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Evolution of Scaphinotus Petersi (Coleoptera: Carabidae) and the Role of Climate and Geography in the Madrean Sky Islands of Southeastern Arizona, USA

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Evolution of Scaphinotus Petersi (Coleoptera: Carabidae) and the Role of Climate and Geography in the Madrean Sky Islands of Southeastern Arizona, USA YQRES-03387; No. of pages: 10; 4C: 3, 5 Quaternary Research xxx (2012) xxx–xxx Contents lists available at SciVerse ScienceDirect Quaternary Research journal homepage: www.elsevier.com/locate/yqres Evolution of Scaphinotus petersi (Coleoptera: Carabidae) and the role of climate and geography in the Madrean sky islands of southeastern Arizona, USA Sara Gran Mitchell ⁎, Karen A. Ober Department of Biology, College of the Holy Cross, PO Box B, 1 College St., Worcester, MA 01610, USA article info abstract Article history: Geographically isolated environments such as the conifer forests atop the Madrean “sky islands” in southeastern Received 21 May 2012 Arizona provide natural laboratories for studying factors involved in speciation and origins of biodiversity. Using Available online xxxx molecular and geospatial analyses, we examine beetle population phylogeny, regional climate records, and the Quaternary paleobiogeography of forests to evaluate four hypothetical scenarios regarding the current Keywords: geographic and population genetic patterns of Scaphinotus petersi. Scaphinotus petersi is a large, flightless beetle Arizona that resides in the Madrean conifer forests above ~1900 m asl. Our results do not support the current hypothesis Sky islands Quaternary that S. petersi populations found on seven separate mountain ranges are genetically distinct and separated as Beetles temperatures warmed after the Last Glacial Maximum (LGM). Rather, we show that only some of the ranges Molecular evolution hold genetically distinct populations, and the timing of separation among the populations does not appear to Population divergence coincide with specific climatic events such as warming trends. In addition, we show that predicted changes to Paleobiogeography the climate of the Madrean sky islands may result in the disappearance of S. petersi from some of the lower ranges Climate by the end of this century. © 2012 University of Washington. Published by Elsevier Inc. All rights reserved. Introduction first hypothesis, cooler and/or wetter climate conditions in the south- western USA during the late Pleistocene favored more widely distributed The Madrean “sky islands” region of southeastern Arizona consists of conifer forests, and thus S. petersi habitat, at lower altitude. Divergence isolated forested mountain ranges that rise above a relatively flat prairie, occurred after post-Last Glacial Maximum (LGM) warming caused the scrub, and desert environment. Steep precipitation and temperature lower limit of conifer forests to contract uphill, eventually disconnecting gradients with altitude give rise to a series of ecological successions on forests and thus separating the beetle populations. If true, we would sky islands, and the ecosystems atop these ranges are sufficiently sepa- predict the following results from our genetic and paleobiogeographic rated from other suitable habitats such that the remoteness promotes analyses: 1) Beetle populations on different ranges will be genetically differentiation of organisms that live there. Such environments provide distinct from the others (in other words, each of the populations will natural laboratories for studying the processes involved in speciation be “monophyletic”). 2) Populations on ranges with conifer forests that and origins of biodiversity of endemic organisms (e.g., Brown, 1971; connect with the least amount of lowering or are close together will be Warshall, 1994). One such organism is the flightless, ground-dwelling more genetically similar and more recently separated than populations beetle, Scaphinotus petersi, characterized by Ball (1966).Today,S. petersi on ranges that require more conifer forest lowering to connect or are is found only in the Madrean montane conifer forest ecosystem (here separated by larger distances. 3) Now-separated populations should referred to as “conifer forests”) above ~1900 m elevation. According to have common ancestors during cooler, wetter times such as the Last Ball (1966), beetle populations on several sky island mountain ranges Glacial Maximum (LGM) and divergence should occur after the initiation have distinct morphological characteristics; therefore, Ball concluded of warmer or drier times during the Holocene. 4) Independent analyses that these ranges now host separate S. petersi subspecies. In this study, of conifer forest paleobiogeography and other paleoecological data will we examine S. petersi phylogeny, regional climate records, and the show that forests were connected during the cooler, possibly wetter paleobiogeography of Madrean montane conifer forests to test four hy- conditions thought to exist during the Pleistocene. potheses that could explain current geographic and population genetic A second possible scenario is that beetle dispersal can occur regard- patterns of this endemic beetle. less of habitat connectivity or proximity (no “island” effect), and is thus Ball (1966) hypothesized that all S. petersi subspecies were derived completely decoupled from changes in climate or forest biogeography. from a common ancestral population that once moved freely between In this scenario, we predict the following: 1) Genetic analysis of beetle mountain ranges when forests were more extensive. According to this populations will show evidence of a diversity of lineages (populations may be “polyphyletic” and contain distantly related individuals). ⁎ Corresponding author. Fax: +1 508 793 2696. 2) There may not be any relationship between mountain range proxim- E-mail address: [email protected] (S.G. Mitchell). ity or saddle altitude and the genetic relatedness of beetle populations. 0033-5894/$ – see front matter © 2012 University of Washington. Published by Elsevier Inc. All rights reserved. http://dx.doi.org/10.1016/j.yqres.2012.11.001 Please cite this article as: Mitchell, S.G., Ober, K.A., Evolution of Scaphinotus petersi (Coleoptera: Carabidae) and the role of climate and geography in the Madrean sky islands of southeastern Arizona, USA, Quaternary Research (2012), http://dx.doi.org/10.1016/j.yqres.2012.11.001 2 S.G. Mitchell, K.A. Ober / Quaternary Research xxx (2012) xxx–xxx 3) Climate conditions were not necessarily favorable for forest connec- conifer forests that occur on the highest mountain ranges in southeastern tivity during the timeframe represented by the phylogenetic tree. 4) Arizona. Evidence for gene flow or dispersal (the movement of one or more indi- viduals from one range to another) can occur during warm and/or dry Study area climate conditions. 5) The genetic data may be consistent with the re-population of ranges that could have lost and then regained their The Madrean sky islands area encompasses a nearly 200,000 km2 forests during the Holocene. area of southeastern Arizona, southwestern New Mexico and north- A third possible scenario is that the S. petersi populations were ern Mexico. This region is notable for its relatively small but high once fully connected and are now isolated, but they have not been (>2000 m) mountain ranges separated by low (b1000 m) topogra- separated long enough to diverge into distinct phylogenetic groups. phy. S. petersi populations are restricted to a subset of ranges: the In this scenario, 1) there would be no population genetic structure Pinal, Pinaleño, Santa Catalina, Rincon, Santa Rita, Chiricahua, and or genetic divergence seen among the mountain ranges and isolated Huachuca Mountains (Ball, 1966;Fig.2).S. petersi has been sighted populations would not fall into distinct monophyletic groups, and but not formally recorded in the Galliuros, Sierra Anchas, and Mogollon 2) there will be no geographic relationship to genetic structure, and Rim (K. Will and G. Ball, pers. comm., 2010). 3) the timing of divergence will not be dependent on climatic conditions. Tectonically, the Madrean sky island region is located in the Basin and A final scenario is a combination of the above. Perhaps some ranges RangeprovinceofNorthAmerica.Themountainrangeshereformedasa are more geographically isolated than others, and thus have distinct, result of the extension and uplift, likely in two stages (Wagner and possibly monophyletic beetle populations. But in this scenario, dispersal Johnson, 2006). The last period of peak uplift likely occurred between may occur between some mountains despite unfavorable climates or a 12 and 6 Ma (Wagner and Johnson, 2006). Due to the geometry of ex- lack of evidence for forest connectivity. In this last scenario, we might tension, the ranges are aligned along two major NW–SE trending ridges. see: 1) A complex phylogenetic tree, with some monophyletic and The ridges are separated by broad expanses, and along ridges, peaks are some polyphyletic groups of beetles on individual ranges. 2) A complex often separated by narrow but deep canyons. The San Pedro River sepa- relationship between mountain range geography and genetic similarity rates the two ridges, and the Gila River separates the northernmost Pinal and divergence times. 3) Climate conditions and their relationship to Mountains from all others to the south. forest connectivity may be irrelevant to the timing of beetle divergence. The steep topographic gradients in the area are associated with gradi- We evaluate these four scenarios by utilizing modern DNA sequenc- ents in temperature and precipitation. These climate gradients in turn ing and analysis tools, a GIS-based analysis of forest paleobiogeography, generate a rich diversity of ecosystems in a relatively small geographic and an analysis of geographic and climatic
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