Arboreality in the California Whipsnake (Masticophis lateralis): Implications for Survey Techniques

J. Alvarez, A. Murphy
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Abstract

Historically, new species were defined by mensural characters and descriptive analyses of external features (i.e., coloration, patterning, behavior) that attempted to delineate new species from those previously described. Ecological associations (e.g., habitat, microhabitat, geographic extent or location, etc.) further defined the differences between one species and those closely related (Van Denburgh 1897). Over time, the species’ niche becomes better defined as various researchers use different foci or perspectives to investigate natural history traits (Storer 1925). Occasionally, natural history traits that were once considered putative, particularly in older references, are lost to the cognizance of modern researchers and become rarely referenced. The California whipsnake [Masticophis (=Coluber) lateralis], first described by Hallowell in 1853, provides an example of this phenomenon. This species is a relatively long, narrow-bodied snake that is found throughout the foothills or lowto mid-elevations of portions of the Coast, Cascade, Sierra Nevada, Transverse and Peninsular Ranges in California, as well as northwestern Baja California (Stebbins 1954; Brown 1997; Grismer 2002; Flaxington 2021). Early investigators described the California whipsnake as arboreal (Grinnell and Grinnell 1907; Grinnell and Storer 1924; Ortenburger 1928; Pickwell 1947; Stebbins 1954; Appendix I), but when an extended gap in published work on snake habitat occurred, awareness of the snake’s arboreal traits did not span that gap. Here, we draw upon our own recent investigations to describe arboreal traits of the California whipsnake and use those observations to suggest that it be considered semiarboreal, and that survey and management techniques should reflect this snake’s behavior. Further, we describe survey techniques designed to examine habitat features that best support the arboreal nature of this snake. Following Stebbins (1954) general natural history account, nearly 40 yrs elapsed before new information on California whipsnake habitat and microhabitat was published. Subsequent to subspeciation of M. lateralis, with one subspecies becoming listed as threatened in the San Francisco Bay region by state and federal wildlife agencies, a new focus on the natural history of the species was initiated (Reimer 1954; CDFW 1987; USFWS 1997, 2002). Interest in the listed Alameda whipsnake subspecies (M. l. euryxanthus) sparked a flurry of new observations and research beginning with Swaim and McGinnis (1992) who reported habitat associations for the Alameda whipsnake, including preferred habitat features. However, they made no mention of the arboreal behavior in the species or subspecies, nor the potential effects of this behavior on their methodology for study. Lind (1992) picked up the thread carried by early researchers when she reported that a California whipsnake was “found 2 m above ground in a canyon live oak (Quercus chrysolepis)...” supporting the arboreal nature of the species described in historical accounts. In 1994, Swaim prepared a detailed report on the ecology of M. l. euryxanthus, but arboreality and climbing ability, and its potential impact on the ground-level trapping surveys used to
加州鞭蛇(Masticophis lateralis)的树栖特征:调查技术的意义
从历史上看,新物种是通过测量特征和对外部特征(即颜色、图案、行为)的描述性分析来定义的,这些分析试图从先前描述的物种中划定新物种。生态关联(如生境、微生境、地理范围或位置等)进一步界定了一个物种与那些密切相关的物种之间的差异(Van Denburgh 1897)。随着时间的推移,随着不同的研究人员使用不同的焦点或视角来研究自然历史特征,物种的生态位变得更好。偶尔,曾经被认为是假定的自然历史特征,特别是在旧文献中,被现代研究人员所忽视,很少被引用。加州鞭蛇(Masticophis (=Coluber) lateralis)在1853年由Hallowell首次描述,提供了这种现象的一个例子。该物种是一种相对较长的窄体蛇,遍布加利福尼亚海岸、喀斯喀特、内华达山脉、横向和半岛山脉部分的山麓或中低海拔地区,以及下加利福尼亚州西北部(Stebbins 1954;布朗1997;“2002;Flaxington 2021)。早期的研究人员将加利福尼亚鞭蛇描述为树栖的(Grinnell and Grinnell 1907;格林内尔和斯托弗1924;Ortenburger 1928;Pickwell 1947;史泰宾斯1954;附录1),但是当关于蛇的栖息地的出版工作出现了一个扩大的空白时,对蛇的树栖特征的认识并没有跨越这个空白。在这里,我们利用我们自己最近的调查来描述加州鞭蛇的树栖特征,并使用这些观察结果来建议它被认为是半树栖的,并且调查和管理技术应该反映这种蛇的行为。此外,我们描述了旨在检查栖息地特征的调查技术,这些特征最能支持这种蛇的树栖性质。继Stebbins(1954)的博物通论之后,近40年后才有关于加州鞭蛇栖息地和微栖息地的新信息发表。随后subspeciation m .外侧,与一个亚种成为旧金山湾地区列为威胁由州和联邦野生动物机构,一个新的关注物种的自然历史开始(雷蒙1954;CDFW 1987;Usfws 1997, 2002)。对所列的阿拉米达鞭蛇亚种(m.l. euryxanthus)的兴趣引发了一系列新的观察和研究,始于Swaim和McGinnis(1992),他们报告了阿拉米达鞭蛇的栖息地关联,包括偏好的栖息地特征。然而,他们没有提到物种或亚种的树栖行为,也没有提到这种行为对他们的研究方法的潜在影响。林德(1992)接过了早期研究人员的线索,她报告说,一条加州鞭蛇“在离地面2米的峡谷活橡树(Quercus chrysolepis)中被发现……支持了历史记载中所描述的物种的树栖性质。1994年,Swaim编写了一份详细的euryxanthus生态学报告,但对M. l. euryxanthus的树栖性和攀爬能力及其对地面捕获的潜在影响进行了调查
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