This file was created by scanning the printed publication. l Errors identified by the software have been corrected; however, some errors may remain. f:~~ United States ~.~flWJ" .,.} Department of Biodiversity and \'''',,;,..• .-.1' (I Ag ricultu re Forest Service Management of the Rocky Mountain Forest and Range Madrean Archipelago: Experiment Station The Sky Islands of Fort Collins, CO 80526 Southwestern United States and General Technical Northwestern Mexico Report RM-GTR-264 September 19-23, 1994 Tucson, Arizona USDA Forest Service July 1995 General Technical Report RM-GTR-264 Biodiversity and Management of the Madrean Archipelago: The Sky Islands of Southwestern United States and Northwestern Mexico September 19-23, 1994 Tucson, Arizona Technical Coordinators: Leonard F. DeSano Peter F. Ffolliott Gerald J. Gottfried University of Arizona Robert H. Hamre Carleton S. Edminster Alfredo Ortega-Rubio Rocky Mountain Forest and Range Centro de Investigaciones Biologicas Experiment Station del Noroeste Page Design: Carol LoSapio Rocky Mountain Forest and Range Experiment Station Sponsors: Rocky Mountain Forest and Range School of Renewable Natural Resources Experiment Station University of Arizona U.S. Department of Agriculture Tucson, Arizona Fort Collins, Colorado The Madrean Sky Island Archipelago: A Planetary Overview Peter Warshall 1 Abstract.-Previous work on biogeographic isolation has concerned itself with oceanic island chains, islands associated with continents, fringing archipelagos, and bodies of water such as the African lake system which serve as "aquatic islands". This paper reviews the "continental islands" and compares them to the Madrean sky island archipelago. The geological, hydrological, and climatic context for the Afroalpine, Guyana, Paramo, low and high desert of the Great Basin, etc. archipelagos are compared for source areas, number of islands, isolating mechanisms, interactive ecosystems, and evolutionary history. The history of scientific exploration and fieldwork for the Madrean Archipelago and its unique status among the planet's archipelagos are summarized. In 1957, Joe Marshall published "Birds of the American Prairies Province of Takhtajan, 1986) Pine-Oak Woodland in Southern Arizona and Ad­ and western biogeographic provinces, a wealth of jacent Mexico." Never surpassed, this elegant genetically unique cultivars in the Sierra Madre monograph described the stacking of biotic com­ Occidental, and a myriad of mysteries concerning munities on each island mountain from the the distribution of disjuncts, species "holes," and Mogollon Rim to the Sierra Madre. He defined the species II outliers" on individual mountains (e.g., Madrean archipelago as those island mountains Ramamoorthy, 1993). The northernmost sky is­ with a pine-oak woodland. In 1967, Weldon Heald lands are the only place in North America where (1993), from his home in the Chiricahuas, coined you can climb from the desert to northern Canada the addictive phrase-"sky islands" for these in­ in a matter of hours (Warshall, 1986). The sky is­ sular mountains of the North American lands pose numerous puzzles about vertical borderlands. Weldon Heald's catch phrase immor­ migration strategies used by plants and animals talized Joe Marshall's meticulous observations. both annually and over glacial time periods. Today's conference is the first solely dedicated to These interests in ecology and evolution mix understanding Madrean sky island biology, with the other citizen interests in skiing, grazing, beauty, and needs for management and conserva­ hunting, fishing, escaping the heat, summer tion. homes, telescopes and radio towers, bird watch­ There are about 40 sky islands (fig. 1) between ing, rock-climbing, military practice maneuvers, the Mogollon Rim and the Sierra Madre Occiden­ fuelwood cutting, camping, mining, sacred Native tal (Warshall, in press). Mt. Graham on the American values and ceremonies, archaeological Pinaleno mountains is the tallest peak (10,712 sites, as well as preservation of sky island habitat feet). Relief between valleys and peaks ranges for threatened and endangered species. Underpin­ from 1250 to 6750 feet (McLaughlin, 1992), but is ning all these interests is the exceptional beauty of typically between 3000 and 5000 feet. The Ma­ the sky islands-their layering of peaks in a dusty drean region has exceptional species richness, sunset, lines of vibrant riparian along arid yellow super-species complexes, unusual neoendemics slopes, the contrast of snow and desert, the baf­ and archeoendemics, an exceptional mixture of fling complexity of erratic ridge lines, the power species from the Nearctic and Neotropic regions, of fires, and the subtle tones of blue-gray lime­ important influences from the eastern (North stonE!, speckled granite, and pastel volcanics. 10ffice of Arid Lands Study, Col/ege of Agriculture, University of Ari­ zona, 845 N. Park, Tucson, AZ 85719. 6 coniferous fortst \"~'~"""-'10 .\~"."II' .> j .,11 pine-oak woodland _'\6, ..v,.... 2 encinal ,', -';";'---'---'--- 9ral.land -,,\I .. d ... rt .. H.. "'\11., ,\,,, ,\LII" .\,', o 10 20 30 40 50Miles Figure 1.-The Madrean Sky Island archipelago (based on Marshall, 1951), 7 WHAT ARE SKY ISLANDS? SKY ISLANDS OF THE PLANET Sky islands are a type of continental or inland There are about twenty sky island complexes terrain made up of a sequence of valleys and on the planet (table 1 and fig. 3). All the conti­ mountains. All sky islands have a stack of biotic nents with, perhaps, the exception of Australia, communities that allow vertical (as well as aspect) harbor sky island complexes. The information on migration annually or during one of the planet's the sky island complexes of Eurasia, China, and long-term climatic events. The valleys act as barri­ southeast Asia remains incomplete because of the ers or bridges to the colonization by new species difficulty of obtaining English translations (Sus­ that attempt to cross the intervening valley. The lov, 1961; Aiken, 1992). Most of the literature has valleys become barriers when they contain an focused on the mountains themselves, not the im­ ecology alien to the migrating species. By analogy portance of the valleys between them. There has with the saltwater seas between oceanic islands, been remarkably lithe work comparing the the higher elevation biotic communities of sky is­ planet's continental island ecosystems (Carlquist, land mountains are isolated by each valley'S "sea" 1963), their palaeogeographic history, floristic and of alien vegetation. The mountains, like the Gala­ faunal source areas, and valley barriers. pagos or other oceanic islands, act as isolated By creating a parallel typology with recent cradles of evolution. classifications of oceanic islands such as isolated In the Madrean archipelago, the valleys and island "chains" (Hawaii), "continent associates" mountains are roughly parallel. The stacked biotic (Madagascar, Philippines), and "fringing archipel­ communities (fig. 2) include: montane coniferous agos" (the southern Japanese or Sea of Cortez forests; oak-pine (coniferous) woodlands; tropical island groups), we can approximate a classifica­ deciduous forest; oak savanna (deciduous vs. ev­ tion for the continental islands. Table 1 classifies ergreen oaks predominate); short-grass prairie; continental island clusters on the basis of their subtropical thornscrub; and subtropical desert geographic axis, latitude, whether they are coastal (Brown, 1982). The "heart" of the Madrean archi­ or inland, the number per complex, and the con­ pelago (its defining characteristic) is the oak-pine figuration of each grouping. Configurations woodlands (Marshall, 1957). The barrier" seas" in­ include: stepping stone archipelagos (mountains cl ude the short-grass prairie, the subtropical and valleys spaced between two cordilleras), iso­ thomscrub, and subtropical desert. lated massif(s) with outlier sky islands, linear NORTH SOUTH~ 11.000 h,t-- 10.000--- Wl,---Ab••• contolo' ----.~r-~t.;;;--- AbllS concolor and ---~ij.,l~ Pstudo'lug" lalifolla 8.000--- or chaparral----'\\.I '~---'ncinal----............. 4.0007&------------....Ji\ vrassland ------~ 3.000---------------!.-.;!:.------------_-4r---_____________ WHITE PINALENO HUACHUCA OPOSURA Figure 2.-Example "stacking" of biotic communities of the Madrean archipelago (Marshall, 1957). Note the position of the oak-pine woodlands. 8 chains of outlier mountains/valleys at one end of Table 1.-Sky Island Types a cordillera, and completely isolated groups of ~ =E~xa=m~p~l~e~s ________________ mountains. A surprisingly large number of com­ Stepping stone archipelago Madrean archipelago plexes (e.g., the Adamoua, Baja Peninsula, and between two mountain chains Great Basin archipelago Altai/Tien Shan Basin East African arc) contain both oceanic islands and Meso-American massifs sky islands with both saltwater and vegetative "seas." Like all typologies, there are ambiguously Isolated massif with outliers Ethiopian highlands East African arc categorized groupings. Saharan massifs Atlas Mountains Jabal Lubnan Adoumoua Mountains Planetary Examples Drakenbergs Central European massifs The four descriptive variables used to classify Caucaso-Iranian massifs sky islands act in concert. Compare the Madrean Cordillera with outliers Chaine Annimatique (Vietnam) stepping stone configuration with the isolated ar­ Malay peninsula chipelago of the Western Ghats of India (not Baja California peninsula Coastal Cordillera (SA) illustrated), the isolated cluster of the Pantepuis of