Salamandra venenosa Daudin 1803
Creators: name: Pyron, R. Alexander
name: Beamer, David A.
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Caudata
Salamandridae
Salamandra
Salamandra venenosa

description: Salamandra venenosa Daudin, 1803
[= Lacerta maculata Shaw, 1802]
Holbrook (1838b:105, pl. 24) provided an account of what is now regarded as a junior subjective synonym of the Spotted Salamander, Ambystoma maculatum. Holbrook (1842e:67, pl. 22) reprinted this account nearly verbatim but replaced the relatively crude illustration by T.W. Hill with a new stone lithograph by J. Queen. The naturally posed drawing by Hill seemingly represents a live animal; Holbrook noted seeing it in the White Mountains of New Hampshire, which may be the model. Based on the contorted position of the legs, the Queen etching seems to be of a preserved specimen. The only known extant specimens of this taxon associated with Holbrook are ANSP 1383 and ANSP 4076 (both from “ New York ”), and MNHN-RA 0.4696 (from “ Caroline du sud ”). Additionally, ANSP 1322 from “ New Jersey ” is listed as A. tigrinum in the ANSP catalog but is also actually A. maculatum. We were unable to locate ANSP 4076 (cataloged as A. “ punctatum; ” see below), which bears a note in the electronic catalog “Donor: Dr. Holbrook. Taken from Penna. State Colln. Jan. 1895.”
In the account, Holbrook listed Salamandra subviolacea Barton, 1804 as a synonym, the name by which the species was also commonly known to other early workers such as Green (1825a). Holbrook included some seemingly critical words directed at Benjamin Smith Barton (1766–1815) regarding his role in the nomenclatural history of this species. That history includes some complexities that have never been fully explored in print to our knowledge. Thus, while Holbrook himself was likely unaware of some of these convolutions, contextualizing his comments requires us to explore them in detail here.
The Spotted Salamander (Ambysoma maculatum) was first described by Catesby (1747:10, pl. 10, fig. 2), figured in the beak of the heron “ Ardea Cristata Maxima Americana ” [= Ardea herodias Linnaeus, 1758]. Catesby’s pre-Linnaean work referred to the species as “ Stellio aquaticus minor Americanus. The Spotted Eft. ” The description of an animal 5 inches long, with a large head and double row of white, round spots from head to tail clearly refers to the Spotted Salamander. The illustration also clearly shows a Spotted Salamander. However, a reddish coloration of the plate in some editions or printings such as Catesby (1754:110, pl. 110, fig. 2, in the North Carolina Collection, Wilson Library, University of North Carolina at Chapel Hill), and possibly Catesby’s use of the word “eft,” caused some later workers such as Say (1819) to mistakenly identify the “ Stellio ” as Notophthalmus viridescens (see account for Salamandra symmetrica).
Subsequently, Linnaeus (1758:209) erected Lacerta punctata for a skink from Asia and Africa (Bauer 2003). He later reprinted this description verbatim (Linnaeus 1766:369); both entries clearly refer to an Old-World lizard. Curiously, he then described L. punctata Linnaeus, 1767:370 with explicit reference to Catesby (1747:10, pl. 10, fig. 2)’s “ Stellio.” Thus, L. punctata Linnaeus, 1767 is a junior primary homonym of L. punctata Linnaeus, 1758, and is therefore permanently invalid (Article 57.2; ICZN 1999). In a final posthumous edition (Catesby 1771), a later reviser (“Mr. Edwards, of the Royal College of Physicians, London”) added a “Linnaean Index of the Animals and Plants” which listed “The Spotted Eft. Stellio aquaticus minor Americanus. Lacerta punctata. L. ” (Edwards in Catesby 1771:2). As the ICZN was not in effect in the 18th or early 19th centuries, L. punctata Linnaeus, 1767 was thereafter used for the salamander by many subsequent authors (e.g., Lacépède 1788:237; Bonnaterre 1789:63; Gmelin 1789:1076; Sonnini de Manoncourt and Latreille 1801a:252).
Even then, the problem of preoccupation was obvious to some workers, leading Bechstein (1800:544) to erect Salamandra palustris as a replacement name for Lacerta punctata Linnaeus, 1767. Ironically, this name was also already preoccupied by S. palustris Schneider, 1799:60. The next replacement name erected for L. punctata Linnaeus, 1767 was L. maculata Shaw, 1802:304, again based explicitly on Catesby (1747) ’s “ Stellio.” This is consequently the senior-most available, valid name for the species, but was not recognized as such until much later (Stejneger 1902; see below).
On 15 April 1803, Barton read a paper to the American Philosophical Society describing a Spotted Salamander he had found near Philadelphia and considered to be a new species. Prior to the Society’s meeting, he also apparently informed Rafinesque of his discovery, who relayed it to Daudin in a letter on 2 April 1803, calling it the “Lézard aquatique.” Daudin then published a description in his own volume as Salamandra venenosa Daudin, 1803:229, based on Barton’s notes from Rafinesque’s letter. We must assume that Barton had not yet named his species, as Daudin writes (translated): “... and it was from him [Barton] that I called her a poisonous salamander, because of a liquor which he [Barton] believed to be poisonous, and which oozing from all the warts spilled on his [the specimen’s] back.”
Daudin apparently gave no consideration to priority, as he listed the senior names Lacerta punctata from Gmelin (1789) [= L. punctata Linnaeus, 1767], Catesby (1747) ’s “ Stellio,” and other usages of L. punctata Linnaeus, 1767 from Daubenton (1782), Lacépède (1788), and Sonnini de Manoncourt and Latreille (1801a) in synonymy. He stated (translated): “The poisonous salamander is doubtless very similar to the preceding one. It is here that the punctuated salamander described by Gmelin, Daubenton and Lacépède, from a figure published by Catesby, in his work on the natural history of Carolina, must be brought together.” In a later account for “ La salamandra ponctuée,” Daudin (1803:257) then allocated Salamandra punctata Latreille, 1800 (itself originally intended as a replacement name for Triton parisinus Laurenti, 1768) to “var. B” of L. “ salamandra ” aquatica from Gmelin (1789:1066) [= L. aquatica Linnaeus, 1758]. All of these are junior subjective synonyms of L. vulgaris Linnaeus, 1758, now placed in Lissotriton Bell, 1839 (= Li. vulgaris). This is a European newt; Daudin’s use of the specific epithet “ punctata ” for this species was distinct from his use of L. punctata Linnaeus, 1767 for the Spotted Salamander (see below).
Daudin finally noted that Barton had submitted his paper for publication for the following year in the sixth volume of the Academy’s Transactions. Thereafter, Lacerta subviolacea Barton, 1804:108 became the name by which the Spotted Salamander was known to most American naturalists (e.g., Green 1827, 1830; Harlan 1827, 1835). Barton claimed that his species is distinct from Catesby (1747) ’s “ Stellio ” and L. punctata Linnaeus, 1767, but this was based only on minor disagreements with Catesby’s somewhat crude illustration. He did not, however, seem to think that either represented a newt. He also reported tasting the milky white fluid that his specimen exuded (and inducing his students and colleagues to do so), finding it astringent, caustic, and corrosive, the taste lasting nearly a day, and inducing excessive salivation. This was clearly communicated by Rafinesque to Daudin, resulting in the latter’s name for the species. In contrast, Barton’s name was based on the animal’s purportedly dark purple coloration, but this was apparently not mentioned in Rafinesque’s letter.
Confusion began to arise with Say (1819), who stated that Daudin (1803), for reasons unknown to Say, had intentionally substituted Salamandra venenosa Daudin, 1803 for Lacerta subviolacea Barton, 1804. Obviously, the dates contradict this as 1803 is before 1804; the impetus for Say’s statement is therefore unclear. Contrariwise, it seems most likely that Daudin was unaware of Barton’s intended name, and merely erected his own in its absence. Say then noted that Daudin considered Barton’s species the same as Catesby (1747) ’s “ Stellio ” (i.e., the Spotted Salamander) and asserted that Daudin had therefore assigned the name “ Salamandra punctata ” from Gmelin, 1789 [= L. punctata Linnaeus, 1767] to “var. B” of L. “ salamandra ” aquatica from Gmelin, 1789 [= L. aquatica Linnaeus, 1758], i.e., a European newt referred to by Gmelin (1789:1066). However, Say confused the name L. punctata from Gmelin, 1789:1076 [= L. punctata Linnaeus, 1767] (the Spotted Salamander) with the junior secondary homonym S. punctata Latreille, 1800 [= L. vulgaris Linnaeus, 1758] (the Smooth Newt), which Daudin had synonymized with L. aquatica Linnaeus, 1758. In reality, Daudin considered S. venenosa Daudin, 1803 and L. punctata Linnaeus, 1767 to be the same species—the Spotted Salamander—while Say erroneously concluded that Daudin had allocated the latter to the Smooth Newt.
Compounding the confusion, Say (1819) also rejected Daudin (1803) ’s acceptance of the prevailing conclusion that Lacerta subviolacea Barton, 1804 was Catesby (1747) ’s “ Stellio ” [= L. punctata Linnaeus, 1767],and agreed with Barton (1804) that they were distinct species. Say then described his belief (as noted in the account for Salamandra symmetrica) that Catesby’s “ Stellio ” is the red eft stage of the Eastern Newt (Notophthalmus viridescens), which was then thought to be a different species. He stated that L. maculata Shaw, 1802 was also this species (the red eft), and that its name should therefore be “ Salamandra punctata ” from Gmelin (1789) [= L. punctata Linnaeus, 1767]. Finally, he resurrected Triton parisinus Laurenti, 1768 for “var. B” of L. “ salamandra ” aquatica from Gmelin, 1789 [= Lacerta aquatica Linnaeus, 1758].
Holbrook (1838a) noted this mixup by Say (1819), stating that the red eft had apparently been confused with Catesby (1747) ’s “ Stellio ” (i.e., by Say) until the description of Salamandra symmetrica Harlan, 1825c (see account above), but that the two were clearly distinct. Subsequently, Holbrook (1838b) noted that Daudin (1803), Harlan (1827, 1835), and others considered Catesby’s “ Stellio ” to represent S. venenosa Daudin, 1803, while Say (1819) and others considered Catesby’s “ Stellio ” to represent S. dorsalis Harlan, 1829. Holbrook then incorrectly stated that it is neither, but a third, distinct, as-yet unnamed species “peculiar to the south” that he had only observed “about the Sea Islands of South Carolina.” No further reference to this supposed new species appears in Holbrook’s later writings.
Holbrook (1838b) then took seemingly unwarranted aim at Barton, beginning by describing the history (enumerated in part above) of Barton’s address to the American Philosophical Society in Philadelphia and Rafinesque’s subsequent letter to Daudin. Holbrook apparently concluded that it was Barton who gave the species the name “ venenosa,” stating that Daudin had published Barton’s description “under the name it now bears, which was probably given by Barton, and communicated through Rafinesque to Daudin, for the latter gives the specific name, as imposed by Barton.” However, as noted above, it seems from Daudin’s account as though Barton’s description communicated to him through Rafinesque lacked a name, and that Daudin himself christened it “ venenosa ” based on Barton’s anecdote of its noxious secretions.
Holbrook finally stated that Barton published his description six years after Daudin in the sixth volume of the Transactions of the American Philosophical Society, though it was in 1804 and not 1809 that part I of this volume appeared with Barton’s paper. Finally, Holbrook stated that Barton “had no right to change a name, even though imposed by himself, after it had been once published to the scientific world.” Yet it seems likely that Barton was unaware of Daudin’s intention to establish a different name, as Volume VI, Part I of the Transactions of the American Philosophical Society was published on 23 March 1804, less than a year after Barton’s first address to the American Philosophical Society. Accordingly, Daudin (1803) was published sometime after 11 July 1803, likely August or September (Harper 1940), wherein Daudin was already aware that Barton’s paper was going to press in the forthcoming sixth volume of the Transactions of the American Philosophical Society. Thus, the blame might be placed instead on Daudin for inadvertently “scooping” Barton’s name, or even that Daudin knew of Barton’s name and gave his own anyway, given Daudin’s evident lack of consideration for priority.
Few if any subsequent authors followed Holbrook (1838b, 1842e)’s lead in recognizing Salamandra venenosa Daudin, 1803 as the correct name for this species. Baird (1850) correctly reinstated Lacerta punctata Linnaeus, 1767 as the senior-most valid, available subjective synonym; no widely accepted Code forbidding homonymy was in force at the time. Consequently, Ambystoma punctatum became widely applied to this taxon thereafter (e.g., Seiss 1881). Stejneger (1902) was the first to note the invalidity of the junior primary homonym L. punctata Linnaeus, 1767 and restored L. maculata Shaw, 1802 as the senior-most available, valid synonym for the species under modern nomenclatural rules. However, use of A. punctatum nonetheless persists over a century later, occurring in at least 725 studies since 1902 (Google Scholar 29 January 2022) as recently as Huysseune et al. (2022). Many are simply repeating citations from older literature and are from fields such as organ regeneration that are far-removed from the complexities of herpetological nomenclature, but this nevertheless underscores the importance thereof for accurate communication in science.
Finally, several authors (e.g., Stejneger 1902; Anderson 1967) discussing the nomenclatural history of this species erroneously cited Lacerta punctata Linnaeus, 1766 [= L. punctata Linnaeus, 1758], a lizard (see Bauer 2003), when they clearly intended to reference L. punctata Linnaeus, 1767 based on Catesby (1747) ’s “ Stellio ” for the Spotted Salamander. This caused considerable confusion for recent authors attempting to untangle the complex series of names applied to this species as enumerated above (Darrel R. Frost, pers. comm.). We propose this to be a complete accounting of the possible synonyms and homonyms with correct citations, but it is nevertheless possible that additional details will emerge in the future.
description: Published as part of Pyron, R. Alexander & Beamer, David A., 2022, A nomenclatural and taxonomic review of the salamanders (Urodela) from Holbrook's North American Herpetology, pp. 151-196 in Zootaxa 5134 (2) on pages 159-161, DOI: 10.11646/zootaxa.5134.2.1, http://zenodo.org/record/6534805