Coral Snake Mimics Research Articles

Genetic mechanisms and correlational selection structure trait variation in a coral snake mimic.

Covariation among traits shapes both phenotypic evolution and ecological interactions across space and time. However, rampant geographical variation in the strength and direction of such correlations can be particularly difficult to explain through generalized mechanisms. By integrating population genomics, surveys of natural history collections and spatially explicit analyses, we tested multiple drivers of trait correlations in a coral snake mimic that exhibits remarkable polymorphism in mimetic and non-mimetic colour traits. We found that although such traits co-occur extensively across space, correlations were best explained by a mixture of genetic architecture and correlational selection, rather than by any single mechanism. Our findings suggest that spatially complex trait distributions may be driven more by the simple interaction between multiple processes than by complex variation in one mechanism alone. These interactions are particularly important in mimicry systems, which frequently generate striking geographical variation and genetic correlations among colour pattern traits.

Geographic variation in mimetic precision among different species of coral snake mimics

Batesian mimicry is widespread, but whether and why different species of mimics vary geographically in resemblance to their model is unclear. We characterized geographic variation in mimetic precision among four Batesian mimics of coral snakes. Each mimic occurs where its model is abundant (i.e. in 'deep sympatry'), rare (i.e. at the sympatry/allopatry boundary or 'edge sympatry') and absent (i.e. in allopatry). Geographic variation in mimetic precision was qualitatively different among these mimics. In one mimic, the most precise individuals occurred in edge sympatry; in another, they occurred in deep sympatry; in the third, they occurred in allopatry; and in the fourth, precise mimics were not concentrated anywhere throughout their range. Mimicry was less precise in allopatry than in sympatry in only two mimics. We present several nonmutually exclusive hypotheses for these patterns. Generally, examining geographic variation in mimetic precision - within and among different mimics - offers novel insights into the causes and consequences of mimicry.

Diverse Evidence for the Decline of an Adaptation in a Coral Snake Mimic

Losses of adaptations in response to changed selective pressures are evolutionarily important phenomena but relatively few empirical examples have been investigated in detail. To help fill this gap, we took advantage of a natural experiment in which coral snake mimics occur on two nearby tropical islands, one that has coral snake models (Trinidad) and one that lacks them (Tobago). On Tobago, an endemic coral snake mimic (Erythrolamprus ocellatus) exists but has a relatively poor resemblance to coral snakes. Quantitative image analysis of museum specimens confirmed that E. ocellatus is a poor mimic of coral snakes. To address questions related to the functional importance of this phenotype, we conducted a field experiment on both islands with snake replicas made of clay. These results clearly indicated a strong inter-island difference in predator attack rates where snake replicas that resembled coral snakes received protection in Trinidad but not in Tobago. Further, a molecular phylogenetic analysis of the ancestry of E. ocellatus revealed that this poor coral snake mimic is deeply nested in a clade of good coral snake mimics. These data suggest that the lack of coral snakes on Tobago altered the selective environment such that the coral snake mimicry adaptation was no longer advantageous. The failure to maintain this ancestral feature in allopatry provides a compelling example of how losses of complex adaptations can occur.

A survey on some biochemical and pharmacological activities of venom from two Colombian colubrid snakes, Erythrolamprus bizona (Double-banded coral snake mimic) and Pseudoboa neuwiedii (Neuwied's false boa)

Colombian colubrid snake venoms have been poorly studied. They represent a great resource of biological, ecological, toxinological and pharmacological research. We assessed some enzymatic properties and neuromuscular effects of Erythrolamprus bizona and Pseudoboa neuwiedii venoms from Colombia. Proteolytic, amidolytic and phospholipase A2 (PLA2) activities were analyzed using colorimetric assays and the neuromuscular activity was analyzed in chick biventer cervicis (BC) preparations. The venom of both species showed very low PLA2 and amidolytic activities; however, both exhibited high proteolytic activity, which in E. bizona venom surpassed that of P. neuwiedii venom. E. bizona and P. neuwiedii venoms provoked partial neuromuscular blockade, which was more prominent in P. neuwiedii venom. E. bizona venom (30 μg/ml) induced a significant potentiation of the contracture response to exogenous ACh (110 μM), which was not accompanied by twitch height alteration, whereas the highest venom concentration (100 μg/ml) inhibited contracture responses to both ACh and KCl (40 mM). In contrast, P. neuwiedii venom (30 and 100 μg/ml) caused significant reduction in the contracture responses to exogenous ACh and KCl. The morphological analyses showed high myotoxic effects in the muscle fibers of BC incubated with either venoms; however, they are more prominent in the P. neuwiedii venom. Our results suggest that the myotoxicity of the venom of the two Colombian species can be ascribed to their high proteolytic activity. An interesting data was the potentiation of the ACh-induced contracture, but not the twitch height, caused by E. bizona venom, at a concentration that is harmless to muscle fibers integrity. This phenomenon remains to be further elucidated, and suggest that a possible involvement of post-synaptic receptors cannot be discarded. This work is a contribution to expand the knowledge on colubrid venoms; it allows envisaging that the two venoms offer the potential to go further in the identification of their components and biological targets.

Unlinked Mendelian inheritance of red and black pigmentation in snakes: Implications for Batesian mimicry

Identifying the genetic basis of mimetic signals is critical to understanding both the origin and dynamics of mimicry over time. For species not amenable to large laboratory breeding studies, widespread color polymorphism across natural populations offers a powerful way to assess the relative likelihood of different genetic systems given observed phenotypic frequencies. We classified color phenotype for 2175 ground snakes (Sonora semiannulata) across the continental United States to analyze morph ratios and test among competing hypotheses about the genetic architecture underlying red and black coloration in coral snake mimics. We found strong support for a two-locus model under simple Mendelian inheritance, with red and black pigmentation being controlled by separate loci. We found no evidence of either linkage disequilibrium between loci or sex linkage. In contrast to Batesian mimicry systems such as butterflies in which all color signal components are linked into a single "supergene," our results suggest that the mimetic signal in colubrid snakes can be disrupted through simple recombination and that color evolution is likely to involve discrete gains and losses of each signal component. Both outcomes are likely to contribute to the exponential increase in rates of color evolution seen in snake mimicry systems over insect systems.

More than mimicry? Evaluating scope for flicker-fusion as a defensive strategy in coral snake mimics

Abstract Coral snakes and their mimics often have brightly colored banded patterns, generally associated with warning coloration or mimicry. However, such color patterns have also been hypothesized to aid snakes in escaping predators through a “flicker-fusion” effect. According to this hypothesis, banded color patterns confuse potential predators when a snake transitions from resting to moving because its bands blur together to form a different color. To produce this motion blur, a moving snake’s bands must transition faster than the critical flicker-fusion rate at which a predator’s photoreceptors can refresh. It is unknown if coral snakes or their mimics meet this requirement. We tested this hypothesis by measuring the movement speed and color patterns of two coral snake mimics, Lampropeltis triangulum campbelli and L. elapsoides, and comparing the frequency of color transitions to the photoreceptor activity of the avian eye. We found that snakes often produced a motion blur, but moving snakes created a blurring effect more often in darker conditions, such as sunrise, sunset, and nighttime when these snakes are often active. Thus, at least two species of coral snake mimics are capable of achieving flicker-fiision, indicating that their color patterns may confer an additional defense aside from mimicry.

More than mimicry? Evaluating scope for flicker-fusion as a defensive strategy in coral snake mimics

More than mimicry? Evaluating scope for flicker-fusion as a defensive strategy in coral snake mimics

Spatial and Temporal Drivers of Phenotypic Diversity in Polymorphic Snakes

Color polymorphism in natural populations presents an ideal opportunity to study the evolutionary drivers of phenotypic diversity. Systems with striking spatial, temporal, and qualitative variation in color can be leveraged to study the mechanisms promoting the distribution of different types of variation in nature. We used the highly polymorphic ground snake (Sonora semiannulata), a putative coral snake mimic with both cryptic and conspicuous morphs, to compare patterns of neutral genetic variation and variation over space and time in color polymorphism to investigate the mechanistic drivers of phenotypic variation across scales. We found that strong selection promotes color polymorphism across spatial and temporal scales, with morph frequencies differing markedly between juvenile and adult age classes within a single population, oscillating over time within multiple populations, and varying drastically over the landscape despite minimal population genetic structure. However, we found no evidence that conspicuousness of morphs was related to which color pattern was favored by selection or to any geographic factors, including sympatry with coral snakes. We suggest that complex patterns of phenotypic variation in polymorphic systems may be a fundamental outcome of the conspicuousness of morphs and that explicit tests of temporal and geographic variation are critical to the interpretation of conspicuousness and mimicry.

Selection overrides gene flow to break down maladaptive mimicry

Predators typically avoid dangerous species, and batesian mimicry evolves when a palatable species (the 'mimic') co-opts a warning signal from a dangerous species (the 'model') and thereby deceives its potential predators. Because predators would not be under selection to avoid the model and any of its look-alikes in areas where the model is absent (that is, allopatry), batesian mimics should occur only in sympatry with their model. However, contrary to this expectation, batesian mimics often occur in allopatry. Here we focus on one such example--a coral snake mimic. Using indirect DNA-based methods, we provide evidence suggesting that mimics migrate from sympatry, where mimicry is favoured, to allopatry, where it is disfavoured. Such gene flow is much stronger in nuclear genes than in maternally inherited mitochondrial genes, indicating that dispersal by males may explain the presence of mimetic phenotypes in allopatry. Despite this gene flow, however, individuals from allopatry resemble the model less than do individuals from sympatry. We show that this breakdown of mimicry probably reflects predator-mediated selection acting against individuals expressing the more conspicuous mimetic phenotype in allopatry. Thus, although gene flow may explain why batesian mimics occur in allopatry, natural selection may often override such gene flow and promote the evolution of non-mimetic phenotypes in such areas.

PREDATION BY FREE-RANGING BIRDS ON PARTIAL CORAL SNAKE MIMICS: THE IMPORTANCE OF RING WIDTH AND COLOR.

Many of the venomous New World coral snakes (Micrurus and Micruroides) have a distinctive pattern of red, black, and yellow rings (Campbell and Lamar 1989; Savage and Slowinski 1992), which typically appear in the sequence red-yellow-black-yellow (the tricolor monad of Savage and Slowinski 1992) repeated multiple times on each snake. Relatively harmless coexisting snakes in several different genera have a similar appearance, and most recent investigations have concluded that these are cases of Batesian mimicry (Greene and McDiarmid 1981; Pough 1988a; Campbell and Lamar 1989; Savage and Slowinski 1992). Though it is clear that a precise mimic of a dangerous or unpalatable model often gains protection from predation (see Waldbauer 1988 for a review), the gradual evolution of mimicry requires that partial mimics gain some fitness benefit from even a poor resemblance to model species (Fisher 1958; Sheppard 1959). There is evidence that partial mimics gain limited protection from predation in some insect systems (e.g., Morrell and Turner 1970; Pilecki and O'Donald 1971; Shideler 1973). A large number of Neotropical snakes have some elements of the coral snake pattern (Pough 1988b), but there is very little information on the protective effects of partial mimics, largely due to the extreme difficulty of observing predation events in the field. Brodie (1993, following Madsen 1987) pioneered the use of plasticine replicas of coral snakes to gather data on rates of predation by free-ranging birds in the natural habitat of coral snakes. The soft plasticine retains the imprint of any attempted predation, which can be used to identify the predator as bird, mammal, etc. (Brodie 1993). Using this method, Brodie (1993) showed for the first time that the coral snake pattern reduces the rate of avian predation for replicas of both true coral snakes and coral snake mimics. Here we extend Brodie's (1993) method to determine bird attack rates on partial coral snake mimics that have color and pattern combinations not found in any living snake. The coloration of coral snakes includes a number of elements that can vary independently: ring color, ring width, and order of the arrangement of the rings. There is no historical infor-

Evolution of coloration, urotomy and coral snake mimicry in the snake genusScaphiodontophis(Serpentes: Colubridae)

The Neotropical hinged-tooth, coral snake mimics of the genus Scaphiodontophis are characterized by extremely long and disproportionately thick tails that are extremely fragile. Both the coloration and tail structure are putative antipredator devices. While all examples have components of the coloration that match those of the venomous coral snakes (family Elapidae), the range of variation is extreme, leading to controversy on the status of various populations, including nine named taxa. Individual, ontogenetic and geographic variation in scutellation and head, body and tail coloration were analysed to evaluate population status and possible evolutionary trends based on a sample of 183 examples from Mexico, Central America and Colombia. Variation in subcaudal counts show population differences (higher in Mexico and upper Central America) but are not congruent with geographic variation in coloration. Generally snakes from north of Nicaragua and from central and eastern Panama have a pattern of dyads (black-light-black bands separating red bands), those from Atlantic slope Nicaragua to western Panama a pattern of monads (light-black-light bands separating the red ones) and those from Colombia have both pattern types on the same snake. The dyads and/or monads may be present the length of the body and tail, restricted to the anterior part of the body or on the entire body or on the anterior part of the body and on the tail. Two or more of these variants may occur at a single geographic locality or only a single one may be present. Head and nuchal colour patterns (Z, A, V, and Du) are relatively consistent geographically. The Atlantic slope Guatemala, Belize and Honduras population have the A pattern, those of Nicaragua, Costa Rica and western Panama the V pattern, and those in Colombia a Du pattern. Other populations have the Z coloration. Intermediate conditions in coloration of the body and tail and head and neck are found at localities intermediate between the main pattern types, indicating intergradation among adjacent populations. Consequently, we regard these snakes as representative of a single species, Scaphiodontophis annulatus Duméril and Bibron and the eight other names applied to various populations and individuals as synonyms. Analysis of colour pattern leads us to the conclusion that the tricolour pattern evolved from a uniform one through a lineate-spotted condition (usually present on the non-tricolour portions of the snake) through a bicolour red and black pattern to the dyadal condition. The monadal pattern in turn was derived from the dyadal one. The data further indicates that tricolour components first appeared anteriorly and progressively expanded posteriorly. The evolutionary sequence for the head and nuchal pattern appears to be A → Z → V → Du S. annulatus has a series of jaw and tooth specializations designed for rapid processing of hard-bodied prey found during diurnal foraging in the leaf-litter. Urotomy in this species involves intervertebral tail-breakage (pseudoautotomy) without regeneration. Evidence is presented supporting the long-tail multiple break hypothesis as applicable to Scaphiodontophis and other snakes with similar tail morphology (specialized pseudoautotomy). This is in contrast to snakes with similar tail morphology (specialized pseudoautotomy). This is in contrast to Coniophanes and other snakes with a high incidence of urotomy having long but unspecialized tails (unspecialized pseudoautotomy) without multiple breaks over time. All Scaphiodontophis colour patterns have a general resemblance to that of venomous coral snakes and offer protection from generalizing predators having innate or other triggered responses to coral snake colours. The aposematic effect is enhanced by tail thrashing and head twitching behaviours. The characteristic foraging pose of S. annulatus , which tends to expose the head and anterior body, makes even the incomplete tricolour pattern effective as an antipredator defense. No evidence supports the idea that tail thrashing or the incomplete tricolour pattern directs the predator attacks to the tail to expedite pseudoautotomy. Coral snake mimicry and specialized pseudoautotomy are shown not to be co-evolved and pseudautotomy seems to have evolved long before mimetic coloration in this genus.

Evolution of coloration, urotomy and coral snake mimicry in the snake genus Scaphiodontophis (Serpentes: Colubridae)

The Neotropical hinged-tooth, coral snake mimics of the genusScaphiodontophisare characterized by extremely long and disproportionately thick tails that are extremely fragile. Both the coloration and tail structure are putative antipredator devices. While all examples have components of the coloration that match those of the venomous coral snakes (family Elapidae), the range of variation is extreme, leading to controversy on the status of various populations, including nine named taxa. Individual, ontogenetic and geographic variation in scutellation and head, body and tail coloration were analysed to evaluate population status and possible evolutionary trends based on a sample of 183 examples from Mexico, Central America and Colombia. Variation in subcaudal counts show population differences (higher in Mexico and upper Central America) but are not congruent with geographic variation in coloration. Generally snakes from north of Nicaragua and from central and eastern Panama have a pattern of dyads (black-light-black bands separating red bands), those from Atlantic slope Nicaragua to western Panama a pattern of monads (light-black-light bands separating the red ones) and those from Colombia have both pattern types on the same snake. The dyads and/or monads may be present the length of the body and tail, restricted to the anterior part of the body or on the entire body or on the anterior part of the body and on the tail. Two or more of these variants may occur at a single geographic locality or only a single one may be present. Head and nuchal colour patterns (Z, A, V, and Du) are relatively consistent geographically. The Atlantic slope Guatemala, Belize and Honduras population have the A pattern, those of Nicaragua, Costa Rica and western Panama the V pattern, and those in Colombia a Du pattern. Other populations have the Z coloration. Intermediate conditions in coloration of the body and tail and head and neck are found at localities intermediate between the main pattern types, indicating intergradation among adjacent populations. Consequently, we regard these snakes as representative of a single species,Scaphiodontophis annulatusDumeril and Bibron and the eight other names applied to various populations and individuals as synonyms. Analysis of colour pattern leads us to the conclusion that the tricolour pattern evolved from a uniform one through a lineate-spotted condition (usually present on the non-tricolour portions of the snake) through a bicolour red and black pattern to the dyadal condition. The monadal pattern in turn was derived from the dyadal one. The data further indicates that tricolour components first appeared anteriorly and progressively expanded posteriorly. The evolutionary sequence for the head and nuchal pattern appears to be A → Z → V → DuS. annulatushas a series of jaw and tooth specializations designed for rapid processing of hard-bodied prey found during diurnal foraging in the leaf-litter. Urotomy in this species involves intervertebral tail-breakage (pseudoautotomy) without regeneration. Evidence is presented supporting the long-tail multiple break hypothesis as applicable toScaphiodontophisand other snakes with similar tail morphology (specialized pseudoautotomy). This is in contrast to snakes with similar tail morphology (specialized pseudoautotomy). This is in contrast toConiophanesand other snakes with a high incidence of urotomy having long but unspecialized tails (unspecialized pseudoautotomy) without multiple breaks over time. AllScaphiodontophiscolour patterns have a general resemblance to that of venomous coral snakes and offer protection from generalizing predators having innate or other triggered responses to coral snake colours. The aposematic effect is enhanced by tail thrashing and head twitching behaviours. The characteristic foraging pose ofS. annulatus, which tends to expose the head and anterior body, makes even the incomplete tricolour pattern effective as an antipredator defense. No evidence supports the idea that tail thrashing or the incomplete tricolour pattern directs the predator attacks to the tail to expedite pseudoautotomy. Coral snake mimicry and specialized pseudoautotomy are shown not to be co-evolved and pseudautotomy seems to have evolved long before mimetic coloration in this genus.

The colouration of the venomous coral snakes (family Elapidae) and their mimics (families Aniliidae and Colubridae)

The bright coloured, highly venomous coral snakes, Leptomicrurus, Micrurus and Micruroides (family Elapidae) and a series of harmless or mildly toxic mimics form an important component of the snake fauna of the Americas. Coral snake patterns are defined as any dorsal pattern found in any species of venomous coral snake and/or any dorsal pattern containing a substantial amount of red, pink or orange distributed so as to resemble that of some species of venomous coral snake. The components of coral snake colouration are described and four principal dorsal patterns are recognized: unicolour, bicolour, tricolour and quadricolour. The tricolour patterns may be further clustered based on the number of black bands or rings separating the red ones as: monads, dyads, triads, tetrads or pentads. A detailed classification of all coral snake colour patterns is presented and each pattern is illustrated. The taxonomic distribution of these patterns is surveyed for mimics and the 56 species of highly venomous coral snakes. Among the latter, the most frequent encountered patterns are tricolour monads, tricolour triads and bicolour rings, in that order. No venomous coral snakes have a tricolour dyad, tricolour tetrad or quadricolour pattern. As many as 115 species of harmless or mildly toxic species, c. 18% of all American snakes, are regarded as coral snake mimics. The colouration and behavioural traits of venomous coral snakes combine to form a significant antipredator defence of an aposematic type. The mimics in turn receive protection from predators that innately or through learning avoid coral snake colour patterns. The precise resemblances in colouration between sympatric non-coral snakes and venomous coral snakes and the concordant geographic variation between the two strongly support this view. Batesian mimicry with the highly venomous coral snakes as the models and the other forms as the mimics is the favoured explanation for this situation. It is further concluded that a number of species in the genera Elaphe, Farancia, Nerodia and Thamnophis, although having red in their colouration, should not be included in the coral snake mimic guild.

The status of Pliocercus and Urotheca (Serpentes: Colubridae), with a review of included species of coral snake mimics

The generic name Urotheca Bibron, 1843 is revived for a group of Neotropical colubrid snakes diagnosed by a long, thickened but fragile tail and the presence of a specialized naked pocket on the asulcate surface of the hemipenial capitulum. Urotheca includes those species previously placed in the lateristriga group of the genus Rhadinaea and the coral snake mimics usually referred to the genus Pliocercus. The many names based upon the coral snake mimics are shown to represent two species at most: Urotheca elapoides, a bicolour (red and black) or tricolour (red, yellow and black) banded or ringed form found in Mexico and northern Central America and U. euryzona, which is usually bicolour (red, yellow or white and black) and ranges from Nicaragua to western Ecuador. Coloration in U. elapoides resembles closely that of sympatric species of venomous coral snakes. Local variation in coloration and a geographic trend in the colour of the light rings (usually red in the north, white to the south) in U. euryzona parallels similar colour variation in the sympatric venomous coral snake Micrurus mipartitus. These patterns of variation add strong support to the idea that the two species are mimics of the highly venomous coral snakes. Urotheca, including the non-mimetic species U. decipiens, U. fulmceps, U. guentheri, U. lateristriga, U. multilineata and U. pachyura, shares the characteristic of a very long and disproportionately thickened and fragile tail with the coral snake mimics of the distantly related genus Scapkiodontophis. Members of both genera have a very high proportion (about 50%) of the tails broken indicating a probable predator escape device. Breakage is intercentral, with a calcified cap developing over the tip of the distal surface of the new terminal vertebra unlike the situation in many lizards where there is an intracentral fracture septum and the tail is regenerated.

An Alternative Solution to the Coral Snake Mimic Problem (Reptilia, Serpentes, Elapidae)

A review of various solutions proposed by investigators of the coral snake mimic problem precedes a suggested alternative explanation. Pseudomimicry is the term applied to the condition under which similar color patterns have arisen independently of natural selection in unrelated sympatric species occupying similar habitats. It is further suggested that the resemblances among Micrurus fulvius, Cemophora coccinea, and Lampropeltis triangulum are an example of pseudomimicry.

THE CORAL SNAKE “MIMIC” PROBLEM IN PANAMÁ

student of evolution, if his attention is called to contemporary phenomena in Animal Kingdom, cannot fail to note cases of extraordinary superficial resemblance between animals of quite diverse relationships. term and voluminous body of literature has resulted. In most general terms may be called word to describe situation (or an evolutionary problem) in. which external appearance, confined to certain area and certain size range, is shared by species of diverse ancestry. In many of these situations (and coral snake problem is case in point) some species are definitely venomous, poisonous, or distasteful as food while others are innocuous and edible. In all these situations evolutionary problem of superficial resemblance (the observational data) challenges evolutionary student. An almost complete acquaintance with history, facts, and status of coral snake mimic problem may be gained by perusal of four papers; those of Cope 1860, Wallace 1867, Gadow 1911, and Sternfeld 1913. Cope first called attention to existence of problem in following words (p. 262): The subject of prevalence of peculiar shades and arrangements of colors throughout certain geographical districts is one of much interest to zoologists, the Elaps of South America is represented in same region by black and red ringed Oxyrhopes, Erythrolamprus, Pliocercus, Lampropeltis, etc. Wallace, in discussing problem, proposed solution: a very remarkable and instructive case of true Gadow, writing after extensive field experience in Mexico, objected to idea of mimicry on number of grounds as follows: There are more mimickers . . . than species of Elaps, there are large districts of unquestionably Elaps-terrain where harmless species and individuals form great majority. The range of harmless species in typical dress of Elaps often extends far beyond that of nearest species of Elaps. have not yet come across single instance of what may be called occurrence by side. This in spite of fact that he had previously written (1908) several kinds of harmless snakes, belonging to different groups, inhabit same districts together with poisonous coral snake, and, what is more, they lead same kind of life under rotten stumps and in ants nests. He proposed two other solutions, as follows: (1) at dusk . . . black, alternating with red produces an effacive blur; i.e., coloration of these snakes is concealing; and (2) because America [sic] environment favours production of red in snakes . . . Scientist . . . means, but does not like to say: loci. Sternfeld's paper is defense of mimicry against some of Gadow's criticisms, and an attack on second of Gadow's solutions. He suggests that superiority in number of species of mimics over coral snakes is point in favor of, not against, mimicry. He claims that coincidence of range is much better than Gadow implied, and adds important point of coincidence in size, coral snake coloration being manifested only in snakes of coral snake dimensions. Gadow's by side criticism he considers absurd, and ridicules it in most amusing paragraph I have ever read in scientific paper. With regard to environment in America and Genius loci he says, concerning this geheimii-