Emergent patterns between salamander prey and their predators

Journal Pre-proof Emergent patterns between salamander prey and their predators

Kasey L. Jobe, Carmen G. Montaña, Christopher M. Schalk PII:

S2352-2496(19)30046-1

DOI:

https://doi.org/10.1016/j.fooweb.2019.e00128

Reference:

FOOWEB 128

To appear in:

Food Webs

Received date:

25 October 2019

Revised date:

29 October 2019

Accepted date:

29 October 2019

Please cite this article as: K.L. Jobe, C.G. Montaña and C.M. Schalk, Emergent patterns between salamander prey and their predators, Food Webs(2019), https://doi.org/10.1016/ j.fooweb.2019.e00128

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Journal Pre-proof Emergent patterns between salamander prey and their predators Kasey L. Jobea, Carmen G. Montañaa, and Christopher M. Schalkb,* a

Department of Biology, Stephen F. Austin State University, Nacogdoches, TX, 75962, USA

b

Arthur Temple College of Forestry and Agriculture, Stephen F. Austin State University,

Nacogdoches, TX, 75962, USA

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*Corresponding author. E-mail address: [email protected] (C.M. Schalk)

Journal Pre-proof Abstract. Predator-prey interactions can impose strong selective pressures that scale up to affect the structure and function of communities and ecosystems. Salamanders are prey to a variety of predators that differ in their foraging tactics (e.g., specialists vs. generalists), morphologies, behaviors, and ecologies. Salamanders are an ideal group to explore and quantify patterns that may affect the evolution and persistence of defensive strategies. We examined patterns of predator diversity and body-size relationships of salamander species and their predators by

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compiling a database of observations documenting predator-prey interactions using natural

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history notes published in Herpetological Review (1975-2018), Herpetology Notes (2008-2018)

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and other outside sources. Our database contains 156 salamander predator-prey interactions

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comprised of 69 salamander species that were preyed upon by 89 predator species. Snakes were the most frequently reported predators (35% of predations reported), followed by salamanders

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(24%), and birds (16%). We found that snake predators consistently consumed salamander prey

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that were smaller than themselves, whereas invertebrate predators consumed salamanders across a broader body size range. Snakes are gape-limited predators and unable to eat prey that attain a

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certain size class, whereas invertebrate predators can use other tactics (e.g., use of venom,

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chewing mouthparts) to consume prey larger than themselves. These patterns can serve to help generate hypotheses on the evolution of defensive strategies in salamanders. Key words: Caudata, cannibalism, complex life cycles, defensive strategies, ontogenetic niche shift, predator-prey body size relationships

Journal Pre-proof Predators exert strong selective pressure on prey that can affect different levels of ecological organization, including individuals (e.g., foraging behavior), populations (e.g., abundance and density), communities (e.g., food web structure and function, distribution of species), and ecosystems (e.g., nutrient cycling) (Kenison et al., 2016; Brose et al., 2019). Such adaptive traits may evolve in response to predation pressures, as prey species develop defensive traits to increase their survival and overall fitness (Venesky and Anthony, 2007; Davis and

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Gabor, 2015). For example, gape-constrained predators are unable to consume prey that attain a

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certain size class, thus selection may favor prey attaining larger body sizes (Urban, 2008). Many

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predators avoid this constraint with adaptive feeding mechanisms that allow mastication and

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digestion of prey via various foraging strategies that secure and entrap larger-bodied prey (e.g., venom) (Toledo, 2005; Toledo et al., 2007; Nakazawa et al. 2013; Schalk and Cove 2018).

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The diverse morphologies, ecologies, and life-history strategies of salamanders (Order

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Caudata) makes them model organisms to study predator-prey relationships. Salamanders often reach high abundances representing substantial amounts of biomass in some ecosystems, and

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they are important energetic resources as they are common prey items for many species

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(Hickerson et al., 2017; Davic and Welsh, 2004). Many salamanders have complex life cycles (e.g., biphasic or triphasic stages) that is coupled with a shift in habitat use with each life stage. These habitat shifts expose individuals to different ecological contexts, including shifts in mortality rate-predator regimes (Werner, 1986; Touchon et al. 2013). Vulnerabilities to predators are dependent on a number of factors, including frequency of encounter rates across space or time, prey defenses, and the body sizes of both predator and prey. The interactions of predator type, prey choice, and foraging strategies allow for an opportunity to study these predator-prey dynamics, which could be used to observe the potential defense traits of salamanders.

Journal Pre-proof Salamander predator-prey relationships have contributed greatly to our understanding of the evolution and maintenance of defensive strategies, but much of the knowledge has been produced from intensively studying a few model systems (Brodie and Brodie, 1990; Brodie et al. 2002; Wells 2007). Understanding the relationships and frequencies of predator-prey interactions can be used to develop predictive frameworks of ecological organization across multiple scales (Brose et al. 2019). To determine if broad, generalizable, patterns emerge within this taxon, we

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relationships in instances when salamanders were prey.

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quantified the predator diversity of salamanders and examined patterns of predator-prey size

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Prior studies have synthesized patterns of predator-prey relationships for anuran (Toledo

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et al. 2007), and squamate prey (Schalk and Cove 2018). Therefore, to allow for meaningful comparisons to these other taxa, we use the protocol from these prior studies (Toledo et al. 2007;

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Schalk and Cove 2018) to synthesize data from predation events involving salamanders. We

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reviewed reports when salamanders were preyed upon in articles and natural history notes published in Herpetological Review (vol: 1975 - 2018), Herpetology Notes (vol: 2008 - 2018),

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and other published sources when they contained an unreported predator or salamander family.

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We included salamanders and predators when their taxonomy was reported to the generic level. When reported, we also noted the body size of both predator and prey (i.e., body length [BL] or snout-vent lengths [SVL]), any specialized feeding tactics of the predator (e.g., venom), and instances of cannibalism. We did not include reports of scavenging, experiments, or feeding habits documented in captivity. Taxonomy was updated as reflected in current resources (amphibians: Frost [2019]; birds: Lepage [2019]; mammals: Wilson and Reeder [2005]; reptiles: Uetz et al. [2019]). To ensure our dataset was representative of salamander diversity, we performed a linear regression between salamander familial species richness and the number of

Journal Pre-proof predation events per family (Toledo et al. 2007). We also performed linear regressions using the reported body size data to examine predator-prey size relationships between salamander prey and a subset of predators. We found a significant, positive, relationship between the number of predations reported and familial species richness (R2 = 0.50, P = 0.02, y = 1.08x + 0.65; Fig. 1). Predations were reported from almost every salamander family, except for Proteidae and Hynobiidae. We

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compiled 156 predator-prey interactions representing 69 species of salamanders from 7 families,

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with a total of 89 predator species (Appendices A and B). Overall, snakes (35% of predations

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reported), salamanders (24%), and birds (16%) were the most frequently reported salamander

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predators (Fig. 2). Snakes (30%), birds (15%), and other salamanders (15%) were the most frequently reported predators of adult salamanders. Snakes (26%), other salamanders (22%), and

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insects (19%), were the most frequently reported predators of larval salamander (Fig. 3). When

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salamanders were the reported predator, 56% of the predations were instances of cannibalism. Invertebrate predators consumed salamander prey both larger and smaller than themselves (R² =

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0.27, P = 0.23, y = 0.59x + 0.63) (Fig. 4A), while snakes predators consistently consumed

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salamander prey smaller than themselves (R² = 0.11, P = 0.28, y = 0.38x + 0.86) (Fig. 4B). Prior reviews of anuran, snake, and lizard predators revealed that snakes were the most frequently reported predators of these taxa (Toledo et al., 2007; Schalk and Cove, 2018). We observed a similar pattern, where snakes were the most frequently reported predator of salamanders. Endotherms, such as birds and mammals, are hypothesized to exert stronger selective pressures on ectothermic prey due to their higher ingestion rates resulting from their increased metabolic demands (Greene, 1997). However, endothermic predators are only occasional predators of amphibians, typically encountering and consuming them infrequently,

Journal Pre-proof whereas ectothermic predators, such as snakes, consume amphibians more frequently and with some species even feeding on them exclusively (Toledo et al. 2007). Indeed, the Garter Snake (Thamnophis sirtalis) and Rough-Skinned Newts (genus Taricha) predator-prey relationship has emerged as a model system to examine the coevolution of predator-prey arms races (Brodie and Brodie, 1990; Brodie et al., 2002; Geffeney et al., 2002). The second most common predator reported were salamanders, with most of reports being cannibalistic interactions. Therefore, as a

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whole, snakes and salamanders may be exerting stronger selective pressures on the evolution of

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salamander defensive strategies (sensu Toledo et al., 2007). For example, individuals may

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increase foraging activity to more rapidly attain a size refuge to eliminate predation risk from

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gape-constrained predators such as snakes or salamanders (Urban, 2007b). Due to their morphological and physiological constraints, movement (i.e., dispersal) of

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salamanders are restricted, resulting in isolated populations that must adapt to local pressures

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(Tilley, 2016; Kazitsa et al., 2018). However, even if restricted geographically, the complex life cycles of some species expose them to different pressures due to shifting habitats, predator

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encounter rates, and size- or stage-specific vulnerabilities to mortality (Werner, 1986; Touchon

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et al. 2015; Schalk et al. 2018). For example, insects, crayfish, and other salamanders were more frequently reported as predators of larval salamanders, while birds and mammals were more frequently reported as predators of adult salamanders. If metamorphosis coincides with a shift in habitat, it may expose individuals to a higher diversity of predators across the lifetime of the salamander. A simplified life cycle (e.g., some species in the family Plethodontidae and obligate paedomorphic species) does not consist of a habitat shift during ontogeny and may potentially reduce their exposure to a greater diversity and abundance of predators. However, larval salamanders often use ephemeral aquatic habitats that vary in their community composition (e.g.,

Journal Pre-proof diversity or abundance of predators) across space or time. Many salamanders are capable of producing context-dependent phenotypes in response to these heterogeneous environments, which can be effective at reducing an individual’s predation risk (Van Buskirk and Schmidt, 2000; Bernard 2004; Van Buskirk 2009). Snakes were observed to prey upon salamanders that were smaller relative to their body size. As predators, snakes are incapable of consuming prey that exceed their gape width as they

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are unable to masticate their prey. This gape-limited predation on smaller-bodied prey items

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allows larger individuals to survive and reproduce within a population. This directional selection

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leads to defensive mechanisms to evolve against gape-constrained predators (Urban 2007b,

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2008). Invertebrate predators, however, preyed upon salamanders that were more variable in size, as the salamander prey were both larger and smaller relative to the body size of the

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invertebrate predators. Many invertebrate predators utilize a diversity of feeding mechanisms

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(e.g., digestive venom, chewing mouthparts) or specialized foraging tactics (e.g., entrapping prey in webs) which allow for increased rate of capture success and consumption of a broader size

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range of prey (Toledo, 2005; Nakazawa et al., 2013; Schalk and Cove 2018). However, the

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abundance and frequency of these two broad predator classes in the environment affects the adaptive reaction norms of prey populations. As prey that may escape mortality from gapelimited predators may not escape predation risk from gape-unconstrained predators (Urban, 2007a). Our review provides broad patterns on the diversity of salamander predators across life stages and size-specific predator-prey size relationships. However, these results present patterns that can help formulate hypotheses on the evolutionary trajectory of salamander defensive strategies. For example, testing whether the frequency and intensity of predation pressure is

Journal Pre-proof correlated with fixation of defenses (e.g., production predator-specific noxious compounds; sensu Adams et al. 2011). Alternatively, future studies could test whether a salamander species vulnerable to a broader suite of predators or ecological contexts also have a more extensive defensive repertoire in the form of predator-specific plastic defenses (Van Buskirk and Schmidt, 2000; Relyea 2004; Schalk 2016). In turn, exploring the costs of these defenses within each life stage or for those species with complex life cycles, the carryover effects of the costs of these

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defenses on subsequent life stages (Hagman et al. 2009; O'Connor et al. 2014).

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Acknowledgements

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We thank all our colleagues that reported these valuable natural history observations. We would

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also like to thank C.S. Adams, S.E. Ebert, K. Ireland, K. Kralman, C.A. Layman, D. Saenz, K. Untiedt, and K.J. Ward for their assistance and constructive comments on the manuscript.

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Funding Sources

or not-for-profit sectors.

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Declarations of interest

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This research did not receive any specific grant from funding agencies in the public, commercial,

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None. Conflicts of interest None.

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Appendix A Predator Class Actinopt erygii Actinopt erygii Actinopt erygii Actinopt erygii Actinopt erygii Actinopt erygii Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a

Predator Order Anguillifor mes Cyprinifor mes

Predator Family

Predator Species

Anguillidae

Anguilla rostrata

Cyprinidae Centrarchid ae

Semotilus atromaculatus

Salmonidae

Salvelinus fontinalis

Ictaluridae

Ictalurus furcatus

Ictaluridae

Ictalurus furcatus

Anura

Ranidae

Rana aurora

Anura

Ranidae

Lithobates catesbeianus

Anura

Ranidae

Caudata

Cryptobran chidae

Lithobates catesbeianus Cryptobranchus alleganiensis alleganiensis

Anura

Ranidae

Lithobates sylvaticus

Anura

Ranidae

Lithobates sylvaticus

Anura

Ranidae

Lithobates catesbeianus

Perciformes Salmonifor mes Siluriforme s Siluriforme s

Lepomis cyanellus

l a

n r u

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Salamande r Family Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Ambystoma tidae Salamandrid ae Ambystoma tidae

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Salamander Species Eurycea bislineata bislineata

Plethodontid ae Ambystoma tidae Ambystoma tidae Ambystoma tidae

Reference Russell and Campbell, 2011

Plethodon cinereus

Sharp et al., 2005

Eurycae waterloonesis Desmognathus quadramaculatus

Devitt et al., 2016

Plethodon glutinosus

Sorrell et al., 2005

Plethodon glutinosus Ambystoma californiense Taricha torosa

Sorrell et al., 2005 Baldwin and Stanford, 1987 Jennings and Cook, 1998

Ambystoma gracile

Rowe and Garcia, 2012

Desmognathus quadramaculatus Ambystoma laterale Ambystoma maculatum Ambystoma californiense

Ennen et al., 2015

Hill, 2011 Baldwin and Calhoun, 2002 Baldwin and Calhoun, 2002 Balfour and Stitt, 2003

Journal Pre-proof Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi

Anura Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata Caudata

Ranidae Plethodonti dae Ambystom atidae Salamandri dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae Ambystom atidae

Rana aurora

Ambystoma mavortium

Rhyacotrito nidae Plethodontid ae Ambystoma tidae Salamandrid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Ambystoma tidae

Proteidae Cryptobran chidae Cryptobran chidae Salamandri

Necturus maculosus Cryptobranchus alleganiensis Cryptobranchus alleganiensis Taricha torosa

Proteidae Plethodontid ae Cryptobranc hidae Salamandrid

Plethodon dunni Dicamptodon ensatus Triturus marmoratus Demognathus fuscus Desmognathus quadramaculatus Gyrinophilus porphyriticus

l a

rn

Pseudotriton ruber

u o

Pseudotriton ruber

J

Desmognathus monticola

Rabinowe et al., 2002

Plethodon dunni

Altig and Brodie, 1971

Dicamptodon ensatus

Anderson, 1960

f o

Lissotriton boscai

o r p

e

r P

Desmognathus monticola Desmognathus carolinensis

Rhyacotriton kezeri

Ayres 2007

Demognathus fuscus Desmognathus monticola

Baldauf, 1947 Bernardo and Yoke 2006

Desmognathus ocoee

Bernardo, 2002

Desmognathus wrighti

Bernardo, 2002

Plethodon welleri

Bernardo, 2002

Plethodon cinereus

Bock and Fauth, 1992

Plethodon cinereus Desmognathus monticola

Bock and Fauth, 1992

Ambystoma mavortium

Collins, 1981

Necturus maculosus

Eycleshymer, 1906

Eurycea bislineata Cryptobranchus alleganiensis Taricha torosa

Greathouse et al., 2016 Groves and Williams, 2014 Hanson et al. 1994

Camp, 1997

Journal Pre-proof a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a

Caudata

dae Plethodonti dae Cryptobran chidae Cryptobran chidae Plethodonti dae Plethodonti dae Plethodonti dae Salamandri dae

Caudata

Proteidae

Necturus maculosus

Caudata

Sirenidae Plethodonti dae Plethodonti dae Plethodonti dae Amphiumid ae Salamandri dae Salamandri dae Salamandri dae

Siren lacertina

Caudata Caudata Caudata Caudata Caudata Caudata

Caudata Caudata Caudata Caudata Caudata Caudata Caudata

Plethodon cinereus Cryptobranchus alleganiensis Cryptobranchus alleganiensis Eurycea bislineata Desmognathus welteri Desmognathus welteri Taricha torosa

l a

n r u

Plethodon cinereus Desmognathus ochrophaeus

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Plethodon albagula Amphiuma means Notophthalmus viridescens Triturus marmoratus Taricha granulosa

ae Plethodontid ae Plethodontid ae Cryptobranc hidae Plethodontid ae Plethodontid ae Plethodontid ae Salamandrid ae

Plethodon cinereus Eurycea sp. Desmognathus quadramaculatus

Proteidae Ambystoma tidae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Ambystoma tidae Salamandrid ae Ambystoma tidae

Humphries et al., 2005

Eurycea bislineata

Hutton and Price, 2018

Desmognathus welteri

Hutton et al., 2018

Desmognathus welteri Taricha torosa

Hutton et al., 2018 Kaplan and Shermn, 1980

Necturus maculosus

Lockhart, 2000

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-p

e r P

f o

Heatwole and Test, 1961 Hecht-Kardaz and Nickerson, 2013

Ambystoma talpoideum Luhring, 2007 Maerz and Karuza, Plethodon cinereus 2003 Desmognathus ochrophaeus Martof and Rose, 1963 Plethodon albagula

Milanovich et al. 2007

Eurycea cirrigera

Montaña et al., 2014

Ambystoma tigrinum Ichthyosaura alpestris

Morin, 1983 Orizaola and Rodriguez del Valle, 2000

Ambystoma gracile

Pearl, 2003

Journal Pre-proof Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Amphibi a Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves

Caudata Caudata Caudata Caudata Caudata Caudata Ciconiiform es Passeriform es Strigiforme s Strigiforme s Accipitrifor mes Accipitrifor mes Accipitrifor mes Accipitrifor mes Anseriform es Anseriform es Charadriifo

Salamandri dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae Plethodonti dae

Notophthalmus viridescens

Ardeidae

Ardea herodias

Turdidae

Turdus migratorius

Strigidae

Bubo virginianus

Strigidae Accipitrida e

Strix varia

Plethodon glutinosus Pseudotriton ruber ruber Plethodon dunni Plethodon dunni Gyrinophilus porphyriticus

l a

Buteo lineatus

J

Cathartidae Accipitrida e Accipitrida e

Cathartes aura

Anatidae

Lophodytes cucullatus

Anatidae Recurvirost

Anas platyrhynchos Recurvirostra americana

Buteo lineatus Buteo lineatus

Notophthalmus viridescens

Pitt et al. 2011

Plethodon glutinosus Eurycea longicauda longicauda

Powders, 1973 Reblin and Anthony, 2001

f o

Plethodon dunni

o r p

e

r P

rn

u o

Salamandrid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Salamandrid ae Ambystoma tidae Plethodontid ae Ambystoma tidae Ambystoma tidae Sirenidae Ambystoma tidae Amphiumid ae Sirenidae Salamandrid ae Salamandrid ae Ambystoma

Plethodon dunni Notophthalmus viridescens viridescens

Riesecrer et al. 1996

Plethodon idahoensis

Riesecrer et al. 1996 Secki and Queral-Regil, 1997 Brodman and Pfingsten, 2010 Wilson and Simon, 1985

Ambystoma tigrinum

Holman, 1976

Ambystoma maculatum Siren intermedia nettingi

Jongmsa et al., 2012

Ambystoma mavortium

Duncan, 1999

Amphiuma means

Herman et al., 2016

Siren lacertina Taricha granulosa

Herman et al., 2016 Mebs and YotsuYamashita, 2017

Taricha granulosa Ambystoma

Rombough, 2017 Allaback et al., 2005

Ambystoma tigrinum

Dobbs and Dobbs, 2003

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Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Aves Chilopod a

rmes Charadriifo rmes Ciconiiform es Coraciiform es Passeriform es Passeriform es Passeriform es Passeriform es Passeriform es Pelecanifor mes Pelecanifor mes Pelecanifor mes Pelecanifor mes Podicipedif ormes Strigiforme s Strigiforme s Scolopendr omorpha

ridae Laridae

Sterna forsteri

Ardeidae

Botaurus lentiginosus

Alcedinidae Megaceryle alcyon Icteridae

Quiscalus quiscala

Turdidae

Turdus migratorius

Laniidae

Lanius ludovicianus

Icteridae

Quiscalus quiscula

Corvidae

Aphelocoma californica

Ardeidae

Ardea herodias

Ardeidae

Ardea herodias

l a

n r u

tidae Ambystoma tidae

californiense Ambystoma californiense

Allaback et al., 2005

Sirenidae Salamandrid ae Ambystoma tidae Plethodontid ae Ambystoma tidae Ambystoma tidae Plethodontid ae

Siren lacertina Notophthalmus viridescens

Beane and Pusser, 2013 Tanaka and Means, 2017

Ambystoma maculatum Plethodon cinereus

Duncan, 2005 Fenster and Fenster, 1996

Ambystoma tigrinum

Jensen, 2003

Ambystoma texanum

Mounce, 2004

Aneides lugubris Siren intermedia nettingi

Rubinoff, 1996

o r p

e

r P

f o

Strigidae

Strix varia

Sirenidae Salamandrid ae Taricha granulosa Siren intermedia Sirenidae nettingi Salamandrid Notophthalmus ae viridescens Salamandrid ae Taricha granulosa Amphiumid ae Amphiuma means

Strigidae Scolopocry ptopidae

Strix varia Scolopocryptops sexspinosus

Sirenidae Siren sp. Plethodontid ae Eurycea quadridigitata

Ardeidae

Jo

Ardea alba

Ardeidae Egretta caerulea Podicipedid ae Podilymbus podiceps

Dobbs and Dobbs, 2003

Luhring et al. 2013

Fellers et al. 2007 Frese and Britzke, 2001 Tanaka and Means, 2017 Mcallister et al. 1997 Beane, 2005

Palis, 2011

Journal Pre-proof

Insecta

Staphylinid ae

Insecta

Coleoptera Hymenopte ra

Insecta

Coleoptera

Carabidae

Insecta

Coleoptera

Insecta

Hemiptera

Carabidae Scarites subterraneus Belostomati dae Lethocerus uhleri

Insecta

Mantodea

Insecta Malacost raca Malacost raca Malacost raca Mammal ia Mammal ia Mammal ia Mammal ia Mammal ia Mammal ia Mammal

Trichoptera

Mantidae Phryganeid ae

Decapoda

Cambaridae Procambarus sp.

Decapoda

Cambaridae Procambarus clarkii

Isopoda Artiodactyl a

Asellidae

Cingulata

Formicidae

Platydracus viduatus Lasius umbratus Carabus (Eucarabus) deyrollei

Choeradodis strumaria Ptilostomis sp.

l a

rn

Lirceus fontinalis

u o

Suidae Sus scrofa domestica Dasypodida e Dasypus novemcinctus

Carnivora Soricomorp ha Artiodactyl a

Canidae

Carnivora Carnivora

J

Canis latrans

Plethodontid ae Plethodontid ae Salamandrid ae Ambystoma tidae Salamandrid ae Plethodontid ae Ambystoma tidae Plethodontid ae Plethodontid ae Ambystoma tidae Ambystoma tidae Ambystoma tidae

Jung et al., 2000

Chioglossa lusitanica Ambystoma jeffersonianum Notophthalmus perstriatus

Galan 2007

Bolitoglossa rufescens

Buttenhoff, 1995

Ambystoma maculatum

Murphy, 1961

Eurycea quadridigitata

Franklin, 2000

Eurycea sosorum

Owen et al., 2016

Ambystoma barbouri Ambystoma californiense

Hutton and Price, 2018

f o

o r p

e

r P

Plethodon cinereus Desmognathus ochrophaeus

Soricidae

Blarina carolinensis

Suidae

Sus scrofa

Sirenidae Ambystoma tidae Salamandrid ae

Mustelidae Mustelidae

Neovison vison Neovison vison

Sirenidae Ambystoma

Forester, 1979

Smith, 1946 Jensen, 2017

Wilcox et al. 2008

Ambystoma talpoideum Croshaw 2007 Siren intermedia

Palis, 2013

Ambystoma opacum Salamandra salamandra Siren intermedia nettingi Ambystoma mavortium

Palis, 2011 Carretero and Rosell, 1999 Frese and Britzke, 2001 Fronzuto, 1997

Journal Pre-proof ia Mammal ia Mammal ia

tidae Carnivora

Mustelidae

Neovison vison

Carnivora

Mustelidae

Reptilia

Squamata

Natricidae

Lutra lutra Thamnophis atratus atratus

Reptilia

Squamata

Natricidae

Nerodia sipedon

Reptilia

Squamata

Natricidae

Nerodia sipedon

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Colubridae

Reptilia

Squamata

Dipsadidae

Nerodia sipedon Thamnophis sirtalis sirtalis Farancia abacura reinwardtii

Reptilia

Squamata

Dipsadidae

Rhadinella lachrymans

Reptilia

Squamata

Viperidae

Reptilia

Squamata

Viperidae

Reptilia

Squamata

Dipsadidae

Crotalus oreganus Diadophis punctatus punctatus

Reptilia

Squamata

Elapidae

Micrurus nigrocinctus

Reptilia

Squamata

Colubridae

Diadophis punctatus

Reptilia

Squamata

Viperidae

Sistrurus miliarius

l a

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Agkistrodon piscivorus

Proteidae Salamandrid ae Salamandrid ae Ambystoma tidae Plethodontid ae Ambystoma tidae Ambystoma tidae Plethodontid ae

Necturus maculosus

Ambystoma texanum

Kats, 1986

Pseudotriton montanus

Kats, 1986

Ambystoma gracile

Lefcort et al. 1991

Ambystoma texanum

MaCallum, 1995

Triturus cristatus

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Taricha granulosa

Little, 2002 Bringøes and Nørgaard, 2018 Greene and Feldman, 2009

Plethodon nettingi Siren intermedia Sirenidae nettingi Plethodontid Bolitoglossa ae engelhardti Ambystoma tidae Ambystoma opacum Plethodontid ae Aneides lugubris Plethodontid ae Eurycea lucifuga Plethodontid ae Bolitoglossa striatula Plethodontid ae Plethodon glutinosus Plethodontid Hemidactylium ae scutatum

Messenger et al., 2011 Shively, 2010 Quijano and Rojas, 1995 Kuhns et al., 2018 Mahrdt and Banta, 1997 Nelson et al., 2010 Núñez, 2010 Smith and Robertson, 2018 Trauth and Cochran, 1991

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Reptilia

Squamata

Dipsadidae

Diadophis punctatus

Reptilia

Squamata

Dipsadidae

Rhadinaea fulvivittis

Reptilia

Squamata

Colubridae

Thamnophis atratus

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Natricidae

Natrix maura

Reptilia

Squamata

Colubridae

Thamnophis cyrtopsis

Reptilia

Squamata

Dipsadidae

Heterodon platirhinos

Reptilia

Squamata

Natricidae

Nerodia sipedon sipedon

Reptilia

Squamata

Dipsadidae

Diadophis punctatus

Reptilia

Squamata

Natricidae

Nerodia sipedon

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Dipsadidae

Farancia abacura

Reptilia

Squamata

Dipsadidae

Farancia abacura

Reptilia Reptilia

Squamata Squamata

Natricidae Colubridae

Nerodia fasciata Thamnophis sirtalis

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Thamnophis hammondii Thamnophis sauritus sauritus

Plethodon hoffmani Pseudoeurycea nigromaculata Ambystoma californiense

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Eurycea longicauda

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Plethodontid ae Plethodontid ae Ambystoma tidae Plethodontid ae Plethodontid ae Salamandrid ae Ambystoma tidae Salamandrid ae Plethodontid ae Plethodontid ae Plethodontid ae Plethodontid ae Ambystoma tidae Amphiumid ae Amphiumid ae Ambystoma tidae Plethodontid

Adams, 1999 Alberto et al., 2010 Alvarez and Shea, 2017 Anthony et al. 2007

Eurycea longicauda

Anthony et al. 2007

Lissotriton boscai

Ashega, 2012

Ambystoma velasci Notophthalmus viridescens Gyrinophilus porphyriticus

Banda-Leal et al. 2014

Eurycea lucifuga Desmognathus quadramaculatus

Camp et al. 2015

Barnett et al. 2006 Blackburn et al., 2003

Crawford 2007

Batrachoseps major

Ervin et al., 2003 Greene and Boback, Ambystoma talpoideum 2003 Amphiuma means

Haertle et al., 2015

Amphiuma tridactylum

Haertle et al., 2015

Ambystoma texanum Plethodon

Hampton 2007 Hofmann and

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Reptilia

Squamata

Dipsadidae

Diadophis punctatus Nerodia sipedon insularum

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Colubridae

Reptilia

Squamata

Natricidae

Elpahe quadrivirgata Thamnophis sirtalis sirtalis

Reptilia

Squamata

Natricidae

Thamnophis sumichrasti

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Boidae

Charina bottae

Reptilia

Squamata

Colubridae

Natrix natrix

Reptilia

Squamata

Dipsadidae

Reptilia

Squamata

Dipsadidae

Rhadinaea taeniata Diadophis punctatus stictogenys

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Natricidae

Thamnophis elegans

Reptilia

Squamata

Natricidae

Thamnophis elegans

Reptilia

Squamata

Dipsadidae

Diadophis punctatus

Reptilia

Squamata

Dipsadidae

Diadophis punctatus

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Thamnophis sirtalis Thamnophis ordinoides

ae chlorobryonis Salamandrid ae Taricha torosa

Hofmann, 2018

Proteidae Salamandrid ae Plethodontid ae Plethodontid ae Ambystoma tidae Plethodontid ae Salamandrid ae Plethodontid ae Plethodontid ae Plethodontid ae Rhyacotrito nidae Plethodontid ae Plethodontid ae Ambystoma tidae Plethodontid ae

Necturus maculosus

King et al., 1999

Cynops pyrrhogaster

Kojima, 2017 Konavalinka and Trauth, 2003 Lemos Espinal et al., 2000 Loafman and Jones, 1996

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Plethodon albagula Pseudoeurycea leprosa Dicamptodon copei Ensatina eschscholtzii Salamandra salamandra

Kats et al., 1998

Isthmura bellii

Macey, 1983 Maxinova and Marcel Uhrin, 2018 Mendoza-Quijano et al., 1995

Plethodon albagula

Milanovich et al., 2005

Plethodon vehiculum Rhyacotriton variegatus

Norman, 1988

Aneides hardii Plethodon neomexicanus

Painter et al., 1999

Norman, 2002

Painter et al., 1999

Ambystoma talpoideum Palis, 2016 Desmognathus abditus

Paulson et al., 2015

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Reptilia

Squamata

Natricidae

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Natricidae

Reptilia

Squamata

Colubridae

Reptilia

Squamata

Colubridae

Nerodia floridana Thamnophis sirtalis sirtalis

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Natricidae

Thamnophis sirtalis

Reptilia

Squamata

Reptilia

Testudines

Reptilia

Testudines

Natricidae Chelydrida e Chelydrida e

Thamnophis couchii Chelydra serpentina serpentina Chelydra serpentina serpentina

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Plethodontid ae Salamandrid ae Salamandrid ae Ambystoma tidae Ambystoma tidae Plethodontid ae Plethodontid ae Plethodontid ae Salamandrid ae Ambystoma tidae Ambystoma tidae

Poteet and Bell, 1999

Taricha granulosa

Poteet and Bell, 1999

Taricha granulosa

Poteet and Bell, 1999

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Ambystoma opacum Ambystoma jeffersonianum

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Plethodon dunni

Vogrinc et al., 2013

Ambystoma mavortium

Williams et al., 2013 Wilson and Wilson, 1996 Wilson and Wilson, 1996 Wilson and Wilson, 1996 Wiseman and Pool, 2007 Lewis and Iverson, 2018

Ambystoma maculatum

Spohr, 2005

Plethodon idahoensis Plethodon idahoensis Plethodon idahoensis Taricha torosa

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Appendix B. References of salamander predations. Adams, D.C. 1999. Plethodon hoffmani (Valley and Ridge Salamander). Predation. Herpetological Review. 30 (3): 160. Alberto, A., M. Hernández, U.O. García-Vázquez. 2010. Pseudoeurycea nigromaculata (Blackspotted Salamander). Predation. Herpetological Review 41 (2): 191. Allaback, M.L., D.M. Laabs, S.E. Higgins, T.P. Winfield. 2005. Ambystoma californiense (California Tiger Salamander). Predation. Herpetological Review. 36 (1): 50.

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Altig, R., & Brodie Jr, E. D. 1971. Foods of Plethodon larselli, Plethodon dunni and Ensatina eschscholtzi in the Columbia River Gorge, Multnomah County, Oregon. American Midland Naturalist. 85 (1): 226-228.

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Alvarez, J. A., M. A. Shea. 2017. Ambystoma californiense (California Tiger Salamander). Predation. Herpetological Review. 48(3): 596 Anderson, J.D. 1960. Cannibalism in Dicamptodon ensatus. Herpetologica 16 (4):260.

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Anthony, C.D., M.D. Venesky, J.C. Spetz. 2007. Eurycea longicauda longicauda (Long-tailed Salamander). Predation. Herpetological Review. 38 (2): 175-176.

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Ashega, C.A. 2012. Lissotriton boscai (Viperine Snake). Diet: Scavenging. Herpetological Review. 43 (2): 346-347. Ayres, C. 2007. Triturus marmoratus (Marbled Newt). Newt Predation. Herpetological Review. 38 (4): 434.

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Baldauf, R. 1947. Desmognathus f. fuscus eating eggs of its own species. Copeia 1947:66.

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Baldwin, K.S., R.A. Stanford. 1987. Ambystoma tigrinum californiense (California Tiger Salamander). Predation. Herpetological Review 18 (2): 33. Baldwin, R.F., A.J.K. Calhoun. 2002. Ambystoma laterale (Blue-spotted Salamander) and Ambystoma maculatum (Spotted Salamander). Predation. Herpetological Review. 33 (1): 44-45. Balfour, P.S., E.W. Stitt. 2003. Ambystoma californiense (California Tiger Salamander). Predation. Herpetological Review. 34 (1): 44. Banda-Leal, J., D. Lazcano, M. Nevárezde Los Reyes, C. Barriga Vallejo, C. Aguilera, S.A. Luna-Peña. 2014. Ambystoma velasci (Plateau Tiger Salamander). Predation. Herpetological Review. 45(4): 675-676. Barnett, K.E., A. Breisch, J. Bruchac, B. Abbuhl. 2006. Heterodon platirhinos (Eastern Hognose Snake). Regurgitation of Prey. Herpetological Review. 37 (3): 352. Beane, J.C. 2005. Amphiuma means (Two-toed Amphiuma). Predation. Herpetological Review. 36 (3): 295.

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Beane, J.C., L.T. Pusser. 2013. Siren lacertina (Greater Siren) predation. Herpetological Review. 44 (3): 492. Bernardo, J. 2002. Desmognathus carolinensis (Carolina Mountain Dusky Salamander) and Plethodon welleri (Weller's Salamander); Desmognathus monticola (Seal Salamander) and Desmognathus wrighti (Pygmy Salamander). Gyrinophilus porphyriticus (Spring Salamander) and Desmognathus ocoee (Ocoee Salamander). Intraguild Predation. Herpetological Review. 33 (2): 121.

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Bernardo, J., M.M. Yoke. 2006. Desmognathus monticola (Seal Salamander) and Desmognathus quadramaculatus (Black-bellied Salamander). Intraguild Predation (IGP) and Autourophagy. Herpetological Review. 37 (4): 436.

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Blackburn, J.L., W.D. Brown, J.C. Mitchell. 2003. Nerodia sipedon sipedon (Northern Watersnake). Diet. Herpetological Review. 34 (4): 377.

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Bock, S.F., J.E. Fauth. 1992. Pseudotriton ruber ruber (Northern Red Salamander). Diet. Herpetological Review. 23(2): 58. Bringsøes, H., J. Nørgaard. 2018. Predation of Triturus cristatus (Caudata: Salamandridae) by the Eurasian Otter (Carnivora: Mustelidae). Predation. Herpetology Notes. 11: 279-280. Brodman, R., R.A. Pfingsten. 2010. Ambystoma tigrinum (Tiger Salamander). Predation. Herpetological Review 41 (2): 186. Buttenhoff, P. 1995. Bolitoglossa rufescens (Northern Banana Salamander). Predation. Herpetological Review 26 (4): 197. Camp, C.D. 1997. Desmognathus monticola (Seal Salamander). Oophagy. Herpetological Review 28 (2): 81-82. Carreterro, M.A., C. Rosell. 1999. Salamandra salamandra (Fire Salamander). Predation. Herpetological Review. 30 (3): 161.

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Crawford, J.A. 2007. Desmognathus quadramaculatus (Black-bellied Salamander). Predation. Herpetological Review. 38 (4): 432-433. Croshaw, D.A. 2007. Ambystoma talpoideum (Mole Salamander). Predation. Herpetological Review. 38 (1): 66-67. Dobbs, R.C., D.M. Dobbs. 2003. Siren intermedia nettingi (Western Lesser Siren). Predation. Herpetological Review. 34 (1): 47. Dobbs, R.C., D.M. Dobbs. 2003. Siren intermedia nettingi (Western Lesser Siren). Predation. Herpetological Review. 34 (1): 47. Duncan, R.B. 1999. Ambystoma tigrinum stebbinsi (Sonoran Tiger Salamander). Predation. Herpetological Review. 30 (3): 159. Duncan, R.B. 2005. Ambystoma maculatum (Spotted Salamander). Predation. Herpetological Review. 36 (3): 294.

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Ennen, J.R., B.R. Kuhajda, K.F. Alford, S.C. Hazzard. 2015. Desmognathus quadramaculatus (Black-bellied Salamander). Predation. Herpetological Review. 46 (2): 226-227. Fellers, G.M., J. Reynolds-Taylor, S. Farrar. 2007. Taricha granulosa (Rough-skinned Newt). Predation. Herpetological Review. 38 (3): 317-318. Eycleshymer, A. C. 1906. The habits of Necturus maculosus. American Naturalist. 40:123-136.

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Fenster, T.L., C.B. Fenster. 1996. Plethodon cinereus (Redback Salamander). Predation. Herpetological Review 27 (4): 194. Forester, D.C. 1979. Observations of ant predation upon brooding salamanders. Herpetological Review 10 (1): 3-4. Franklin, C.J. 2000. Eurycea quadradigitata (Dwarf Salamander). Larval Predation. Herpetological Review. 31 (2): 98.

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Frese, P.W., E. Britzke. 2001. Siren intermedia nettingi (Western Lesser Siren). Predation. Herpetological Review. 32 (2): 99.

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Fronzuto, J.A. 1997. Ambystoma tigrinum melanostictum (Blotched Tiger Salamander). Predation. Herpetological Review 28 (2): 81. Franklin, C.J. 2000. Eurycea quadradigitata (Dwarf Salamander). Larval Predation. Herpetological Review. 31 (2): 98.

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Gálan, P. 2007. Chioglossa lusitanica (Golden Striped Salamander). Predation. Herpetological Review. 38 (2): 173-174.

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Greathouse, W. J., C. Byrne, K. Samborsky, A. S. Wendt, S. S. Klein. 2016. Eurycea bislineata (Northern Two-lined Salamander). Predation. Herpetological Review. 47(1): 98 Greene, M.P., S.M. Boback. 2003. Thamnophis sauritus sauritus (Eastern Ribbon Snake). Diet. Herpetological Review. 34 (2): 159. Greene, R.R., C.R. Feldman 2009. Thamnophis atratus atratus (Santa Cruz Gartersnake). Diet. Herpetological Review. 40 (1): 103-104. Groves, J.D., L.A. Williams. 2014. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender). Cannibalism. Herpetological Review. 45: 108-109. Haertle, N.E., P.M. Hampton, P.N. Vogrinc, J.D. Willson. 2015. Farancia abacura (Red-bellied Mudsnake). Feeding Behavior. Herpetological Review. 46 (3): 449-450. Hampton, P.M. 2007. Nerodia fasciata confluens (Broad-banded Watersnake). Diet. Herpetological Review. 38 (1): 90. Hanson, K., J. Snyder, L. Kats. 1994. Taricha torosa (California Newt). Diet. Herpetological Review 25 (2): 62. Heatwole, H., F.H. Test. 1961. Cannibalism in the salamander, Plethodon cinereus. Herpetologica. 17(2): 143-143.

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Hecht-Kardasz, K., M.A. Nickerson. 2013. Cryptobranchus alleganiensis (Hellbender). Larval Diet. Herpetological Review. 44 (3): 490. Herman, J. E., W. Brosse, C. U. Whelan, K. R. T. Whelan, S. Clem, M. Rumbach. 2016. Siren lacertina (Greater Siren) and Amphiuma means (Two-toed Amphiuma). Predation. Herpetological Review. 47(3): 432-433 Hill, R.L. 2011. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender) and Desmognathus quadramaculatus (Black-bellied Salamander). Predator/Prey. Herpetological Review 42 (4): 580. Hoffman, E., P. Hoffman. 2018. Plethodon chlorobryonis (Atlantic Coast Slimy Salamander). Predation. Herpetological Review. 49(1): 91 Holman, J.A. 1976. Owl predation on Ambystoma tigrinum. Herpetological Review 7 (3): 114. Humphries, W.J., M. Solis, C. Caldwell, A. Salveter. 2005. Cryptobranchus alleganiensis (Hellbender). Cannibalism. Herpetological Review. 36 (4): 428.

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Hutton, J., S. Price. 2018. Ambystoma barbouri (Streamside Salamander). Predation Herpetological Review. 49(1): 89 Hutton, J., S. Price. 2018. Eurycea bislineata (Two-lined Salamander). Predation. Herpetological Review. 49(1): 90-91 Hutton, J., R. Leloudis, S. Price. 2018. Desmognathus welteri (Black Mountain Salamander) and Demognathus welteri (Black Mountain Salamander). Predation. Herpetological Review. 49(2): 296 Jennings, M.R., D. Cook. 1998. Taricha torosa torosa (Coast Range Newt). Predation. Herpetological Review 29 (4): 230. Jensen, J.B. 2003. Ambystoma tigrinum (Tiger Salamander). Predation. Herpetological Review. 34 (2): 132-133.

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Jensen, J. B. 2017. Notophthalmus perstriatus (Striped Newt). Predation. Herpetological Review. 48(4): 828 Jongmsa, G.F., D.F. McAlpine, B. Cougle, B. Forsythe. 2012. Ambystoma maculatum (Spotted Salamander). Predation. Herpetological Review. 43 (4): 626-627. Jung, R.E., W.L. Ward, C.O. King, L.A. Weir. 2000. Plethodon cinereus (Redback Salamander). Predation. Herpetological Review. 31 (2): 98-99. Kaplan, R.H., Sherman, P.W. 1980. Intraspecific oophagy in California newts. Journal of Herpetology. 14(2): 183-185. Kats, L.B. 1986. Nerodia sipedon (Northern Water Snake). Feeding. Herpetological Review. 17 (3): 61-64. Kats, L.B., J.A. Goodsell, N. Matthews, C. Barn, A.R. Blaustein. 1998. Taricha torosa (California Newt). Predation. Herpetological Review 29 (4): 230. King, R.B., A. Qurealregil, T.D. Bittner, J.M. Kerfin, J. Hageman. 1999. Nerodia sipedon insularum (Lake Erie Watersnake). Diet. Herpetological Review. 30 (3): 169-170.

Journal Pre-proof Kojima, Y. 2017. Cynops pyrrhogaster (Japanese Firebelly Newt). Predation. Herpetological Review. 48(1): 211 Konavalinka, R.R., S.E. Trauth. 2003. Thamnophis sirtalis sirtalis (Eastern Garter Snake) and Plethodon albagula (Western Slimy Salamander). Predation. Herpetological Review. 34 (4): 378.

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Kuhns, A., C. Phillips, M. Dreslik, M. Corcoran, J. Crawford. 2018. Ambystoma opacum (Marbled Salamander). Predation. Herpetological Review. 49(2): 332-333 Lefcort, H., G. Hokit, J.J. Beatty, C. Hakelev. 1991. Thamnophis sirtalis fitchi (Valley Garter Snake). Feeding Behavior. Herpetological Review 22 (3): 101. Lemos Espinal, J.A., A. Ramírez Bautista, G. Woolrich Pisia, J.E. Gonzalez Espinoza. 2000. Thamnophis sumichrasti (Sumichrast's Garter Snake). Prey. Herpetological Review. 31 (4): 248249.

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Little, M.J. 2002. Necturus maculosus (Mudpuppy). Predation. Herpetological Review. 33 (2): 122.

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Loafman, P. L. Jones. 1996. Dicamptodon copei (Cope's Giant Salamander). Metamorphosis and Predation. Herpetological Review 27 (3): 136. Lockhart, O.M. 2000. Necturus maculosus (Mudpuppy). Egg Predation. Herpetological Review. 31 (2): 98. Luhring, T.M. 2007. Siren lacetina (Greater Siren). Diet. Herpetological Review. 38 3): 317.

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Luhring, T.M., J.W. Gibbons, P.W. Gibbons. 2013. Siren sp. predation. Herpetological Review. 44 (3): 491-492. Macey, R.J. 1983. Charina bottae bottae (Pacific Rubber Boa). Food. Herpetological Review. 14 (1): 19. Maerz, J.C., J. Karuzas. 2003. Plethodon cinereus (Eastern Red-backed Salamander). Cannibalism. Herpetological Review. 34 (1): 46-47. Mahrdt, C.R., B.H. Banta. 1997. Aneides lugubris (Arboreal Salamander). Predation. Herpetological Review 28 (2): 81. Martof, B. S., Rose, F. L. (1963). Geographic variation in southern populations of Desmognathus ochrophaeus. American Midland Naturalist. 69(2): 376-425. Maxinova, E., M. D. Marcel Uhrin. 2018. Salamandra salamandra (Fire Salamander). Predation. Herpetological Review. 49(2): 348 Mebs, D., M. Yotsu-Yamashita. 2017. Taricha granulosa (Rough-skinned Newt). Predation. Herpetological Review. 48(4): 828-829 Mendoza-Quijano, F., A. Loeza-Corichi, O. Flores-Villela. 1995. Pseudoeurycea belli belli (NCN). Predation. Herpetological Review 26 (3): 142. Messenger, K.R., N.A. Shepard, M.A. Yikira. 2011. Plethodon nettingi (Cheat Mountain Salamander). Predation. Herpetological Review 42 (4): 583-583.

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Montaña, C.G., M. Ceneviva-Bastos, C.M. Schalk. 2014. New vertebrate prey for the aquatic salamander Amphiuma means (Caudata: Amphiumidae). Herpetology Notes. 7:755-756.

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Mounce, M.J. 2004. Ambystoma texanum (Small-mouthed Salamander). Predation. Herpetological Review. 35 (1): 51.

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Murphy, T.D. 1961. Predation on eggs of the salamander, Ambystoma maculatum by caddisfly larvae. Copeia. 1961 (4):495-496.

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Nelson, K.T., A.M. Durso, R.V. Horan. 2010. Diadophis punctatus punctatus (Southern Ringnecked Snake). Diet. Herpetological Review 41 (1): 90-91. Norman, B.R. 1988. Plethodon vehiculum (Western Red-backed Salamander). Predation. Herpetological Review 19 (2): 34. Norman, B.R. 2002. Rhyacotriton variegatus (Southern Torrent Salamander). Predation. Herpetological Review. 33 (4): 298.

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Núñez, M.C. 2010. Bolitoglossa striatula (Striated Salamander). Predation. Herpetological Review 41 (2): 187. Orizaola, G., C. Rodríguez del Valle. 2000. Tritururus marmoratus (Marbled Newt). Predation. Herpetological Review. 31 (4): 233. Owen, J. D., T. J. Devitt, L. A. Colucci, N. F. Bendi. 2016. Eurycea sosorum (Barton Springs Salamander). Predation. Herpetological Review. 47(2): 275 Painter, C.W., N.J. Scott Jr., M.J. Altenbach. 1999. Thamnophis elegans vargans (Wandering Garter Snake). Diet. Herpetological Review. 30 (1): 48. Palis, J. G. 2016. Ambystoma talpoideum (Mole Salamander). Predation. Herpetological Review. 47(3): 477-478 Palis, J.G. 2011. Ambystoma opacum (Marbled Salamander). Shrew Predation. Herpetological Review 42 (3): 405.

Journal Pre-proof Palis, J.G. 2011. Eurycea quadridigitata (Dwarf Salamander). Centipede Predation. Herpetological Review 42 (3): 405. Palis, J.G. 2013. Siren intermedia (Lesser Siren) predation. Herpetological Review. 44 (2): 292-293. Paulson, T., S. Drukker, T.W. Pierson. 2015. Desmognathus abditus (Cumberland Dusky Salamander). Predation. Herpetological Review. 46 (3): 408-409. Pearl, C.A. 2003. Ambystoma gracile (Northwestern Salamander). Egg Predation. Herpetological Review. 34 (2): 352-353.

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Journal Pre-proof Smith, A.G. 1946. An unusual predator upon Ambystoma jeffersonianum. Herpetologica 3(3): 67. Smith, C., D Robinson. 2018. Plethodon glutinosus (Northern Slimy Salamander). Predation. Herpetological Review. 49(1): 130 Sorrell, G.G., S.D. Lindey, C.R. Newton. 2005. Plethodon glutinosus (Slimy Salamander). Predation. Herpetological Review. 36 (4): 429. Spohr, S.M. 2005. Ambystoma maculatum (Spotted Salamander). Predation. Herpetological Review. 36 (3): 294-295.

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Figure Captions Fig. 1: Relationship between salamander familial species richness and the number of predation events reported (R2 = 0.50, P = 0.02, y = 1.08x + 0.65). Fig. 2: Percentages of major predator groups reported as predators of salamanders (n = 156 salamander predations).

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Fig. 3: Percentages of predator groups separated by larval and adult salamander life stages (n =

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26 larval salamander predations, n = 130 adult salamander predations).

Fig. 4: Predator-prey size relationships of the salamander prey in relationship to A) invertebrate

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predators (R² = 0.27, P = 0.23, y = 0.59x + 0.63) and B) snake predators (R² = 0.11, P = 0.28, y =

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0.38x + 0.86). A 1:1 predator-prey size relationship is depicted by the solid line. A predator

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consumed a salamander that was larger than itself when a point falls above the solid line. A

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line.

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predator consumed a salamander that was smaller than itself when a point falls below the solid

Figure 1

Figure 2

Figure 3

Figure 4