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Biological Relevance of Antler, Horn, And Journal of Mammalogy, 100(5):1447–1458, 2019 DOI:10.1093/jmammal/gyz123 Biological relevance of antler, horn, and pronghorn size in records programs Downloaded from https://academic.oup.com/jmammal/article-abstract/100/5/1447/5555716 by University of Wyoming Libraries user on 28 October 2019 Tayler N. LaSharr,* Ryan A. Long, James R. Heffelfinger, Vernon C. Bleich, Paul R. Krausman, R. Terry Bowyer, Justin M. Shannon, Eric M. Rominger, Chadwick P. Lehman, Mike Cox, and Kevin L. Monteith Wyoming Cooperative Fish and Wildlife Research Unit, Department of Zoology and Physiology, University of Wyoming, Dept. 3166, 1000 E University Avenue, Laramie, WY 82071, USA (TNL) Department of Fish and Wildlife Sciences, University of Idaho, 875 Perimeter Drive, MS 1142, Moscow, ID 83844, USA (RAL) Arizona Game and Fish Department, 5000 W. Carefree Highway, Phoenix, AZ 85086, USA (JRH) Department of Natural Resources and Environmental Science, University of Nevada Reno, Mail Stop 186, 1664 North Virginia Street, Reno, NV 89557, USA (VCB) School of Natural Resources and the Environment, University of Arizona, 1064 East Lowell Street, Tucson, AZ 85712, USA (PRK) Institute of Arctic Biology, University of Alaska Fairbanks, 902 North Koyukuk Drive, Fairbanks, AK 99775, USA (RTB) Utah Division of Wildlife Resources, 1594 W N Temple, Salt Lake City, UT 84114, USA (JMS) New Mexico Department of Game and Fish, 1 Wildlife Way, Santa Fe, NM 87507, USA (EMR) South Dakota Game Fish and Parks, 13329 US HWY 16A, Custer, SD 57730, USA (CPL) Nevada Department of Wildlife, 6980 Sierra Center Pkwy #120, Reno, NV 89511 (MC) Haub School of Environment and Natural Resources, Wyoming Cooperative Fish and Wildlife Research Unit, Department of Zoology and Physiology, University of Wyoming, 804 East Fremont Street, Laramie, WY 82072, USA (KLM) * Correspondent: [email protected] Long-term datasets are becoming increasingly important for assessing population- and species-level responses to a changing environment. Programs that record morphological measurements of horns, antlers, and pronghorns were established in the early- to mid-20th century to collect biological information about animals that possess large horns, antlers, or pronghorns, which could be used to assess the effectiveness of conservation efforts for large mammals in North America. The general relevance of record books has been questioned because of the minimum size requirements for inclusion in a record book, which may mask trends when changes in the population occur. We compared trends in size of antlers, horns, and pronghorns through time using records from three records programs with different minimum size requirements to evaluate the influence of entry requirements on temporal trends. We also investigated whether horn, antler, or pronghorn size affected the probability of specimens being submitted to a records program. Only two of 17 categories exhibited less-pronounced trends in the record book with the highest size requirements for entry, and in two categories trends were more pronounced. Although societal interest in submitting eligible specimens increased slightly over time in one of six categories, the probability of voluntary entry was largely random and not affected by year of harvest or size of specimen. In contrast to previous criticisms, trends in record books should not be expected to represent the size of all males within a population. Instead, our evaluation indicates that the records programs we examined can provide a useful resource for assessing long-term changes in phenotypic characteristics of ungulates, but importantly, they represent the respective range of sizes within which each program collects data. Key words: antler size, citizen science data, horn size, records programs, selective harvest, temporal trends Long-term (> 20 years) ecological datasets, although rare, are 1959; Franklin 1989; Lindenmayer et al. 2012). Ecological and critically important for understanding ecological and evolu- evolutionary change in wild populations can take many gener- tionary responses of animals to environmental change (Odum ations to manifest, and understanding the complex processes © 2019 American Society of Mammalogists, www.mammalogy.org 1447 1448 JOURNAL OF MAMMALOGY that lead to change at the individual, population, or species one of the longest-running datasets available to wildlife man- level is difficult without access to reliable, long-term datasets agers, and are potentially useful for answering questions about (Lindenmayer et al. 2012). Wildlife populations are valued by effects of harvest practices or environmental change on the size people around the globe, and sustainable harvest of wildlife is an of antlers and horns (Monteith et al. 2013). Much like citizen important management goal. Yet, evolutionary changes resulting science data, records programs represent an enormous source of Downloaded from https://academic.oup.com/jmammal/article-abstract/100/5/1447/5555716 by University of Wyoming Libraries user on 28 October 2019 from hunting have been documented (Kuparinen and Festa- information collected across broad temporal and spatial scales Bianchet 2017). Despite controversy associated with the evolu- (Monteith et al. 2013), and that information may be useful for tionary effects of harvest (Mysterud 2014; Festa-Bianchet 2016; assessing changes in the size of horns and antlers through time. Heffelfinger 2018), harvest strategies that result in evolutionary Despite the potential relevance of record books to wildlife responses have important conservation and management impli- ecology and management, there has been criticism over the cations (Kuparinen and Festa-Bianchet 2017). Unfortunately, validity of using data maintained in record books to assess disentangling the effects of selective harvest from selective pres- trends in sizes (Festa-Bianchet et al. 2015). Data maintained sures imposed by the environment (Berger 2005) is difficult by records programs represent a biased sample, because each without long-term data (Festa-Bianchet and Mysterud 2018). specimen must be a certain minimum size to qualify for entry. Under sufficiently intensive harvest pressure, artificial selec- Specimens documented by records programs only include in- tion can lead to undesirable evolutionary changes, including re- dividuals with large, or in some instances exceptionally large, duced reproductive output (Coltman et al. 2003), earlier sexual horns and antlers that may be approaching their maximum at- maturation (Walsh et al. 2006), reduced allele frequencies tainable size (Festa-Bianchet et al. 2015). Consequently, using (Walsh et al. 2006), potentially altered behaviors (Allendorf and data maintained by records programs to identify more general Hard 2009), and changes in population structure (McCullough temporal trends may be invalid or dilute changes at the popula- 1979; Conover and Munch 2002; Conover et al. 2009). For tion level (Festa-Bianchet et al. 2015). wild populations of large-bodied mammals, empirical evidence Records programs were not designed to document popula- supporting genetic change through artificial selection remains tion- or species-level changes, but to record the size and trends limited. As humans continue to harvest populations of large of horns, antlers, and pronghorns of large males. If temporal mammals around the world, understanding the effects of artifi- trends detected in record books are not representative of what cial selection on populations or species over time will become is occurring among males with large horns or antlers, then they increasingly important (Mysterud 2011; Festa-Bianchet 2016; may not be a useful resource for assessing consequences of har- Heffelfinger 2018). vest or environmental change, a result that would undermine Among ungulates, the potential evolutionary consequences one of the primary goals of records programs: to assess the con- of harvest by humans can be assessed by evaluating temporal sequences of conservation strategies (Monteith et al. 2013). changes in antler, horn, or pronghorn size (hereafter, antlers Hunters have been particularly interested in males that pos- and horns) because those traits are heritable, linked to repro- sess exceptionally large horns or antlers since records programs ductive success, and are often the target of selective harvest were established. If that interest varied through time, it could (Crosmary et al. 2013; Monteith et al. 2013; Festa-Bianchet bias temporal trends in size of horns and antlers detected in et al. 2014; Pigeon et al. 2016). Assessing changes in size over record books. For example, if interest in submitting specimens time is difficult in long-lived species because changes in allele that barely meet the minimum requirements of record books in- frequencies can take decades to manifest as a detectable pheno- creases through time, it could produce a negative trend in horn typic change; moreover, long-term data on phenotypic charac- or antler size (Monteith et al. 2013). If societal interest in en- teristics are often lacking (Hundertmark et al. 1998). tering specimens into a records program has changed through Measurements from harvested animals may provide a rich time, or as a function of the size of the specimen, then trends in source of data for quantifying temporal changes in phenotypic record books may not represent real biological change among characteristics (Torres et al. 1993; Monteith et al. 2013). For large males. most ungulate species in North America, phenotype of har- Our
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