COLUBRIDAE Pliocercus Ekpoides Cope

Home , Micrurus, Micrurus nigrocinctus, Pliocercus, Urotheca

- - Catalogue of American Amphibians and Reptiles. collected by R.M.DeOca, date of collection unknown (not examined by authors). Smith, H.M.and D. Chiszar. 200 1. Pliocercus elapoides. Urotheca elupoides: Boulenger 1894: 182. Urottteca elupoides elapoides: Arnaral 1929: 177 (part). Pliocercus ekpoides Cope Urotheca elupoides: Savage and Crother 1989:352 (pan). Variegated False Coral Snake CONTENT. Five subspecies (P. e, elapoides, P. e. aequalis, Pliocerc~iselapoides Cope 1860:253. Type locality, "Jalapa" P. e. diastemu, P. e. occidenfalis,P. e. nilmurai) are recognized. (= Xalapa), Veracruz, MCxico (amended by Smith and Tay- lor 1950). Four syntypes, Acad. Nat. Sci. Philadelphia 381% 3, an adult, a subadult, and two juveniles (sex not recorded),

FlGURE 5. Pliocercus elapoides aequalis PC2 (KU 187343) from Izabal, Guatemala (photograph courtesy of J.A. Campbell). This pattern class is dominated by mimicry of several subspecies of Micrurus

strongly triad pattern mimics that of Micrurrrs elegans (~ig;2).

FIUUKL L. MICrUrUS tieguns elegans trorn clntalapa. El Mercadito, Chiapas. MCxico (from Roze 1996). Note the contrast.-...... in design of the pattern evolved in mimicry of Micrnrus elegans in ~lior~rcuse. I elcpoides (triad, Fig. 8) and P. e. aeq~talis(modified monad, with black saddles in the red zones. Fig. 5).

FIGURE 3. Micrurus diasrema diaslema from Mexico (from Roze 1996). This model, sympatric in part with P. e. elapoides, is responsible IFIGURE 6. Micrurus diasrema sapperi from 4 km E Chial. Cayo, for the monad variant (Fig. 8). whereas Micrrrrus elegans is responsible Belize (photograph by Paul Freed, courtesy of Carnegie Museum of for the triad variant (Figs. I & 8). Natural History). This Coral Snake senles as a model for Pliocercus ela~oidesaeyualis PC2 (Fig. 5).

Quetzaltenango, Guatemala (photograph by D.G. Barker, courtesy of FIGURE 4. Pliocercus elapor~~.ucrrtirli~ PC I from Huehuetenango, J.A. Campbell). This is the most common variant of the subspecies, Guatemala (photograph by J.A. Campbell). This pattern class is domi- mimicking Micrurlrs hrowni impor~unus(Fig. 9) and others with simi- nated by mimicry of Mirr/ovs c. elegatl.r (Fig. 2). lar patterns. FIGURE 8. Syntypes of Liopliis mricirrctrrs Jan (= Pliocerrrrs elapoides el(rpoides). from Jan and Sordelli (1866: livr. 18, pl. 4. figs. 46),showing three pattern variants and single secondary temporals: no. 4 exemplifies the extreme triad pattern, with the secondary black rings fused across the red rings, much as described by Smith and Langebartel ( 1949) for a specimen from near La Gloria, Oaxaca; no. 5 exemplifies a normal triatl pattern; both nos. 4 and 5 are apparent mimics of Micrlirus clegans; no. 6 exemplifies retluction of the triads to monads, apparently mimicking Miourus diastema.

MAP. Distribution of Pliocercirs elapoides; circles indicate type localities, dots other known records. FIGURE 14. ~lzrcuselapoides wilmarai (UTA 3058) from 9 km ESETebanca, Veracruz, Mtxico (from Smith and Chiszar 1996). This specimen came from the same litter as that illustrated in Fig. 17. It is bicolored, completely lacking yellow rings, but still belongs to the tricolor comnlex. annarentlv a mimic of Micrurus I. limha/un (Fie. 15).

FIGURE 9. Micrurus browni importunus from Duefias, Guatemala (from Roze 1996). This Coral Snake serves as a model for Pliocercus elapoides diastema (Fig. 7).

Tropical "Los Tuxtlas," Veracruz, Mexico. This subspecies is the apparent model for Pliocercus elapoides tvilnlarai (Figs. 13-14)(photo-

FIGURE 10. Pliocercus elapoides diastema (UTA 2 17 16) from Quetzaltenango, Guatemala (photograph by D.G. Barker, courtesy J.A. Campbell). This uncommon variant of the subspecies has yellow rings FIGURE 16. Micrurus I. litnbafus from VolcAn Santa Marta. Los limited to the ventral surface. The other extreme has exceptionally long Tuxtlas, Veracruz, Mtxico (from Roze 1996). This Coral Snake pattern yellow rings, as in the holotype of the synonym I! andrewsipacificus. serves as the model for the ringed variant of Pliocercus elapuides Note that the locality is the same as in Fig. 7. Micrurus nigrocinctus wilmarai (Figs. 13-14).

FIGURE 11. Micrurus ni~rocinctuszunilensis from San Gednimo, Oaxaca, Mexico (from ROG1996). This pattern serves as the apparent model for Lhe sort of variant illustrated in Fig. 10.

F bum I I. rrtocercus elaporaes wrrmarar rrorn Y m c3r. leoanca, Veracruz, Mtxico (photograph courtesy of W.F. Pyburn and J.A. Campbell). This nearly patternless variant, represented by the holotype. mimics Micrurus limbatus soilosomus (Fig. 18).

FIGURE 12. Pliocercris elapoides occiden/alis (UIMNH 61401) from La Concepci6n, near Putla, Oaxaca, Mexico (from Smith and Landy 1965). This is the only known s~ecimen(of l I) with caudal triads. m

FIGURE 18. Micrurus limbarus spilosomus from VolcAn Santa Marta, 3.Pliocercus elapo~...... -.I ESE Tcbancii. Los Tuxtlas, Veracruz, Mexico (photograph courtesy of R.C. Vogt and Veracruz, Mexico (photograph by R. Powell). This fom~mimics Mi- J.A. Campbell). This subspecies is the apparent model for the nearly crur1r.r limbatus limbatus (Figs. 15-16). patternless variant of Pliocercus elapoides wilmarai (Fig. 17). However, no subspecies are recognized by some recent review- and figure 51 1 illustrates P. e. wilmarai). Color figures of P. e. ers (e.g., Wilson and Meyer 1985, Savage and Crother 1989). elapoides (figure I E) and P. e. aequalis (figure 1 F) appeared in who also gave this species more extensive parameters, includ- Mertz (1996) and in Greene and McDiarrnid (1981); the latter ing P. andrewsi and P. bicoloras here interpreted (see also Smith does not show the yellow rings characteristically present and Chiszar 2001a). (Campbell and Lamar 1989, figures 5 14-6, even in figure 5 13, vestiges of yellow rings are present dorsally). Smith and Landy DEFINITION. Pliocercus elapoides is a small, generally tri- (1965) included black and white illustrations of P. e. elapoides, colored colubrid snake of the tricolor I! elapoides complex, with P. e. dirrstema, and P. e. occidentalis (reproduced in Smith and a maximum known SVL of 416 mm (female, A.K. Smith Chiszar 1996). Pliocercus e. aequnlis was illustrated and de- 1969)(TL calculated at about 670 mm, assuming the tail to be scribed in Fischer ( I88 1 ), Bocourt (1886), and Giinther ( 1893). the mean of 38% of TL). The tail in males is 384%of TL ( i Weber (1945) showed in color a P. e. elapoides in the claws of a = 42), in females, 3541% (1= 39)(A.K. Smith 1969). White Snake Hawk. Smith and Langebartel(1949) illustrated a Head scalation is as follows: supralabials usually 8 (91%, N P. e. e1apoide.s from the Isthmus of Tehuantepec. Jan and Sordelli = 607 of 665), occasionally (8%) 9 (N = 52), rarely (I %) 7 (N= (1866) illustrated the three syntypes of Liophis rricincrus (= P. 6); infralabials usually 9 (53%, 340 of 642) or 10 (45%, 286). e. elapoides). Pliocercus e. diastema was illustrated in Bocourt seldom (2%, 10) or I I (1%. 6); preoculars usually 2 (84%, 560 (1 886). Pliocercus e. nequalis was depicted in color in Wilson of 668), occasionally 1 (9%. 61, in one of which the loreal con- and Meyer (1982, 1985). A black and white figure of P. e. di- tacts the eye) or 3 (7%, 47); postoculars usually 2 (99%, 667 of astenla is in Alvarez del Toro (1973, 1982). in Smith and 676), rarely 3 (I%, 9); anterior temporal always single (N = Chrapliwy (1957). in Mertens (1952b), and in Hecht and Marien 340); secondary temporal single on both sides in 80% (N = 27 I), (1956). Greene (1969) portrayed f? e. ~vilmaraiin black and single on one side and double on the other in 9% (N = 30), white. double on both sides in 11% (N = 39); tertiary temporals usually 2 on both sides (94%, 3 19 of 340), single on one side in 1% DISTRIBUTION. Pliocercus el(~poidesis the southernmost (N = 3), on both sides in 1 % (N = 3), triple on one side in 4% (N species of the tricolor group of the genus, occurring in rainforest = 12), on both sides in 1% (N = 3). Ventral scales in males and cloud forest habitats on Atlantic slopes from central number 119-132 (7 = 125.2, N= 148), in females 124-139 (7 = Veracruz, Mexico, to western Honduras, and on Pacific slopes 132.1, N = 184). Subcaudal scales in males number 90-121 (x from western Oaxaca and perhaps adjacent Guerrero, Mexico. = 107.4, N= 86), in females 85-1 12 ( F = 98.7, N = 83). Of the to El Salvador and perhaps Honduras. This geographic range 148 males examined, 58% had complete tails, whereas only 45% marginally overlaps that of P. dimidiafu.r. in western Honduras, of 184 females did. and is extensively sympatric with f! andrewsi in the northern In life, most populations have yellow rings between red and part of the Yucatrin Peninsula. The population in the Los Tuxtlas black rings. Yellow rings vary in length from 0.54 scales in uplift of southern Veracruz (P. e. wilmarai) is completely iso- length, usually 1 more or less. Primary black rings on body lated from all others. number 2-25, and may be complete or not. Red zones lack black, or may have some or all dorsal scales lightly to heavily FOSSIL RECORD. None. black-tipped. Secondary black saddles may be present or absent; when present, they are peripheral in the red zones, form- PERTINENT LITERATURE. Most of the literature on this ing triads with the primary black rings (in some specimens they species is distributional or variational, and is sumniarized in the extend toward each other across the red zones, and rarely com- subspecies accounts. Much of the rest concerns mitnicty, sum- pletely replace the red middorsally, although not ventrally, be- marized in Smith and Chiszar (200121). Motphological varia- cause all secondary black saddles are incomplete ventrally). The tion in the species as a whole was described by Smith and Chiszar nuchal black ring often is somewhat longer than others. A yel- (1996). Savage and Crother (1989) also reviewed some aspects low parietal ring is present. of variation, but included P. arldrervsi and P. hicolor with P. elapoides. Variation in material from Honduras was described DIAGNOSIS. Pliocercus elapoides is a member of the P. by Wilson and Meyer (1982, 1985). An excellent review of elapoides complex (with black rings separated from each other variation in the whole species, although including P. birolor, by complete red rings, typically bordered by yellow rings sepa- was included in an unpublished dissertation by A.K. Smith rating the black rings from the red rings), thus eliminating all ( 1969). members of the P. euryzorrus complex (P. eurjzonus, P. The natural history of mostly this species. although covering dimidiatus). This species is distinguished from other members the whole genus, is reviewed in Smith and Chiszar (1996). of the tricolor complex by having the posterior infralabial sepa- Wilson and McCranie (1998) and Pelcastre-Villafuerte and rate from the posterior labiogenial (thus eliminating P hicolor). Flores-Villela (1992) sumniarized habitat preferences. Seib and black rings not as few as 7, unless they are much less than (1980) reported a severe reaction to the venom off? e. diasfemo. half as long as their pale interspaces (thus eliminating P. noted by numerous authers (e.g., Minton 1990). Wilson (1968) andreu~si).The isolated Los Tuxtlas population (P. e. wilmrrrai) described and illustrated the fracture plane in the caudal verte- lacks yellow rings, as do some individuals in central western brae of P. e. aequalis (reproduced in Bellairs and Bryant, 1985), Guatemala and adjacent Chiapas (P. e. aequalis), but they are although Arnold (1984, 1988) and Savage and Crother (1989) readily distinguished from members of the P. eurjionus com- argued for intervertebral breakage. Thatcher (1966) reported plex by having the primary black rings much shorter than the trematodes. Predation on P. e. elapoides by a White Snake Hawk pale interspaces. was noted by Weber (1945). Vernacular names were suggested in Smith and Chiszar DESCRIPTIONS AND ILLUSTRATIONS. The most de- ( 1996). and Greene ( 1997) applied the name Halloween Snake. tailed descriptions and illustrations (in black and white) are in Other literature includes Flores-Villela (1993, keys), Flores Smith and Chiszar (1996). Several color illustrations appear in et al. (1995, keys), Liner (1994, checklist and common names). Campbell and Lamar (1989). where figure 512 is of P. e. di- Ramirez-Velizquez and Sigler-Morena (1993, Chiapas). Sav- nstema (as labelled), and figures 5 13-6 are all of P. e. aequalis, age (1966, herpetofaunal origins), Wilson and Meyer (1982, as here interpreted (figure 5 10 depicts P bicolor hoharrsmithi, 1985, Honduras), and Zhao et al. (1993, name). Literature explicitly relating to P1iocerclt.s elnpoides elnpoides tioned, but its scope and taxonomic content have been a source includes Alvarez del Toro (1960, 1973, 1982, Chiapas), Booth of considerable disagreement. One extreme is a single species (1959. Chiapas), Campbell et al. (1995. Veracruz), Casas Andreu (including F! bicolor and /? anrlrew~si),with no subspecies, as et al. (1996, Oaxaca. part), Flores et al. (1991, Oaxaca), Fugler understood by Savage and Crother (1 989), as well as many oth- and Dixon (1958. Veracruz), Gonzilez-Romero et al. (1991, ers. The other extreme is exenlplified by Smith and Smith Veracruz), Greene ( 1969, mention), Liner (1960, comparisons), (1976). who accepted 9 taxa that we now know are referable to Pelcastre-Villafuerte and Flores-Villela (1992, Veracruz, habi- P. elapoides. We herein recognize little more than half (5) that tat), Perez-Higareda and Navarro (1980, Veracruz, barrier), number, as initially proposed by Smith and Chiszar (1996). with Pkrez-Higareda and Smith ( 199 I, Veracruz), Peters and Orejas- modifications they adopted later (2000,2001b). Miranda (1970, key), Peterset al. (1986, key), Ram'rez-Bautista Although our argument for acceptance of P: ondrewsi as a et al. (1993, Veracruz), Salvin (1 861, comparisons), Smith (1941, separate species may be questioned, we regard the validity of 1942.1943.1987, Veracruz, taxonomy), Smith et al. (1 989. com- the subspecies we here recognize in F! elnpoides as the chief parisons), Smith and Langebartel (1949, Oaxaca), Smith and bone of contention relative to the tricolor complex of Pliocercus. Smith (1976, 1993, literature). Smith and Taylor (1945, check- Our recognition of five subspecies rests on the observation that list). Smith and Taylor (1950, type localities), Stuart (1950, those five units exhibit unique parameters of variation of at least Guatemala). Tanner ( 1957, Chiapas), Troschel ( 1862, literature), reasonable magnitude (better than 80% separability), and pre- Wake and Johnson (1989, Chiapas), and Zhao (1993, list). sumably reflect distinctive genetic compositions. Each has Literature explicitly relating to Pliocercus elnpoides nequolis evolved unique tendencies apparently influenced for the most includes Alvarez del Toro (1973, 1982, Chiapas), Boulenger part by the associated models for mimicry. (1882, Zoological Record), Cadle (1984, molecular systemat- We have been impressed by the relatively minor variation in ics, Quintana Roo), Campbell and Vannini (1988. Guatemala, two of the subspecies (F! e. diastemu, F! e. occidentnlis). as com- habitat, abundance). Cochran (1961. list of type specimens), pared with the considerable variation in I? e. elnpoides, and the Duellman (1 963, El Peten, Guatemala), Hahn ( 197 1, Hondu- very extensive variation in /? e. neqtrctlis and /? e. uvilrnarai, all ras). Henderson and Hoevers (1975, Belize). Jackson (1973, apparently correlated with multiple sympatric Coral Snake pat- Honduras), Kramer (1978, list of type specimens). Lee (1980, terns (whether monads or triads, for example; see discussion in 1996, 2000, Yucatan Peninsula), Markel (1990, color photo), Smith and Chiszar 2001a). Martin del Campo (1945, Chichen Itza, Yucatan), Marx (1958, In recognizing five subspecies of /? elopoides, we have been 1976, list of type specimens), McCoy (1966, 1970, Guatemala), influenced by the geographic consistency of certain character- Mendelson (1990, Guatemala), Miiller (1973, dispersal center), states in each taxon; geographically erratic variants, particularly Neill(1965, Belize), Neill and Allen (1961, Belize). Peters and as in P. e. oequulis and F! e. wilmnrni, we regard as infrataxo- Orejas-Miranda (1970, 1986. keys), Smith (1943, 1987, nomen- nomic chromotypes. Our analyses sought geographic character clature), Smith and Smith (1976, 1993, literature), Smith and consistencies, whether or not associated with mimicry of Coral Taylor (1945, 1950, checklist, type localities). Stuart (l950,Alta Snakes. Thus the perceived geographic ranges of the five sub- Vera Paz, Guatemala), Taylor (1944, list of type specimens), species of P. elnpoides are allopatric. Some taxa (F! e. w~ilnrarui) Wilson (1968, tail fracture plane), Wilson and McCranie (1998. are strongly dichopatric, others parapatric or presumably so. Honduras), and Wilson and Meyer (1985, Honduras). Good integrades are known from one intermediate locality be- Literature explicitly relating to P1iocerc~t.selal>oides dinsterncr tween the ranges of F! e. elnpoides and P. e. dinstenla (Smith includes Alvarez del Toro (1960, 1973, 1980, Chiapas), and Landy 1965); they are to beexpected between the latter and Campbell and Lamar (1989, photograph), Casas-Andreu et al. P. e. occidenmlis. but isolation of the latter prevents integrada- (1996, Oaxaca), Greene (1969, 1973, eggs, Senckenberg Mu- tion with any subspecies except P. e. diostemo. Intergrades may seum), Hecht and Marien ( 1956, pattern, mimicry, photograph), occur between P. e. elapoides and F! e. aequalis. but probably Landy et al. (1966, food. habitat. mimicry), Lynch and Smith not between the latter and F! e. dinsterna, because of the pres- (1965, Oaxaca). Maldonado (1953, mention). Martinez- ence of an isolating physiographic/ecologicalbarrier. Pliocerc~ts Castellano and Muiioz-Alonzo (1998, mention), Mertens (1952b, e. ~,ilnzaraiis isolated from all other congeneric taxa. hence can El Salvador), Peters and Orejas-Miranda (1970, 1986, keys), have no intergrades. Seib (1980, envenomation), Smith ( 1942, 1959, 1987, key, Gua- After decades of experience with the Guatemala herpetofauna. temala), Smith and Landy (1965, Chiapas, Oaxaca). Smith et Stuart ( 1963) recognized three populations of R elnpoirlrs. al- al. (1964. list of type specimens), Smith and Smith (1976. 1993, though his nomenclature was partly different than that used here. literature), Smith and Taylor (1945, 1950. checklist. type His populations were designated as F! e. dinsterna. exactly the localties). Stuart (1948, 1950, 1963, Guatemala), and Wilson same as interpreted here; I! e. ~eq~mlis.essentially the same as and Meyer (1982, 1985. Honduras). our /? e. crequnlis PC1 (pattern class I); and /? e. salvini, essen- Literature explicitly relating to Pliocercirs elapoides occiden- tially the same as our /? e. trequnlis PC2. More recent material rnlis includes Peters and Orejas-Miranda (1970, 1986, keys), has changed his analysis only by making the latter two insepa- Savage and Crother (1989, rejected), Smith and Smith (1976, rable taxonomically, although they remain two different 1993, literature), Snlith and Taylor (1966, checklist), and Wil- chromotypes. son and Meyer ( 1982. 1985, rejected). Literature explicitly relating to Pliocercirs elapoides n~ilrnar~ti ETYMOLOGY. Cope's ( 1860) name elopoides is an adjec- includes Campbell and Lamar ( 1989, color photograph), Greene tive formed from elrps (Latin for "snake"), the generic name of et al. (1998. mimicry). Pelcastre-Villafuerte and Fiores-Villela the Coral Snake when Cope worked, and the suffix -0ide.7, "like," (1992, habitat). Perez-Higadera (sic)et al. (1987, Los Tuxtlas a Latin term derived from the Greek -eides. Thus the first spe- herpetology), Perez-Higareda and Smith (1990, 1991, Los cies described in the genus was at once recognized as a mimic Tuxtlas), Ramirez-Bautista (1978, Los Tuxtlas), Ramirez- of Coral Snakes. Bautista and Nieto-Montes de Oca (1997, Los Tuxtlas). Vogt The name neqrinlis is a Latin adjective meaning "equal," and (1997, serpentine communities, Los Tuxtlas), and Vogt et al. refers to the "equidistant black bands" (Salvin 1861) of the ho- (1997, herpetofaunal list. Los Tuxtlas). lotype. Salvin ( 186 1 ) and other early workers did not distin- guish between primary and secondary rings, and they also failed REMARKS. The validity of I? einpoides has never been ques- to note the presence of yellow rings. interpreting a11 of the light areas as red. Bocoult's name diastema is a Greek noun mean- b. Primary black rings on body 14 or fewer (79%; minimum ing "space between" or "interval," and was applied in reference number of subcaudals between primary black rings on com- to the widely spaced black rings in the type series. The name is plete tail 9 or more (69%); black rings on tail 9 or fewer not an adjective and therefore does not change its ending to (94%); ratio, middorsal length of the primary black ring agree with the gender of the generic name to which it is re- adjacent to the shortest interval between, divided by the ferred. The Latin word occidentalis ("of the west") was applied latter, 33% or less (94%): ratio, middorsal length on the to its taxon in reference to its geographic position as the primary black ring on body adjacent to the longest inter- westernmost population of the genus on Pacific slopes. The val, divided by the latter, 31% or less (81%) ...... name u~ilrrrarrriis a genitive patronym honoring William P. Mara ...... P e. aequrr1i.s PC2 for his editorial and herpetocultural skills, the latter of which are dedicated to exercise on the present genus, where they are 1. Pliocercus elapoides elapoides Cope so important to its taxonomy. Deppe's False Coral Snake

KEY TO SUBSPECIES. The two pattern classes of P1iocercrt.s elal~oidesCope 1860:253. See species synonymy. Pliocerccrs elapoiries oeq~ra1i.scan be seen to differ substantially. E1upochr~t.sdely~ei Peters 1860:293. Type locality, "~Mexico," as is evident in the following key. However, because they are restricted to "Xalapa, Veracmz." by Smith and Taylor ( 1950). inconsistent in geographic occurrence. occurring both together Holotype, Zoologischen Museum Berlin (ZMB) 241 1 (Bauer and by themselves over considerable areas, they are not et al. 1995). an adult female. collected by F. Deppe, date of populationally distinctive and hence do not qualify as subspe- collection unknown, but between 1824 and 1830 (Taylor cies. A parallel exists in the distinctive chromotypes of Lnrnpi-o- 1969)(not examined by authors). peltis getuln caIifr,rrrioe. P1iocercct.s deppei: Gunther 1862:53. Liophis (Cos~niosophis)rriciilcrtrs Jan 1863a:289. Type local- I. a. No black rings or saddles on body or tail except nuchal ity, "Messico," restricted to "Xalapa, Veracruz." by Smith and and anal; or, if present. no yellow rings, pale zones all red, Taylor (1950). Syntypes, one adult and two juveniles (sex even ventrally ...... I? e. ~vilmnrrri not recorded), cited in the original description and also by b. Black rings or saddles present throughout body and tail; Jan ( 1863b) to be in the Nati~rhistorischesMuseum in Vienna, yellow rings present at least ventrally between red and black the Copenhagen University Museum, and the Museo Civico rings or saddles ...... 2 di Storia Naturale in Milan. Collectors and dates not recorded (not examined by authors). 2. a. Minimum middorsal or paravertebral scale lengths in Urorheca elapoides elrlpoides: Amaral 1929: 177 (part). nuchal black ring, posterior to parietals, 6 or more (P e. P1iocercu.s elapoides elrrpoides: Smith 194 1: 1 19. diasretrra. 94%: P e. occidentolis, 100%; I? e. elapoides. E1apochrolt.s deppei: Smith and Taylor 1945: 110. Lup.s~t.sCO- 38%) ...... 3 latni. b. Minimum middorsal or paravertebral scale lengths in Pliocerclts e1iipoide.s: Greene and McDiarmid 198 1: 121 1 (part). nuchal black ring, posterior to par~etals,5 or fewer (I? e. Urotheca elcrpoides: Savage and Crother 19893356 (part). aequcrlis PC I, 95%: P e. creclualis PC2, 9 1%; I? e. clap- oide.~,62%) ...... 5 DIAGNOSIS. This subspecies of I? elapoides is distingukhed from all others by the combination of a relatively short parietal 3. a. No distinct triads on tail (828) ...... P. e. occideiltct1i.s pale ring not involving scales anterior or posterior to parietals b. Distinct triads on tail (I? e. dirr.sten~n.92%; I? e. elapoides, (95%); distinct triads on body (92%) and tail (97%), yellow 97%) ...... 4 rings distinct; a relatively short nuchal ling. covering 5 or fewer nliddorsal or paravertebral scale lengths (66%); and infralabials 4. a, lnfralabials 9-9 or fewer (87%); outer rings of triads at usually 9-9 or fewer (87%). least 1.5 dorsal scales in length on at least part of body (74%) ...... P e. elrrpoi~le~ REMARKS. This subspecies is extensively mimetic of two b. Infralabials 9-10 or more (80%); outer rings of triads very differently patterned Coral Snakes. Micntrus elegcrr~s(with shorter than 1.5 dorsal scales in length throughout body triads, resulting in evolution of supernumerary black saddles (958) ...... I? e. dirrsretna appearing as triads; on the contraly in P. e. nequalis, mimicking the same species, they appear usually as modified monads) and 5. a. Parietal pale ring narrow, not involving any scale anterior M. dinsrernu (with monads, resulting in reduction, even loss, of or posterior to parietals (95%) ...... P e. clapoides the supernumerary black saddles). Variation in 76 specimens b. Parietal pale ring longer, involving 95% or more of the was described by Smith and Chiszar (1996). length of parietals, or parts of scales anterior (frontal. supraoculars, postoculars) or posterior (occipitals) to the pa- 2. Pliocercus elapoides aequalis (Salvin) rietal~(P e. czeq~t~1i.sPC l, 75%; P e. neqlralis PC2, 84%) Salvin's False Coral Snake

Pleiocerc,us oequalis Salvin 1861:227. Type locality, "San 6. a. Primary black rings on body 15 or more (91'3%): minimum Ger6nimo and the neighboring mountains ... of Vera Paz," in number of subcaudals between primary black rings on com- Baja Vera Paz, Guatemala. Holotype, British Museum of plete tail 6 or fewer (91 %); maximum number of subcaudals Natural History (BMNH) 1946.1.1.4. an adult female, col- between primary black rings on complete tail 8 or fewer lected by R. Owen. IS60 (not examined by authors). (88%); black rings on tail 10 or more (88%); ratio. rnid- Pliocetr,rrs ~eqrrn1i.c:Cope 1862:72. dorsal length of the primary black ring on body adjaccnt to Elapncht-us aequrr1i.s. varietas: Miiller 1878x662. the shortest interval between. over the latter. 34% or more Pliocercus saltlirlii Muller 1878a:709 (substitute name for (79%): ratio, middorsal length of the primary black ring Elcrpochr-trs a~clunlis,varietas Muller (1878). Type locality. on body adjacent to the longest interval. di\.ided by the "Vera Paz. Guatemala." Holotype. Naturhistorischen Mu- latter, 38% or more (86%) ...... F! e. ireqrtnlis PC1 seum Berlin (ZMB) 1495. an addlt (sex not recorded). collected by G. Bernoulli, date of collection unknown (not ex- present in the middle of the red zones, which in some speci- amined by authors). mens are completely obliterated dorsally. The yellow rings are Liophis tricinctus: Miiller 1878b:750 (nee Jan). frequently very short and may be absent dorsally, although al- Pliocercrrs sawii Fischer 1881:225. Type locality, "Cobhn," ways present ventrally. Pattern Class 2 (PC2) occurs mostly Alta Vera Paz, Guatemala. Holotype, Naturhistorischen Mu- outside of the range of M. elegans, but also erratically within seum Stuttgart 2012, an adult female, collected by F. Sarg, that range. It mimics the tricolor monad M. dinstemo and is date of collection unknown (not examined by authors). characterized by a simple pattern of widely spaced primary black Li0phi.r elcrpoides neqrralis: Bocourt 1886:637. rings and minimal pigmentation in the intervening zones (see Elapochrus arqrralis: Giinther 1893: 106. Smith and Chiszar 1996). Urotheca elapoides aeqrtalis: Werner 1896:348. P1iocercrr.s e. nequalis has at times been regarded as bicol- Urothecn neqrmlis: Gadow 19 1 1 :17. ored. lacking yellow rings, and has been so illustrated in several Urothecrr e1clpoide.s elupoides: Taylor 1939 ( 1940):469 (part). works (e.g., Giinther 1893). This taxon. however, is not a mern- Pliocercrrs elnpoiclrs loticollnris Smith 194 1 :122. Type local- ber of the P. errryzorrus complex. Of 88 specimens of P e. ity. "Tenosique. Tabasco. Mexico." Holotype. National Mu- nequtrlis we examined, 19 (22%) showed no clear dorsal evi- seum of Natural History (USNM) 110767, an adult female, dence of yellow rings, although in some of those exceptions the collected by H.M. Smith, 30 June 1939 (examined by au- yellow may have been too faded or discolored to be evident. thors). However, the yellow rings were evident, at least ventrally, in Pliocercus elcrpoides schmiclti Smith 1942: 16 1. Type locality, all. None of the 32 specimens examined of PC2 had yellow "Chichen Itzi, Yucatin. Mexico." Holotype, Museum of rings reduced dorsally; that variation appeared only in some of Comparative Zoology (MCZ) 26843. an adult female, col- PC1, mostly in those from Chiapas and in Guatemala from Alta lected by E.N. Thompson, date of collection unknown (ex- Vera Paz and Baja Vera Paz. amined by authors). If these two pattern classes were clearly allopatric, they would Pliocercrts elcrpoiclrs sernicinctrrs Schmidt 194 1502. Type lo- qualify as si~bspecies(per Stuart 1963). but we now know that cality, "Double Falls, west of Stann Creek. Belize." Holo- they are pop~~lationallyinseparable. The situation is much like type, Field Museum of Natural History (FMNH) 72471, an that in Lrtrnpropeltis getula cciliforniae. adult female?collected by I.T. Sanderson, 13 December 1939 Variation in 88 specimens was reported by Smith and Chiszar (examined by authors). (I966). PIiocerc~rseltrl~oides snlt~inii: Stuart 1948:67. P1iocercrr.s elnpoir1e.s: Stuart 1948:71 (part). 3. Pliocercus elapoides diastema (Bocourt) P1iocercrr.s eLrt?zonu.s oequalis: Stuart 1948:72. Bocourt's False Coral Snake Pliocerrrts bicolor: Stuart 1948:72 (nee Smith). Pliocercrrs elopoides elapoides: Smith 1960:223 (nec Cope). Liophis elnpoides [elcrpoides]: Bocourt 1886:635 (part; four of Pliocercus Inticollrris: Liner 1960:2 17. five specimens and the figures are of P. e. dinsterna). Pliocrrcus elnpoides dinsremrts: Henderson and Hoevers 1975: Liophis elapoides diastema Bocourt 1886:636. Type locality, 44 (nee Bocourt). "Plateau of Guatemala." Syntypes, adult males, MusCum Na- Pliocrrcus nequolis aequnlis: Snlith 1987:~~~~. tional d'Histoire Naturelle, Paris (MNHN) 132-133, collec- Urotheco elapoides: Savage and Crother 1989:356 (part). tor and date of collection unknown (not examined by authors). Pliocrrcus elupoides neqrralis: Smith and Chiszar 199653. Urotheccr e1apoide.s elnpoides: Slevin 1939:402 (nee Cope). Urotheca (Pliocerc~ts)elcipoides: Stafford 1999:48. Pliocvrcus elnpoides dicrstenrrtr: Smith 1941: 120. Unjustified Pliocercus psychoides Smith and Chiszar 1996:34. Type local- emendation of the subspecific name (Mertens 1952b). ity, unknown. given as "Brasil or Venezuela." but obviously Pliocercus elnpoides elapoides: Stuart 1948:7 1 (nee Cope). in error. probably central western Guatemala (Smith and Pliocercus elapoidrs diastemn: Mertens 1952x70. Chiszar 200 I b). Holotype. American Museum of Natural His- Pliocercus elapoides salt~adorensisMertens 1952a:9 1. Type tory (AMNH) 4433. an adult female, collector and date of locality, "Finca San JosC, 1150 m, nr Santa Tecla, Depto. La collection unknown (examined by authors). Libertad, El Salvador." Holotype, Natur-Museum Sencken- berg (SMF) 42301, an adult female, collected by C. Holtmann. DIAGNOSIS. This subspecies of P. elapoides is distinguished 22 July 1950 (not examined by authors). from all others by the combination of a short nuchal black ring, Pliocerc~rsnndrewsi pacijicus Smith and Chrapliwy 1957:233. usually (91%) extending no more than 6 dorsal scale lengths Type locality, "Finca Custepeque, 40 mi ESETonalL. Chiapas, posterior to the parietals, and a long, pale parietal ring. usually Mexico." Holotype, University of Illinois Museum of Natu- (75% in PC 1. 84% in PC2) involving 95%: or more of the pari- ral History (UIMNH) 40832, an adult male. collected by T. etals or parts of scales anterior (frontal, supraoculars, MacDougall, 18 November 1956 (examined by authors). postoculars) or posterior (occipitals) to the parietals. Pliocerc~rselapoides e1opoide.s x diasterr~us:Lynch and Smith 1965: 168. REMARKS. This subspecies is extensively mimetic of Mi- Pliocercrrs elupoides: Greene 1973: 15 1 (part). crrtrrts dinstrrnrr and M. hippoerepis (both tricolor monads; Sav- Urotkeccr elcrpoides: Savage and Crother 1989:356 (part). age and Slowinski 1990) and M. ele~ons(basically a tricolor triad, but incompletely a tricolor pentad), hence populations are DIAGNOSIS. This subspecies of P. elopoides diff'ers from highly variable where both species occur together. but are less others of the species as follows: from P. e. aequolis PC1 in hav- so elsewhere. ing 13 or fewer primary black rings on body (98% in 159 speci- Two pattern classes occur in this subspecies. Pattern class 1 mens versus 5% in 58); from P e. crec~ualisPC2 in having a (PCI) is mostly limited to the range of the tricolor triad species relatively long nuchal ring. at least 6 dorsals long near midline Micrrrrtr.~elegctr7.s. where the tricolor modad M. di~istern(1also (94% in 160 versus 9% in 32); from fl e, occiderltnlis in having serves as a model. and is characterized by a complex and erratic well developed triads on tail (91% in 159 versus 9% in 11); pattern of numerous black rings or saddles, primary and sec- from f! e. elapoides in having 9-10 or more infralabials (8 1 % ondary, more or less matching one or both of the model patterns in 155 versus 13% in 69). and triads on body weaker or absent, (see Smith and Chiszar 1996). Black saddles are frequently the outer (incomplete) rings less than 1.5 dorsals long (95% in 160 versus 26% in 74): and from F! e. wilmnrrri in regularly Pliocercus wilmarui Smith, Perez-Higareda, and Chiszw 1996: possessing complete yellow rings (versus absence). 76. Type locality, "5.6 road mi ESE Tebanca (= Tabanca), Los Tuxtlas region, southern Veracruz, [Mexico]." Holotype REMARKS. This subspecies is extensively mimetic of the University of Texas at Arlington (UTA) 3159, an adult fe- tricolor monad Coral Snakes Micrurus bogerti, M. browni, M. male, collected by J. Darling, R. Harris, and W.F. Pyburn, 20 Irrrifrrsciatus,M. ni~rocinctus,and M. stuarti. The minimal vari- June 1966 (examined by authors). ability of F! e. diastema is most likely correlated with the basic Pliocercus elapoides wilmarai: Smith and Chiszar 2000: 18. pattern similarity of all species of sympatric Coral Snake species. The usual pattern in F! e. diastema does embrace a varia- DIAGNOSIS. This subspecies off? elapoides differs from tion in number of primary black rings from 6-14, and the yel- all conspecifics by lacking black rings or saddles except for the low rings vary from a maximum of 3-4 dorsal scale lengths to a nuchal and anal marks, or, if they are present, yellow rings are minimum of complete absence dorsally. These variations to a missing both dorsally and ventrally. This subspecies, from the certain extent reflect different Micrurus species with different Los Tuxtlas uplift in southern Veracruz, is geographically iso- monad patterns. Variation in 160 specimens was reported by lated from all other populations of the genus. Smith and Chiszar (1996). REMARKS. This subspecies is extensively mimetic in its 4. Pliocercus elapoides occidentalis Smith and Landy ringless lnorph of the equally ringless, spotted, sympatric Mi- MacDougall's False Coral Snake crrrrus limbatus .spilosornus; and in its ringed morph with the sympatric tricolor triad M. elegnns and tricolor monads M. di- Pliocerc~rsekipoides occider~talisSmith and Landy 1965: I. Type rtsternn (oligocricoid) and M. I. limborrts (polycricoid). The locality, "La Concepci6n. nr Putla, Oaxaca, Mexico." Holo- characteristic loss of yellow rings in F! e. wilmrrrai may have type, University of Illinois Museum of Natural History resulted from mimicry of M. elegans, in which yellow is not or (UIMNH) 61410, an immature male, collected by T. Mac- scarcely evident. Dougall, 24 May 1965 (examined by authors). This taxon was regarded in its original description and in Pliocercus elapoides: Greene and McDiarmid 198 1 : 12 11 (part). Smith and Chiszar (1996) as a full species thought to occur sym- Urotheca elnpoides: Savage and Crother 1898:356 (part). patrically in the Los Tuxtlas region with an equally isolated population of I? e. aequalis PC I that Perez-Higareda and Smith DIAGNOSIS. This subspecies of f? elapoides differs from F! earlier (1986) regarded as R bicolor, because of the absence of e. aequnlis PC1 in having 12 or fewer primary black rings on yellow rings, before the defining scale character of that species body (100% versus 2%) and 7 or fewer on tail (100% versus was known. Both taxa mimic the same Coral Snake models 0%); from F! e. neqtmlis PC2 in having a longer nuchal ring, and, not surprisingly, resemble one another. They differ, how- near midline covering a minimum of at least 6 dorsal scale ever, in apparently lacking yellow rings both dorsally and ven- lengths (100% versus 9%); from F! e. elapoides in usually lack- trally (see color figures in Greene and McDiarmid 1981, and ing distinct triads on the tail (91% versus 0%). and triads on Perez-Higareda and Smith 1991), whereas apparently all F! e. body weak or absent, outer saddles less than one dorsal scale in ctequalis PC1 possess yellow rings at least ventrally if not dor- length where present (100% versus 8%); from F! e. rvihnarni in sally. All chromotypes occur sympatrically within the Los regularly possessing yellow rings (versus absence); and from ?! Tuxtlas area (PCrez-Higareda and Smith 1991), hence they are e. diastema in usually lacking distinct triads on the tail (91% most likely consubspecific. The apparent overlap of variation versus 9%), having a lesser maximum number of subcaudals in in the Los Tuxtlas population and F! e. aequalis PC1 in reduc- the primary black rings (4 or fewer, 89% versus 14%), lesser tion of the yellow rings suggests that the two populations are minimum number of subcaudals in the primary black rings (3 best regarded as distinct subspecies, not species, until consider- or fewer. 89% versus 35%). and a longer light parietal ring (cov- ably more material is at hand. ering 95% of the parietals. or extending onto scales anterior or Most subspecies of F! elapoides that we recognize are distin- posterior to the parietals, 84% versus 33%). guished on the basis of characters that seem not to be affected by mimicry, but F! e. wilmnrai is an exception, being distin- REMARKS. The only species of Micrtirus likely to occur guished solely on the basis of mimicral features. However, F! e. within the range of F! e. occider~talisis M. b. bogerti (Roze 1996). wilmarai is distinct from other taxa of Pliocercus, just as M. although no records exist at present for any Coral Snake in that lirnhatus (and its two distinctive subspecies) is from other Coral area. If none occur. their absence may have been influential in Snake species, and both taxa are equally limited geographically. the differentiation off? e. occidentnlis from the widely distrib- Consequently, recognizing both Los Tuxtlas populations as taxo- uted R 4. diastema, the only other taxon of the genus occuring nomically distinct is not unreasonable. on Pacific slopes, and sympatric with several species of Coral Variation is summarized by Smith et al. (1996) and supple- Snakes. Variation in 11 specimens was reported by Smith and mented by Smith and Chiszar (1996). Chiszar (1996). ACKNOWLEDGMENTS. We are indebted to Kraig Adler 5. Pliocercus elapoides wilmarai Smith, Perez-Higareda, and for counsel and bibliographic aid. to Ernest A. Liner for help Chiszar with the literature, to Frank Van Breukelen for technical assis- Mara's False Coral Snake tance, to Deborah Aguiar for preparation of the map. to Comstock Publishing Company for permission to reproduce figures 37. Pliocerc~tselapoides: Greene 1969327 (part). 73, 5 11, and 5 15 from Ca~npbelland Lamar (1989), and to Pliocercus elapoides schmidti: Ramirez-Bautista 1977: 137 (nec Krieger Publishing Company for permission to reproduce pat- Smith). terns 8. 13.22. and 30 from Roze (1996). P1iocercrt.s elopoides scilvir~ii:Perez-Higareda and Smith 1986: 125 (part). LITERATURE CITED Pliocercus hicolor: PCrez-Higareda and Smith 1986: 127 (nec Smith). Alv;~rezdelToro. M. 1960. Los Reptiles de Chiapas. Inst. Zool. Estado. Urorhecn elapoides: Savage and Crother 1989:356 (part). Tuxtla Gutierrez, Chiapas. -. 1973. Los Reptiles de Chiapas. 2nd ed. Gob. 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