Neotropical Bird Migration During the Ice Ages: Orientation and Ecology
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Swarthmore College Works Biology Faculty Works Biology 1-1-1996 Neotropical Bird Migration During The Ice Ages: Orientation And Ecology Timothy C. Williams , '64 Swarthmore College, [email protected] T. Webb III Follow this and additional works at: https://works.swarthmore.edu/fac-biology Part of the Biology Commons Let us know how access to these works benefits ouy Recommended Citation Timothy C. Williams , '64 and T. Webb III. (1996). "Neotropical Bird Migration During The Ice Ages: Orientation And Ecology". Auk. Volume 113, Issue 1. 105-118. https://works.swarthmore.edu/fac-biology/254 This work is brought to you for free by Swarthmore College Libraries' Works. It has been accepted for inclusion in Biology Faculty Works by an authorized administrator of Works. For more information, please contact [email protected]. Neotropical Bird Migration during the Ice Ages: Orientation and Ecology Author(s): Timothy C. Williams, Thompson Webb and III Source: The Auk, Vol. 113, No. 1 (Jan., 1996), pp. 105-118 Published by: American Ornithologists' Union Stable URL: http://www.jstor.org/stable/4088939 Accessed: 27-04-2015 14:44 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/4088939?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Ornithologists' Union is collaborating with JSTOR to digitize, preserve and extend access to The Auk. http://www.jstor.org This content downloaded from 130.58.65.20 on Mon, 27 Apr 2015 14:44:17 UTC All use subject to JSTOR Terms and Conditions The Auk 113(1):105-118, 1996 NEOTROPICALBIRD MIGRATION DURING THE ICE AGES: ORIENTATIONAND ECOLOGY TIMOTHY C. WILLIAMS' AND THOMPSON WEBB III2 'Department of Biology, SwarthmoreCollege Swarthmore,Pennsylvania 19081, USA; and 2Departmentof GeologicalSciences, Brown University, Providence,Rhode Island 02912, USA ABsTRAcr.-Reconstruction of breeding habitat of North American Neotropical migrants 18,000 years ago and 9,000 years ago indicated major shifts in both location and composition of plant communities relative to present conditions. Increased vegetation in xeric areas may have compensated, at least in part, for the reduction in breeding habitat due to glaciation. Autumnal flights of Neotropical passerine migrants flying on constant headings from North America to Central and South America were simulated under present wind conditions and for winds during periods of glaciation at 18,000 and 9,000 years ago. The 1550average headings currently observed for Atlantic migrants were found to function well during periods of glaciation and may have been more generally useful during those times than at present. Received 15 June 1994, accepted 27 January 1995. CYCLESOF QUATERNARYGLACIATION during the ing the fitness of current bird populations over last 1.6 million years have influenced the de- their short lifetime? Would the orientation sys- velopment of present routes of both Neotrop- tem(s) used by birds under present conditions ical and Paleotropical migration (Moreau 1972, have been effective under past conditions? Gauthreaux 1982). North American Neotropical We examine whether fixed-heading orienta- migrants were most clearly affected because most tion would be sufficient to guide Neotropical of their present breeding grounds were covered migrants during periods of glaciation. We first by a permanent ice cap only 18,000 (radiocar- estimate breeding habitats of Neotropical mi- bon) years ago (18 kya) and because the ecology grants in North America at 18 kya (full glacia- of North America has undergone several major tion) and at 9 kya (partial glaciation). We then changes during the period of deglaciation (Im- simulate migratory flights at fixed headings for brie and Imbrie 1979, Emery et al. 1988, Webb these birds in order to examine the constraints 1988). Modern species assemblages or plant for- posed on orientation at those times in the past. mations are of recent origin in North America. The Neotropical migrants we discuss are a sub- Most developed after early Holocene warming: set of the type A Neotropical migrants defined modern tundra first appeared at about 8 kya, by the Research Working Group of Partners in boreal forest at 7 kya, and modern mixed forest Flight (1992). We consider only autumnal at 6-8 kya. Only modern prairie, first appearing (southward) migration. We exclude diurnal mi- about 10 kya, and modern deciduous forest (12 grants such as raptors, which are clearly influ- kya) existed during periods of significant gla- enced by local topography, and exclude birds ciation (Webb 1988). After their first appear- moving southwest within North America. We ance, plant formations increased in area and focus on shorebirds and passerines flying south approached their present distributions only in to southeast from North America to Central and the past 4,000 years (Van Devender 1986, Webb South America. We refer to this subset of birds 1988). The present distributions of plant taxa as Neotropical migrants. were relatively rare events during the Quater- Several lines of evidence, including radar and nary and, like the interglacial periods with radio-telemetry observations of migration, the which they are associated, have existed for only experimental manipulation of captive birds, and 10% of the 100,000-year glacial/interglacial cy- simulated migratory flights, suggest that at least cles (Ruddiman and Raymo 1988). These find- some songbirds and shorebirds guide long-dis- ings pose two major problems for the study of tance migrations by maintaining a fixed com- the evolution of bird migration. What behavior pass heading. Drift by winds produces a vari- would allow bird species to track changes in able track, but by selecting synoptic weather habitat over thousands of years, while preserv- conditions for take-off, the great majority of birds 105 This content downloaded from 130.58.65.20 on Mon, 27 Apr 2015 14:44:17 UTC All use subject to JSTOR Terms and Conditions 106 WILLIAMSAND WEBB [Auk, Vol. 113 can move reliably between breeding and win- METHODS tering areas each year (Williams and Williams We used the mapped pollen data from Webb (1988) 1978, 1990, Alerstam 1981, Stoddard et al. 1983, and Overpeck et al. (1992) to estimate bird habitats. Cochran and Kjos 1985, Alerstam et al. 1986, We labeled these habitats as desert, prairie, tundra, Richardson 1991, Williams 1991). Direction and coniferous forest, mixed forest, and deciduous forest some cue for the cessation of migration (dis- to describe structural features, but do not imply pres- tance, time, latitude, or habitat) appear to be ent species assemblages for each region in the past (Baker 1983, Webb et al. 1987, Thompson 1988, Webb genetically encoded (Wiltschko and Wiltschko 1988). We set the southern limit of breeding habitat 1978, Gwinner 1986, Berthold 1988, 1990). One for Neotropical migrants as the southern limit of kill- or more such vectors direct first-year birds to ing frost and set no northern continental limit. Breed- the general location of stopover and wintering ing habitat includes all forest types, wetlands, prairie areas, where they then search for suitable hab- and tundra, but excludes desert and permanent ice. itat. In subsequent years, birds return to these We considered Arctic coastal and North Slope areas local habitats using familiar landmarks or some as suitable habitat for shorebirds, but not Neotropical other orientation system operational within the passerine migrants. areas specified by the coded vector (Wiltschko To simulate migratory flights, we first created a and Wiltschko 1978, Williams and Williams 1990, wind matrix for North America representative of those winds actually used by Neotropical migrants. We then R. Wiltschko 1992). A variety of approaches have simulated flights through these wind fields with an shown that such fixed headings could be main- iterative computer program. Finally, we modified the tained by use of a compass based on the sun, wind matrix to estimate conditions during glaciation stars, wind, or magnetic field of the earth (Able and repeated the simulations with these wind con- 1980, Alerstam 1981, Wiltschko and Wiltschko, ditions. 1988). The methods for creation of a wind matrix under Although birds may use other techniques for present conditions are described in detail in Williams orientation (Rab0l 1978, Able 1980, Wiltschko (1991). The 50 latitude x 50 longitude matrix for pres- and Wiltschko 1988, 1991, Richardson 1991, ent conditions was an average wind velocity for those Wallraff 1991, R. Wiltschko 1992), orientation North American weather systems known to have sup- of and by a fixed compass heading is consistent with ported major autumnal migrations passerines shorebirds moving to the south and southeast as ob- available data for Neotropical migrants moving served by radar. Weather systems associated with such to south to southeast across North American movements were determined from radar observations Central and South America