upwelling, where they have been found in large numbers. One explanation for the different occurrences of Paraeuchaeta and heterorhabdid endemics in the Southern Ocean is that the heterorhabdids may be relatively late colonizers of the Southern Ocean; species of Paraeuchaeta may already have established themselves as the dominant carnivores before colonization of the Southern Ocean by species of Heterorhabdidae. In summary, within the two families of pelagic calanoids that have been studied worldwide, the highest number of endemic species is found in the Southern Ocean. All Southern Ocean endemics of Paraeuchaeta are found in large numbers, except for P. similis and three species found near the ice edge adjacent to Antarctica where these ice edge species may have been undersampled. In contrast, the six endemic species of Heterorhabdidae found in the Southern Ocean are rare. However, beyond the Southern Ocean the endemics of both families of carnivores are associated with the waters of high primary and secondary productivity, where they may be collected in large numbers. On the basis of these observations, the endemicity of many of the very common bathypelagic calanoids of the Southern Ocean, like the endemicity of Southern Ocean epipelagic calanoids, is suggested to have resulted from the adaptation to conditions of high primary and secondary productivity. EVOLUTION OF THE PELAGIC CALANOID FAUNA WITHIN THE SOUTHERN OCEAN Among the 184 deepwater calanoid species found in the Southern Ocean, 50 species (27%) occur exclusively in the Southern Ocean; 20 of those species (40%) are rare or very rare (Table 3). Several factors may be responsible for the evolution of the deepwater pelagic calanoid fauna, restraining their dispersal throughout the deepwater of the world’s oceans and selecting for this endemicity. Water temperature, as represented by the rather abrupt changes at the Antarctic Convergence or the Subtropical Convergence, is unlikely to affect the structure or the distribution of the deepwater calanoids because water temperatures below 1,000 m are uniformly cold within the Southern Ocean, and this uniformly cold deepwater is continuous with the deepwater of the adjacent Pacifi c, Atlantic, and Indian oceans. The proposed relationship between habitat productivity and endemism may be a more useful initial condition. The majority (60%) of deepwater endemic species of the Southern Ocean are common or very common. As mentioned earlier, this may be the product of the high primary and secondary productivity of the Southern Ocean, especially south of the Antarctic Convergence, resulting in the evolution and adaptation of an oligotrophic species to this eutrophic habitat. Endemism of deepwater pelagic calanoids in the Southern Ocean, therefore, is hypothesized to have evolved as rare species that are widely distributed in oligotrophic habitats throughout the world’s oceans became adapted to exploit high primary and secondary productive habitats (Park, 1994); these adaptations have resulted in an increased population size of the eutrophic species. A second explanatory condition for the evolution of the pelagic, marine calanoid fauna in the Southern Ocean depends on whether polar species within a single genus are monophyletic, having evolved from a single ancestral species that initially colonized the Southern Ocean, or polyphyletic, with each species having evolved independently from an ancestor distributed outside of the Southern Ocean or by evolving from more than one initially colonized ancestral species. There is evidence that supports this latter model of independent colonizations for Southern Ocean endemic species of the families Euchaetidae and Heterorhabdidae (see the Comparative Endemicity of the Southern Ocean Fauna section), although the situation is more complex for the antarctica species group of Paraeuchaeta. Further evidence can be found in the phenomenon of sibling species pairs. When morphological details are closely compared, one species often can be found outside the Southern Ocean that is very similar to each Southern
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Smithsonian at the Poles Contributi
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Contents FOREWORD by Ira Rubinoff i
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Elaina Jorgensen, Alaska Fisheries
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CONTENTS vii Watching Star Birth fr
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x SMITHSONIAN AT THE POLES blooms i
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xii SMITHSONIAN AT THE POLES Change
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xiv SMITHSONIAN AT THE POLES Museum
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Advancing Polar Research and Commun
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James P. Espy (1785- 1860), the fi
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ology fueled hopes that the scienti
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Meteorologists also provided critic
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visualize weather patterns remotely
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in ice sheets. The latest collapse
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Cooperation at the Poles? Placing t
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British Association for the Advance
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cations, in which the Smithsonian s
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ence” (Robinson, 2006: 76), he ha
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Taylor, C. J. 1981. First Internati
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24 SMITHSONIAN AT THE POLES / KORSM
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36 SMITHSONIAN AT THE POLES / De VO
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From Ballooning in the Arctic to 10
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ern Svalbard in 1896 (Capelotti, 19
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FIGURE 2. The Andrée campsite in 1
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National Zoo in Washington, D.C.—
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FROM BALLOONING TO 10,000-FOOT RUNW
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e rapidly deployed to South America
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“Of No Ordinary Importance”: Re
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1942). Ethnological collecting had
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group. More importantly, Murdoch’
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gan to study the question of Indian
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small museums and culture centers i
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fer of Alaskan objects and informat
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fi rst time in polar research— di
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Boas, Franz. 1888a. “The Central
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Lindsay, Debra. 1993. Science in th
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80 SMITHSONIAN AT THE POLES / FIENU
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90 SMITHSONIAN AT THE POLES / BURCH
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100 SMITHSONIAN AT THE POLES / CROW
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Page 208 and 209: ing such cryptic speciation suggest
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214 SMITHSONIAN AT THE POLES / WINS
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Considerations of Anatomy, Morpholo
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Many theories have been hypothesize
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FIGURE 4. A three dimensional recon
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are wider and more bulbous in the a
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mimicked the shape and profi le of
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faces for SEM observation. The spec
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DISCUSSION Imaging and dissection o
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out its length. The pulp chamber al
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pulpal neurons and dentin tubules.
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Scientifi c Diving Under Ice: A 40-
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TABLE 2. Principal Investigators an
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attaching ice anchors to the chunks
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dumping of weight under water. The
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MARINE LIFE HAZARDS Few polar anima
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Urine should be copious and clear a
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Environmental and Molecular Mechani
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face (�1.8°C) are likely to pose
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FIGURE 2. Illustrated selection shi
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FIGURE 4. Distribution of respirati
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invertebrates were placed in the ba
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Meehan, R. R., D. S. Dunican, A. Ru
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266 SMITHSONIAN AT THE POLES / KOOY
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272 SMITHSONIAN AT THE POLES / DUNT
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286 SMITHSONIAN AT THE POLES / QUET
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300 SMITHSONIAN AT THE POLES / NEAL
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310 SMITHSONIAN AT THE POLES / SMIT
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320 SMITHSONIAN AT THE POLES / KIEB
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Capital Expenditure and Income (For
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tal breeding systems (Boyd, 1998; T
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FIGURE 3. Pup mass gain in relation
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MONITORING FOOD CONSUMPTION DURING
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primary productivity in McMurdo Sou
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Non-steady-State Systems. Journal o
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Latitudinal Patterns of Biological
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and perhaps others, may have invade
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colonizing the Arctic Ocean under c
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due in part to the great distances
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and (2) human-mediated responses to
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Lewis, P. N., M. Riddle, and C. L.
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Cosmology from Antarctica Robert W.
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There they found better observing c
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TELESCOPES AND INSTRUMENTS AT THE S
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Gaier, T., J. Schuster, and P. Lubi
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J. M. Kovac, C. L. Kuo, A. E. Lange
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370 SMITHSONIAN AT THE POLES / MART
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372 SMITHSONIAN AT THE POLES / MART
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374 SMITHSONIAN AT THE POLES / WALK
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Watching Star Birth from the Antarc
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STAR BIRTH FROM THE ANTARCTIC PLATE
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is constructing and deploying the P
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Antarctic Meteorites: Exploring the
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at the Field Museum, and pieces wer
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the description and curation of Ant
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led these committees throughout the
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Index AAUS. See American Academy of
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temperature, 350-355 tourism, 354 B
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Fishing. See also Biological invasi
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McMurdo Station, xiv, 265-267, 393
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Rogick, Mary, 206 Ronne, Finn, 55 R
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U.S. Naval Support Force Antarctica
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