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Systematics Of The Common Kingsnake (Lampropeltis Getula; Serpentes: Colubridae) And The Burden Of Heritage In Taxonomy
Abstract and Figures
We present a systematic revision of the Lampropeltis getula group, based on a recent range-wide phylogeographic analysis. We define our theoretical and operational concepts of species delimitation, and provide diagnoses based on mitochondrial DNA evidence, ecological niche modeling, morphology, and historical precedence. We find support for the recognition of five distinct species, which bear the name of the nominate subspecies found primarily within the range of each phylogeographic lineage: the Eastern lineage (Lampropeltis getula, Eastern Kingsnake), the Mississippi lineage (L. nigra, Black Kingsnake), the Central lineage (L. holbrooki, Speckled Kingsnake), the Desert lineage (L. splendida, Desert Kingsnake), and the Western lineage (L. californiae, California Kingsnake). Interestingly, all of these taxa had originally been described as distinct species and recognized as such for up to 101 years (in the case of L. californiae) before being demoted to subspecies. We discuss the impact that increasingly detailed genetic information from phylogeographic analyses may have on traditional taxonomy.
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... Recent taxonomic revisions in this group have further increased the total number of species and have been largely clustered within Lampropeltis and Pantherophis. Since 2000, the number of species of Lampropeltis has been recommended for increase from 9 to 24 (Bryson et al., 2005;Pyron and Burbrink, 2009b;Myers et al., 2013;Ruane et al., 2014;McKelvy and Burbrink, 2017). The clade that is now Pantherophis was increased from 3 species to 9 (Burbrink, 2001;Burbrink, 2002;Crother et al., 2011). ...
... The evolutionary history of two particular genera, Arizona and Rhinocheilus, in this tribe are notable because of the close co-distribution of their species (Fig. 2). The divergences associated with two major clades of Arizona elegans and two major clades of Rhinocheilus lecontei correspond to the Cochise filter barrier, a proposed barrier to gene flow in many other desert taxa (Liebherr, 1986;Castoe et al., 2007;Pyron and Burbrink, 2009b). The dates associated with these divergences in our analysis were very different from each other (9.78 MYA for Arizona, and 3.97 MYA for Rhinocheilus), and older than expected. ...
... The snake tribe Lampropeltini is one of the most heavily studied snake clades in the world and, over the last 30 years, the number of potential species in this tribe has increased from 23 to 51 with the majority of these being in Lampropeltis (Pyron and Burbrink, 2009b;Myers et al., 2013;Ruane et al., 2014). Additionally, many papers have been published that have explored historical biogeography (Rodriguez-Robles and De Jesus-Escobar, 1999, Burbrink and Lawson, 2007, Hansen and Salmon, 2017, morphological evolution (Keogh, 1996;Manier, 2004) and character evolution such as mimicry (Davis Rabosky et al., 2016), oviparity , and diet (Rodriguez-Robles and De Jesus-Escobar, 1999). ...
Accurate representation of lineage diversity through complete taxon sampling is crucial to understanding the evolution of biodiversity, particularly when using molecular phylogenetics to estimate evolutionary relationships. In this interest, taxonomic diversity is often used as a proxy for lineage diversity even though the two concepts are not synonymous. We explore this within the snake tribe Lampropeltini which includes some of the most conspicuous and heavily studied snakes in North America. Both the taxonomy and hypothesized relationships within this tribe have been in flux. The number of species has increased from 23 to 51 over the last thirty years, predominately within three of the nine genera (Lampropeltis, Pantherophis, Pituophis). The remaining six depauperate genera (Arizona, Bogertophis, Cemophora, Pseudelaphe, Rhinocheilus, and Senticolis) have been poorly represented in phylogenetic studies. To estimate evolutionary relationships and determine if the dichotomy in depauperate and speciose genera within Lampropeltini is a function of taxon sampling or truly represents the lineage diversity, we estimated the phylogeny of this group using nuclear and mitochondrial loci in a concatenated and coalescent framework with the largest sampling of the six depauperate genera to date. In addition, we estimated the divergence dates among the genera to assess whether the instability of Lampropeltini phylogenetic relationships is due to an adaptive radiation. While some nodes still remain unresolved, the generic-level relationships we recovered agree with those of a recent next-generation study that used a much larger set of loci for fewer individuals. We also tested two putative species, Arizona pacata and Pseudelaphe phaescens, for the first time phylogenetically and find evidence that they are distinct lineages. Overall, we find that the taxonomic and genetic diversity are not correlated in Lampropeltini and that representing putative diversity in phylogenies will lead to a better estimate of evolutionary histories, especially in groups with complex radiations.
... Lampropeltis getula.-The authors do not recognize the partitioning of this transcontinental species complex by Pyron and Burbrink (2009). ...
... When Bachia were tested using molecular data (Kohlsdorf and Wagner 2006;Galis et al. 2010;Kohlsdorf et al. 2010), paraphyly was suggested, although no consensus was reached. This has been termed Bthe burden of heritage in taxonomy^by Pyron and Burbrink (2009). ...
The phylogenetic and systematic relationships of the reduced limbed lizards of the genus Bachia are poorly understood. Here, we investigate the eastern Caribbean Bachia assigned to the B. heteropa and B. flavescens groups, whose members are characterized by a band of hexagonal or quadrangular scales on the dorsum, respectively. The polytypic Bachia heteropa is redefined, and the previous subspecies in the Grenadines (Bachia heteropa alleni) and Trinidad (B. h. trinitatis) are demonstrated to be species-level lineages. One new species of hex-scaled Bachia was formerly assigned intergrade status between B. heteropa and Bachia trinitatis. Here, it is described as a new species from Caripito, Venezuela. Bachia h. heteropa, B. h. lineata, and B. h. marcelae are elevated to species status. The Tobago species formerly considered a member of the Bachia flavescens species group is described as a new species. In this paper, we increase the number of species in the genus Bachia from 25 to 31 with the description of two new species and the elevation of four previously described species from the synonymy of Bachia heteropa. This work will greatly improve the understanding of the systematics and evolution of Bachia in the eastern Caribbean.
... tropical vs. temperate). Several other studies have found divergent climatic niches between closely related species, including studies of frogs (Hua and Wiens, 2010), lizards ( Knouft et al., 2006), and snakes (Pyron and Burbrink, 2009). Several studies have also found interesting patterns of within-species phenotypic divergence and environmental variation that may lead to parapatric speciation (e.g., Schneider et al., 1999;Ogden and Thorpe, 2002). ...
Speciation processes have long been inferred from phylogenetic, phylogeographic, and biogeographic pattern-driven perspectives. Now much current speciation research is attempting to more directly describe the underlying processes and mechanisms of divergence leading to speciation. Ideally, researchers should integrate both process- and pattern-based approaches for a more comprehensive understanding of speciation. To this end, a symposium was organized during the 7 th World Congress of Herpetology in Canada with the goal of bringing leading experts together to share successful examples of these perspectives and to promote a more cohesive understanding of reptile and amphibian speciation. Here we present a joint paper of short and updated summaries of each of these contributions with the aim of providing a reference source and launching pad for students and researchers interested in speciation in amphibians and reptiles.
... tropical vs. temperate). Several other studies have found divergent climatic niches between closely related species, including studies of frogs (Hua and Wiens, 2010), lizards (Knouft et al., 2006), and snakes (Pyron and Burbrink, 2009). Several studies have also found interesting patterns of within-species phenotypic divergence and environmental variation that may lead to parapatric speciation (e.g., Schneider et al., 1999;Ogden and Thorpe, 2002). ...
Speciation processes have long been inferred from phylogenetic, phylogeographic, and biogeographic pattern-driven perspectives. Now much current speciation research is attempting to more directly describe the underlying processes and mechanisms of divergence leading to speciation. Ideally, researchers should integrate both process-and pattern-based approaches for a more comprehensive understanding of speciation. To this end, a symposium was organized during the 7 th World Congress of Herpetology in Canada with the goal of bringing leading experts together to share successful examples of these perspectives and to promote a more cohesive understanding of reptile and amphibian speciation. Here we present a joint paper of short and updated summaries of each of these contributions with the aim of providing a reference source and launching pad for students and researchers interested in speciation in amphibians and reptiles.
... al. (1996, Dessauer and Pough (1975), Duméril and Bibron (1835), Enge (2009), Fitch (1936, Franklin (1998), Green and Pauley (1987), Grismer (1999), Gutberlet and Franklin (1996), Hallmen (2005Hallmen ( , 2006, Hay (1902), Hibbitts (1998), Irwin (2004), Jan (1865a, 1865b), Klauber (1938), Kreutz (2005), Krysko (1998), Krysko and Hurt (1998), Krysko and Judd (2006), Lara-Gongora et. al. (1993), Lazell and Musick (1973), LeClere (1995), Liner (1996), Linné (1766), Lönnberg (1894), Mattison (2007), Means (1998), Meierkord (2010), Mitchell (1994), Murphy and Ottley (1984), Neill and Ross (1949), Palmer and Braswell (1995), Phillips and Petzing (1998), Price (1987), Pyron and Burbink (2009a, 2009b, 2009c, Schmidt (2004Schmidt ( , 2005, Jauch (1980a, 1980b), Shoop (1957), Skubowius (2009Skubowius ( , 2010, Slevin (1950), Smith (1956), Snyder (1945, Stebbins (1985), Stejneger (1902), Stevens (1994), Tanner (1927), Tanner (1958), Taylor (1952), Thissen and Hansen (2001), Thornton and Smith (1993), Thums (2004), Van Denburgh and Slevin (1921), Werner (1924), Wilgers et. al. (2006, Woodbury (1928), Yarrow (1882), Young and Iverson (1997) and Zweifel and Norris (1955). ...
The King and Milk Snakes, Lampropeltis Fitzinger, 1843 are familiar to most American
herpetologists. Notwithstanding their familiarity and general abundance. the taxonomy of
the genus has remained unstable to the present time.
Confusion and dispute remains in terms of the exact number of species.
Even the generic placement of members has been unstable in recent years.
In 2009, Pyron and Burbink placed the short-tailed snake, known widely as Stilosoma
extenuatum within the synonymy of Lampropeltis.
Other available genus names for subgroups and species groups have generally not been
used.
Most recently the detailed evidence published by Pyron et. al. (2011) led the authors to
note that they viewed the genus Lampropeltis to be paraphyletic at the genus level as
currently defined.
Viewing this evidence and the obvious morphological and behavioral differences between
the species groups, this paper divides the genus as currently accepted in three ways.
Lampropeltis retains the type species getula and several others, including Stilosoma which
remains subsumed as does Ophibolus Baird and Girard, 1853. Oreophis Dugès, 1897 is
resurrected to contain the type species mexicana and several others. Finally the divergent
taxon, calligaster is placed within its own monotypic genus Eksteinus gen. nov.
Keywords: new genus; Kingsnake; Milksnake; Lampropeltis; Stilosoma; Ophibolus;
Oreophis; Eksteinus; calligaster; Prairie Kingsnake; Mole Snake; Florida Mole Snake.
... Of the squamate prey where ingestion direction potentially was determinable, 18 were consumed head first and prey direction was indeterminable for one small Diadophis punctatus. interpretation of the activity patterns, reproductive biology, and prey base of the species at this site and throughout the range of this species and its closely related congeners (Pyron and Burbrink 2009a). ...
Eastern Kingsnakes (Lampropeltis getula) are an important component and predator in herpetofaunal communities∗but many Eastern Kingsnake populations have declined precipitously in the last few decades, particularly in the southeastern United States. Here, we describe an intensive capture-mark-recapture study of L. getula conducted during 1974-1978 in a canal bank-Water Hyacinth (Eichhornia crassipes) community at Rainey Slough in southern Florida, where annual capture probabilities of adults ranged from 0.662-0.787. Population size and structure, seasonal activity, movements, microhabitat use, behavior, thermal ecology, and predator-prey relationships are described. At this site kingsnakes were susceptible to capture mostly in winter and spring, were diurnal, used rodent (Sigmodon hispidus) burrows on canal banks as nocturnal retreats, and emerged from burrows on 13-26% of the sampling days. Overlap of burrow use by both sexes was extensive with no evidence of territoriality. Kingsnakes readily entered the Water Hyacinths to bask, pursue mates, and forage. At Rainey Slough only snakes were detected in the diet of kingsnakes. Concurrent sampling of potential snake prey in the hyacinths and on canal banks revealed 10 species that varied in use of the two sampled habitats and in body size. A range-wide analysis confirmed that in descending order snakes, reptile eggs, and lizards dominate the diet of L. getula in Florida (94.8%) and remain important prey types elsewhere (80.2%). At Rainey Slough the density of six species of semiaquatic snakes in Water Hyacinths averaged 3534 individuals/ha with a mean annual biomass of 135.8 kg/ha, and kingsnake biomass was only 2.2-3.9% of prey snake biomass. We estimated that the kingsnake population consumed 36.82-63.58 kg/yr, or about 10.0-17.2% of the standing crop of snakes in the Water Hyacinth community. Adult male L. getula lost on average 39.3% of their body mass associated with the spring reproductive season, whereas females lost only 3.4% in the same period. Body condition indices for both sexes improved substantially thereafter. In follow-up surveys at Rainey Slough during 2006-2010 no kingsnakes were found. Semiaquatic snake densities in the Water Hyacinths were 77.2% lower (807.4/ha) than in the 1970s and consisted of only three species. Compared to the enigmatic declines and extirpation of L. getula populations elsewhere, at Rainey Slough the primary cause likely was unsustainable mortality from road reconstruction and paving in the winter-spring of 1979 and subsequent roadkill. Other potentially causative agents of extirpation of L. getula in this system are discussed.
... While taxonomy-based tests may represent a natural progression from earlier studies (i.e., validation of described morphological variation), they may also violate assumptions of some analyses, especially when paraphyly or gene flow are present. Furthermore, tests predicated on previously described variation often miss cryptic variation (Leache et al., 2009;Pyron and Burbrink, 2009). Tests predicated solely on morphology may best reflect phenotypic responses to environments, but may also fail to provide the optimal basis for exploring evolutionary history or establishing taxonomy (Gotthard and Nylin, 1995;Mayr, 1956;Simpson, 1951), although there are exceptions (Hoekstra et al., 2005;Patton and Smith, 1994). ...
Delimiting species can be challenging, but is a key step for the critical examination of evolutionary history and for prioritizing conservation efforts. Because systematic relationships are often determined iteratively using tests based on taxonomy, such methods can fail to detect cryptic variation and result in biased conclusions. Conversely, discovery-based approaches provide a powerful way to define operational taxonomic units and test species boundaries. We compare both approaches (taxonomy-based delimitation - TBD and discovery-based delimitation - DBD) within North American jumping mice (Zapodinae) using broad sampling, multilocus analyses, and ecological tests. This group diversified through the dynamic glacial-interglacial periods of the Quaternary and phylogeographic tests reveal 28 lineages that correspond poorly with current taxonomy (4 species, 32 nominal subspecies). However, neither the 4-species or 28-lineage hypotheses are optimal for species-level classification. Rather, information theoretic approaches (Bayes Factors) indicate a 15-species hypothesis is best for characterizing genetic variation in this group, with subsequent iterative pairwise ecological tests failing to confirm four species pairs. Taken together, evolutionary and ecological tests capture divergence among 11 putative species that, if upheld by additional tests, will lead to taxonomic revision and reevaluation of conservation plans.
Phylogeography is a relatively young fi eld that investigates the historical and contemporary processes that affect the geographic distribution of genea-logical lineages, particularly those at the intraspecifi c level (Graves et al. 1984; Avise et al. 1987; Avise 1998). Phylogeography occupies a place between mi cro -evolutionary (demography, population genetics, and ethology) and macro-evolutionary (systematics, historical biogeography, and paleoecology) fi elds (Avise 2000). Ironically, since the term was coined, the lines that demarcate phylogeography from phylogenetic and population genetic studies has sub-stantially blurred, and it may be more reasonable to consider this sub discipline to be research that incorporates both macro-and microevolutionary processes rather than occupying a discrete space between these two scales. The main benefi t of phylogeographic studies is that they reveal patterns that are too diffi cult to discover using other less integrative approaches. For instance, phylogeographic studies can detect cryptic genetic diversity in the geographic range of a taxon, which may be an early step in the recogni-tion of new species (e.g., Zamudio and Greene 1997; Burbrink et al. 2000; Parkinson et al. 2000; Burbrink 2001; Rodríguez-Robles et al. 2001; Feld-man and Spicer 2002; Castoe et al. 2003, 2005). In this sense, phylogeog-raphy may provide the initial information about the geographic range of a newly defi ned lineage; this in turn may supply critical information used to prioritize conservation efforts aimed at maintaining viable populations of newly discovered lineages (Fig. 2.1). Because assessing species boundaries and recognizing the true biodiversity of a region is a primary goal for con-servationists, phylogeographic research is tied intimately to conservation bi-ology.
Texas populations of slimy salamanders are isolated from other members of the Plethodon glutinosus complex and are currently placed in the species P. albagula with populations from the Ozark Plateau and Ouachita Mountains of Missouri, Oklahoma, and Arkansas. We sequenced a portion of the mitochondrial protein-coding gene ND4 for 52 P. albagula from 25 localities in Texas and one locality in Missouri. Bayesian MCMC analysis identified five parapatric lineages of Plethodon in Texas. Several morphologically distinctive populations are phylogenetically embedded within the Texas radiation. Additional studies of the interactions among the lineages identified in this study will be necessary to resolve their taxonomic status.
