Bioacoustics of Incilius nebulifer

Biology of The Gulf Coast Toad. The catalogue account for Bufo valliceps ( Porter, 1970 ) , as well as Porter’s (1962; 1964a–c) overall concept of the species, is a composite of I. nebulifer , I. valliceps , and additional species of Incilius ( Mendelson 1998 ) ; however, those records and literature cited that pertain to specimens from the currently known range of I. nebulifer are valid for this latter species. Morphology. Morphological descriptions and analyses of morphological data were presented by Bauldauf (1958), McAlister (1961) , W. Martin (1967; 1971), R. Martin (1972a , b ), Martin & Gans (1972) , Gans (1973) , Emerson (1982) , Hutchinson & Savitzky (2004) R. Martin (1972b) . The analyses of sternal morphology and inguinal fat body organs by da Silva & Mendelson (1999) included specimens referable both to I. nebulifer and I. valliceps , however no differences in these structures are evident between the two species ( Mendelson et al. 2011 ). The osteological review provided by Martin (1973) is problematic because it was based on specimens referable to I. nebulifer , but also included a skeletal specimen identified as Bufo valliceps (the specimen in Martin 1973 :fig. 2D is TNHC 41994; T. LaDuc, in lett. ); this specimen bears the locality data “ Mexico : Sinaloa” which is greatly extralimital for the either I. nebulifer or I. valliceps . Blair (1962 ; 1970 ) discussed the importance of various types of data to inform understanding of this species, with respect to other bufonids. Cytological comparisons with other bufonid species were presented by Sanders & Cross (1964) . A morphologically anomalous specimen was described by Freed (1992) and larval deformities by Drake et al. (2007) . Effects of long-term preservation on specimens were reported by Deichmann et al. (2009) . Reproduction, development, and hybridization. Various aspects of the reproductive biology, including the advertisement calls, of I. nebulifer have been documented by A. P. Blair (1947) , Wright & Wright (1949) , Duran (1954) , Martin (1958) , W. F. Blair (1956a ; 1960a ), Thornton (1960) , Awbrey (1963) , Porter (1964a) , Hubbs & Martin (1967) , Grubb (1970 , 1973a ), W. Martin (1972), Wiest (1982) , Sullivan & Wagner (1988) , Dundee et al. (1989) , Ryan & Sullivan (1989) , Branson (1995) , Wagner & Sullivan (1992 ; 1995 ), Foley (1994) , Greuter (2004) , Lannoo et al. (2005), Salinas (2009) , Jones & Ratnam (2009) , Pierce & Hall (2012) , and Oyervides & Zaidan (2013) . Development, metamorphosis, and ontogeny and factors affecting them have been documented by Taylor (1942) , Wright & Wright (1949) , Volpe (1957), Limbaugh (1956) , Limbaugh & Volpe (1957) , Conant (1958 , 1975 ); Blair (1953 ; 1956b ; 1960a ; 1963 ), Turner (1960 , 1962 ), Hubbs et al. (1963) , Ballinger & McKinney (1966) , Yew (1966 ; 1969 ), Licht (1967) , Grubb (1973b) , Dundee et al. (1989) , Pierce & Montgomery (1989) , Nelson (1993) , Rosenberg (1990) , Rosenberg & Pierce (1995) , and Conant & Collins (1998) . Various aspects of hybridization with other anurans have been discussed by Blair (1941) , Orton (1951) , Liner (1954) , Thornton (1954 , 1955 ), A. P. Blair (1941) , W. G. Blair (1956b ; 1958 ; 1959 ; 1960a ; 1961b ; 1963 ; 1964 ; 1966 ; 1972 ), Moore (1955) , Volpe (1956 ; 1959 ; 1960 ), Gosner & Black (1958) , Fox et al. (1961) , Kennedy (1962) , Brown (1971a , b ), Guttman (1972) , Brown & Brownell (1971) , Brownell (1971), Sanders (1978 ; 1985 ; 1986 ), Hillis et al. (1984) , Dundee et al. (1989) , Greuter (2004) , Brown & Mesrobian (2005) , Mulcahy et al. (2006) , and Mendelson et al. (2011) . This older literature regarding hybridization must be interpreted in the context of the taxonomic separation of I. nebulifer and I. valliceps by Mulcahy & Mendelson (2000) , and also in the context of the re-analyses of data from the laboratory of W. F. Blair presented by (2008) and subsequently re-analyzed by Brandvain et al . (2014) . Additional, recent, studies of hybridization were presented by Vogel (2007) and Vogel & Johnson (2008) . Habitat use. Habitat use by the adults has been discussed by Pope (1919) , Wright & Wright (1949) , Thornton (1960) , Awbrey (1963) , Brattstrom (1963) , Wilks (1963) , Grubb (1968 , 1970 ), Neill & Grubb (1971) , Grubb (1976) , Moore (1976) , Whiting et al. (1987) , Dundee et al. (1989) , McAllister et al. (1989) , Foley (1994) , Reid & Whiting (1994) , Irwin (1997) , Mendelson (1998) , Mulcahy & Mendelson (2000) , Means (2005) , Mulcahy et al. (2006) , Lemos-Espinal & Smith (2007) , Vogel & Johnson (2008) , Salinas (2009) , Oliver-López et al (2009) , Gehlbach (2010) , Walls et al. (2011) , Hernández-Salinas et al. (2012) , and Milko (2012) . Associations with caves has been reported by Mohr (1948) , McAlister (1954) , Baker (1957) , Redell & Knox (1962) , Redell & Finch (1963) , and Reddell (1970) . Specific associations with saltwater were documented by Viosca (1926) , Allen (1932) , Burger et al. (1949) , reviewed by Neill (1958) , Mueller (1985) , Alexander et al. (2012) and Hua & Pierce (2013) . Responses to fire were reported by Brown et al. (2011 ; 2014 ). Potential effects of exposure to introduced plant species were reported by Cotton (2009) and Cotten et al. (2013) . Community ecology. Feeding ecology has been documented by Strecker (1927a) , Campbell & Davis (1968) , Clark (1969) , and McGehee et al. (2001). Predators and possible antipredator mechanisms have been documented by Strecker (1927b) , Wright & Wright (1949) , Licht (1968) , Neill (1968a , b ), Grubb (1972) , Clark (1974) , Brown (1974) , Platt & Fontenot (1993) , Tucker (1994) , Adams (2005) , and Mendelson (2005) . Palatability of the tadpoles was tested by Adams et al. (2011) . Parasites have been reported by Walton (1929) , Harwood (1930) , Hoffpauir & Morrison (1966) , McAllister et al. (1989) , McAllister & Donn (1990) , and Martin & Dresser (1991) . Pathogens were reported by Gaertner et al (2010) , and Saenz et al. (2010) . Interspecific associations with other anurans have been discussed by Glass (1946) , Wright & Wright (1949) , Axtell (1958) , Blair (1961a) , James (1966) , Moore (1976) , Gambs & Littlejohn (1979) , Foley (2005), Mitchell & Lannoo (2005a , b ), Schwalbe & Goldberg (2005) , Franklin & Killpack (2005) , Sredl (2005), Sredl & Field (2005) , Sullivan (2005) , Wallace (2005) , Vogel & Pechmann (2010) , and Preston et al. (2014) . Associations with other species caught in traps with this species were reported by Blair (1960b) . Population trends were discussed by Boundy (2005) . Camper & Dixon (1988) evaluated a microchip marking technology in the species. Road and industrial mortality was reported by Flickinger (1981) and Ray et al. (2006) ; road mortality and scavenging by Thamnophis proximus was reported by Watson (2007) . Physiology and behavior. Various aspects of the physiological parameters and responses in this species have been presented by Morgan & Stokes (1936) , Svedberg & Hedenius (1934) , Sanders (1962) , Wittliff (1964) , Campbell & Davis (1971) , Guttman (1974) , Withers (1978) , Prasad et al. (1984) , Wygoda (1989) , Williams (1992) , Williams & Wygoda (1993) , Verma & Pierce (1994) , Moore (1997) , and Rowson et al. (2001) . Veterinary protocols were presented by Brannelly et al. (2012) and Brannelly (2014) and notes on captive care were provided by Walsh et al. (1992) . Phototactic responses were presented by Jaeger & Hailman (1973) and Hailman & Jaeger (1974) ; however, the source for the specimens used by Jaeger & Hailman (1973, presented as B. valliceps ) is not reported, so referral of data as relevant to I. nebulifer is speculative. Interspecific effects of injections of pituitary extracts were presented by Quinn & Mengden (1984) . Hemoglobins and transferrins were described by Fox et al. (1961) . Resistance to cardiotoxins was reported by Moore et al. (2009) The parotoid secretions were analyzed by Porter (1964c) , but results are based on specimens referable both to I. nebulifer and I. valliceps . Serological comparisons were made by Durio (1960) and biochemical components in the skin were analyzed by Cei et al. (1968) . Use of this species for human pregnancy testing was evaluated by Aguirre-Pequeno (1950) . Effects of potentially toxic chemicals were evaluated by Chen & Chen (1933) , and Shaw & Grushkin (1957) . Effects of radiation were reported by Blair (1960c) and Clayton (1960) . Clark (1971) reported on use of branding as a means of marking individuals. Greding (1971) reported on learning in this species. Evolution. The phylogenetic literature, with respect to I. nebulifer , can be difficult to interpret if the authors did not state the provenance of the samples used in their analyses. For example, Baldauf (1959) implies that the individuals used in his morphological analyses were collected in the USA (and possibly may be the same individuals reported by Baldauf, 1958 ), but one cannot be certain. Maxson (1984) and Maxson et al. (1981) included “ Bufo valliceps ” in their studies from Texas, so they can be assigned with certainty to I. nebulifer . All phylogenetic treatments of the taxon “ valliceps ” published prior to 2000 were based, variously, on specimens or samples representing either I. valliceps (sensu stricto) or I. nebulifer . The modern reader of those papers is encouraged to check the geographic source of the material included in any particular analysis and assign the appropriate taxonomic name. Because of the well-supported sister-taxon relationship between I. nebulifer and I. valliceps ( Mulcahy & Mendelson 2000 ; Mulcahy et al. 2006 ; Mendelson et al. 2011 ), the taxonomic differences typically have virtually no bearing on the overall evolutionary discussions presented by those authors. The exception, however, is the result presented by Van Bocxclaer et al. (2010), wherein they reported that the species pair valliceps + nebulifer ) was rendered paraphyletic by a sample identified as “ I. macrocristata ” (= I. macrocristatus ). Mendelson et al. (2011) reviewed all of the relevant material, presented a clarification of the issue and determined that Van Boxclaer et al. (2010) had included a misidentified specimen in their analyses (i.e., their sample of “ I. macrocristata ” actually represented I. valliceps ). In their broad analysis of amphibian relationships, Pyron & Wiens (2011) chose to compile various published genetic sequences (i.e., GenBank) to create chimera to represent the taxa “ Bufo valliceps ” and “ Bufo nebulifer ” for their analyses; their compilation of sequences from various sources into singular OTUs included the misidentified material from Van Bocxlaer et al. (2010) , with the result that Pyron & Wiens presented “ valliceps ” and “ nebulifer ” not to be sister taxa. Because their analyses include clearly misidentified material, this section of their overall phylogenetic tree is inadmissible. Thus, the sister relationship between I. valliceps + I. nebulifer remains is supported by all modern phylogenetic analyses that are based on accurately identified material. Phylogenetic relationships, biogeographic history, and evolutionary natural history of I. nebulifer were presented and discussed by Mulcahy & Mendelson (2000) , Mulcahy et al. (2006) , and Mendelson et al. (2011) . The relative phylogenetic diversity of I. nebulifer , with respect to other North American bufonids, in terms of conservation prioritization was presented and discussed by Goebel (2005) . Conservation and education. Summary of possible conservation threats to this species were presented by Hammerson & Canseco-Márquez (2004) , Mendelson (2005) , Frías-Alvarez et al. (2010) . Use of habitat by this species following mining activities was reported by Walton (2012) . Use of these common toads in conservationawareness education programs was described by Rommel (2012) .