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Lampropeltis triangulum (LACÉPÈDE, 1789)

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Higher TaxaColubridae, Colubrinae, Lampropeltini, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesLampropeltis triangulum gentilis (BAIRD & GIRARD 1853)
Lampropeltis triangulum syspila (COPE 1889)
Lampropeltis triangulum triangulum (LACÉPÈDE 1789) 
Common NamesE: Eastern Milksnake
E: Louisiana Milksnake [amaura]
E: Red Milksnake [syspila]
G: Rote Königsnatter, Milchschlange, (Östliche) Dreiecksnatter
E: Mexican Milksnake [annulata]
E: Central Plains Milksnake [gentilis]
E: Pale Milksnake [multistriata]
E: Utah Milksnake [taylori]
E: New Mexico Milksnake [celaenops]
S: Barila 
SynonymColuber triangulum LACÉPÈDE 1789: 86
Coluber doliatus — SHAW & NODDER 1795: plate 254
Coluber eximius HARLAN 1827
Ophibolus clericus BAIRD & GIRARD 1853
Ablabes triangulum — DUMÉRIL, BIBRON & DUMÉRIL 1854: 315 (?)
Lampropeltis triangula — COPE 1860: 256
Lampropeltis amaura COPE 1861: 258 (footnote)
Coronella eximia — JAN 1866 ?
Ophibolus doliatus amaura — COPE 1875: 36 (part.)
Ophibolus triangulus — GARMAN 1884: 66
Ophibolus doliatus parallelus COPE 1889: 385
Ophibolus doliatus collaris COPE 1889: 383
Coronella annulata — GÜNTHER 1893 (fide VILLA et al. 1988)
Ophibolus doliatus annulatus — COPE 1893: 1067
Ophibolus doliatus parallelus — COPE 1893: 1067
Ophibolus doliatus collaris — COPE 1893: 1068
Ophibolus doliatus clericus — COPE 1893: 1068
Coronella triangulum collaris — BOETTGER 1898: 72
Osceola doliata parallela — COPE 1900: 893
Osceola doliata triangula — MCATEE 1907
Lampropeltis triangulum amaura — STEJNEGER & BARBOUR 1917: 90 (part.)
Lampropeltis polyzonus — DITMARS 1934 (fide VILLA et al. 1988)
Lampropeltis triangulum amaura — BURT 1935 (part.)
Lampropeltis polyzona blanchardi — SCHMIDT & ANDREWS 1936 (VILLA et al.)
Lampropeltis doliata amaura — KLAUBER 1948: 11 (part.)
Lampropeltis triangulum — STEBBINS 1985: 194
Lampropeltis triangulum triangulum — CONANT & COLLINS 1991: 207
Lampropeltis triangulum amaura — CONANT & COLLINS 1991: 210 (part.)
Lampropeltis triangulum — LINER 1994
Lampropeltis triangulum triangulum — CROTHER 2000: 64
Lampropeltis triangulum amaura — CROTHER 2000: 64 (part.)
Lampropeltis triangulum amaura — TENNANT & BARTLETT 2000: 434 (part.)
Lampropeltis triangulum triangulum — TENNANT & BARTLETT 2000: 425
Lampropeltis triangulum triangulum — CROTHER et al. 2012
Lampropeltis triangulum — CROTHER et al. 2012
Lampropeltis triangulum amaura — CROTHER et al. 2012 (part.)
Lampropeltis triangulum — RUANE et al. 2014
Lampropeltis triangulum — WALLACH et al. 2014: 359
Lampropeltis triangulum triangulum — GUYER et al. 2018
Lampropeltis triangulum — CHAMBERS et al. 2022

Lampropeltis triangulum gentilis (BAIRD & GIRARD 1853)
Ophibolus gentilis BAIRD & GIRARD 1853: 90
Lampropeltis annulata COPE 1860: 257 (refers to ms. of KENNICOTT) (part.)
Lampropeltis annulata KENNICOTT 1861: 329 (part.)
Lampropeltis multistriata KENNICOTT 1861: 328
Lampropeltis multistrata — COPE 1861: 566
Coronella doliata var. gentilis — JAN 1866
Ophibolus multistratus — COPE 1875: 36
Ophibolus triangulus var. gentilis — GARMAN 1884: 66
Ophibolus multistratus — GARMAN 1887: 128
Lampropeltis multistrata – STEJNEGER 1891: 502
Ophibolus doliatus gentilis — COPE 1893: 1068
Lampropeltis pyrrhomelana celaenops STEJNEGER 1902: 153
Lampropeltis pyromelana celaenops STEJNEGER & BARBOUR 1917: 89
Coronella multistrata – WERNER 1929: 129
Lampropeltis doliata multistrata — TANNER & LOOMIS 1957: 12
Lampropeltis doliata celaenops — TANNER & LOOMIS 1957: 15
Lampropeltis doliata taylori TANNER & LOOMIS 1957: 31
Lampropeltis triangulum celaenops — STEBBINS 1966: 16
Lampropeltis triangulum multistrata — STEBBINS 1966: 161
Lampropeltis triangulum gentilis — MEHRTENS 1987: 127
Lampropeltis triangulum multistriata — CONANT & COLLINS 1991: 209
Lampropeltis triangulum celaenops — CONANT & COLLINS 1991: 209
Lampropeltis triangulum gentilis — CONANT & COLLINS 1991: 209
Lampropeltis triangulum annulata — CONANT & COLLINS 1991: 209 (part.)
Lampropeltis taylori — COLLINS 1991: 43
Lampropeltis triangulum taylori — WILLIAMS 1994
Lampropeltis triangulum multistrata — WILLIAMS 1994
Lampropeltis triangulum celaenops — WILLIAMS 1994
Lampropeltis triangulum multistriata — CROTHER 2000: 64
Lampropeltis triangulum gentilis — CROTHER 2000: 64
Lampropeltis triangulum celaenops — CROTHER 2000: 64
Lampropeltis triangulum taylori — CROTHER 2000: 64
Lampropeltis triangulum annulata — CROTHER 2000: 64
Lampropeltis triangulum multistrata [sic] — TENNANT & BARTLETT 2000: 442
Lampropeltis triangulum annulata — TENNANT & BARTLETT 2000: 436 (part.)
Lampropeltis triangulum celaenops — TENNANT & BARTLETT 2000: 438
Lampropeltis triangulum gentilis — TENNANT & BARTLETT 2000: 440
Lampropeltis triangulum gentilis — BARTS 2006
Lampropeltis triangulum multistriata — CROTHER et al. 2012
Lampropeltis triangulum celaenops — CROTHER et al. 2012
Lampropeltis triangulum gentilis — CROTHER et al. 2012
Lampropeltis triangulum taylori — CROTHER et al. 2012
Lampropeltis triangulum annulata — CROTHER et al. 2012 (part.)
Lampropeltis gentilis — RUANE et al. 2014
Lampropeltis triangulum gentilis — CHAMBERS et al. 2022

Lampropeltis triangulum syspila (COPE 1889)
Ophibolus doliatus syspilus COPE 1889: 381 (part.)
Lampropeltis doliatus syspilus — HAY 1892: 516 (part.)
Ophibolus doliatus syspilus — COPE 1893: 1067
Osceola doliata syspila — COPE 1900: 891 (part.)
Lampropeltis triangulum syspila — RUTHVEN 1919: 3 (part.)
Lampropeltis triangulum syspila — BURT 1935 (part.)
Lampropeltis triangulum syspila — MEHRTENS 1987: 128
Lampropeltis triangulum syspila — CONANT & COLLINS 1991: 208 (part.)
Lampropeltis triangulum syspila — CROTHER 2000: 64 (part.)
Lampropeltis triangulum syspila — TENNANT & BARTLETT 2000: 432 (part.)
Lampropeltis triangulum syspila — CROTHER et al. 2012 (part.)
Lampropeltis triangulum syspila — GUYER et al. 2018
Lampropeltis triangulum syspila — CHAMBERS et al. 2022 
DistributionCanada (Ontario, S Quebec),
USA (N Texas, W Oklahoma, C/W Kansas, E Colorado, and SC / SW Nebraska, Illinois, Minnesota, Michigan, Iowa, Missouri, Tennessee, Arkansas, N Louisiana, N Mississippi, N Alabama, N Georgia, N South Carolina, N North Carolina, Virginia, West Virginia, Kentucky, Indiana, Illinois, Wisconsin, Ohio, Maryland, Pennsylvania, New Jersey, New York, Rhode Island, Connecticut, Massachusetts, New Hampshire, Vermont, S Maine)
Mexico (Coahuila)

Type locality: “America”

amaura (invalid): SW Arkansas, SE Oklahoma, E Texas and Louisiana. Type locality, original entry of type reads "?Mississippi," restricted by Schmidt (1953) to the vicinity of New Orleans, Louisiana. Williams (1978) suggested that the type locality be considered as unknown, inasmuch as the characteristics of the holotype do not agree with data taken from specimens from New Orleans and vicinity.

celaenops (invalid): USA (SW Texas, E New Mexico, SE Arizona). Type locality: Mesilla Valley, New Mexico.

gentilis: Kansas, W Oklahoma, E Colorado, Texas panhandle, C/SW Nebraska, C/W Kansas; Type locality: "north fork of the Red River, near Sweetwater Creek, Wheeler County, Texas."

multistrata (invalid): USA (C/NW Nebraska, S Montana, SW North Dakota, N/C Wyoming); Type locality: "Fort Lookout, Nebraska," now in Lyman County, South Dakota.

syspila: USA (Nebraska, Oklahoma, Kansas)

taylori (invalid): C/NE Utah, W Colorado, N Arizona; Type locality: "from approximately 2 miles north of Alpine, Utah County, Utah."  
Reproductionoviparous. KREUTZ (2005) reports hybrids between E. guttata and Lampropeltis getulus californiae, between E. guttata and L. pyromelana, and between E. guttata and L. triangulum sinaloae/nelsoni! 
TypesHolotype: lost or never designated (fide WILLIAMS 1994)
Holotype: USNM 5282, an adult male [amaura]
Syntypes: USNM 2192 etc [Ophibolus doliatus collaris]
Holotype: USNM 13380 [Ophibolus doliatus syspilus]
Holotype: USNM 10544 [Ophibolus doliatus parallelusf]
Holotype: USNM 2380 [Ophibolus clericus]
Lectotype: USNM 131737 (old catalog number 1852 or 1853), adult male (designated by BLANCHARD 1921) [gentilis]
Holotype: USNM 22375 [celaenops]
Holotype: USNM 1842, young male [multistriata]
Holotype: BYU 10533, adult female [taylori] 
DiagnosisDiagnosis: Lampropeltis triangulum differs from L. rutheni, L. mexicana, and L. alterna in having the head less distinct from the body. In addition, this species differs from L. ruthveni (within the range of the latter) in having less uniform red rings, often obscured by black pigment middorsally. Another difference between L. triangulum on the one hand, and ruthveni and L. mexicana on the other, is that in the latter two the black rings are edged with a paler color (usually a pale lime green). Lampropeltis ruthveni has a lower ventral count (182-196) than sympatric L. triangulum (196-221). All members of the mexicana group tend to have considerable gray in the body colors, such is not so in triangulum, except some triangulum in the northeastern United States. Lampropeltis triangulum differs from L. zonata in that the white rings widen laterally in the former; from L. pyromelana in having a black, black and white, or red snout, rather than white. Lampropeltis pryomelana has 46-85 white rings on body, more than L. triangulum (10-32). Both L. pyromelana and L. zonata have a higher ventral scute number (213-238, and 194-227, respectively) than any of the adjacent western United States members of L. triangulum (174-208). Lampropeltis triangulum differs from both L. calligaster and L. getula in having an alternating pattern of white (yellow), red and black rings or blotches, the latter reaching the ventrals or nearly so, and in having the posterior two maxillary teeth longer and stouter than the preceding teeth. (Williams 1994)

Definition. A species of Lampropeltis with a body and tail color pattern consisting of rings or blotches involving the following sequence starting from the nape: yellow (or white), black, red Cor orange, brown, or gray), black. If red (or equivalent) blotches are present, they are completely margined by black pigment, with pale pigment surrounding each blotch. The venter in blotched forms varies from an almost immaculate white to a checkerboard pattern of black and white rectangular markings. The red and white scales on the body and tail vary from having no black pigment on them to being nearly obscured by it. Dorsal scale rows normally range from 19-23 at midbody, reducing posteriorly (usually by fusion of rows 4 + 5) to 17 or 19. Supralabials usually are 7, infralabials 9. Preoculars usually are 7, postoculars 2, loreals (frequently absent in individuals from the southeastern United States). Temporals normally are 1 + 2 or 2 + 3. Ventrals in males range from 154-236, in females from 161-244; subcaudals in males from 32-63, in females from 31-60. Maxillary teeth range from 11-15, the last two are larger than those preceding them. Adult size varies geographically from small individuals (ca. 500 mm) inhabiting the southeastern United States to large individuals (1500-1900 mm) in southern México and Central America. The head shape in relation to body width varies from slightly distinct to distinct. Tail length averages about 13.5% of total length in males, 14.4% in females. (Williams 1994)

Descriptions. Williams (1988) provided detailed descriptions of all subspecies. Quinn (1983) named and gave detailed descriptions of L. I. campbelli and L. I. dixoni. Powell and Parmerlee (1981) described a color variant L. t. blanchardi from Quintana Roo, México, which was banded only over the posterior portion of the body. They also suggested that this pattern is in agreement with a possible trend (unpublished information) toward a uniform red coloration in populations in the northeastern portion of the Peninsula de Yucatán. Frank (1994) included a table listing characteristic patterns of all subspecies. (Williams 1994) 
CommentSynonymy after PETERS & OREJAS-MIRANDA 170, WILLIAMS 14, and RUANE et al. 2014, and Chalmers et al. 2022 (although they did not comment on synonyms beyond moving gentilis back to triangulum as subspecies).

Distribution: L. gentilis is not in Nuevo León but annulata is, fide Nevárez-de los Reyes et al. 2016. See map in Chalmers et al. 2022: 359 (Fig. 1).

Distribution: revised after genetic analysis of Ruane et al. 2014. Previously, this species was also reported from Mexico (e.g. Jalisco, San Luis Potosí, Nuevo Léon, Coahuila, Nayarit, Sonora etc., e.g. by Lemos-Espinal et al. 2016), south to Colombia (Valle del Cauca, Valencia et al. 2014).

Subspecies: Lampropeltis triangulum elapsoides (HOLBROOK 1838) has been elevated to species status. Ruane et al. (2014) elevated a number of subspecies of L. triangulum to full species and synonymized the remaining subspecies as follows: Subspecies synonymized under L. micropholis include L. t. gaigeae and L. t. andesiana. Subspecies synonymized under L. abnorma include L. t. blanchardi, L. t. hondurensis, L. t. oligozona and L. t. stuarti. Subspecies synonymized under L. polyzona include L. t. arcifera, L. t. conanti, L. t. campbelli, L. t. nelsoni, L. t. smithi, and L. t. sinaloae. Subspecies synonymized under L. annulata include L. t. dixoni. Subspecies synonymized under L. gentilis include L. t. celaenops, L. t. multistriata, L. t. taylori, L. t. amaura (in part), L. t. syspila (in part) and L. t. annulata (in part). Subspecies synonymized under L. triangulum include L. t. syspila (in part) and L. t. amaura (in part). L. gentilis and triangulum have a broad contact zone with little reproductive isolation which does not warrant recognition as full species fide Chalmers et al. 2022.

Nomenclature: LACÉPÈDE’s name conserved through ICZN opinion 1463. Coluber doliatus was suppressed via Opinion 804.

Mimicry: Lampropeltis triangulum mimicks Micrurus bernadi, M. limbatus and M. diastema (TORRE-LORANCA et al. 2006).

NCBI taxonID: 165043 [gentilis] 
EtymologyThe name triangulum is derived from the Latin meaning "triangle," in reference to the shape of the anterior edge of the first blotch.

The name gentilis is derived from the Latin, meaning "of the same race or clan", presumably in reference to its relationship to other members of the species.

The name celaenops may be derived from the Greek word celaeno, which means "black or dark," and the ending -ops, meaning eye.

The name multistrata is derived from the Latin multus, meaning "many," and the Latin stratum, meaning "layer," apparently in reference to the number of dorsal scale rows present.

The name taylori isa patronym honoring Edward H. Taylor, designated "our friend and teacher" by describers. 
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