Imperfect Mimicry and the Limits of Natural Selection

Mimicry—when one organism (the mimic) evolves a phenotypic resemblance to another (the model) due to selective benefits—is widely used to illustrate natural selection’s power to generate adaptations. However, many putative mimics resemble their models imprecisely, and such imperfect mimicry represents a specific challenge to mimicry theory and a general one to evolutionary theory. Here, we discuss 11 nonmutually exclusive hypotheses for imperfect mimicry. We group these hypotheses according to whether imperfect mimicry reflects: an artifact of human perception, which is not shared by any naturally occurring predators and therefore is not truly an instance of imperfect mimicry; genetic, developmental, or time-lag constraints, which (temporarily) prevent a response to selection for perfect mimicry; relaxed selection, where imperfect mimicry is as adaptive as perfect mimicry; or tradeoffs, where imperfect mimicry is (locally) more adaptive than perfect mimicry. We find that the relaxed selection hypothesis has garnered the most support. However, because only a few study systems have thus far been comprehensively evaluated, the relative contributions of the various hypotheses toward explaining the evolution of imperfect mimicry remain unclear. Ultimately, clarifying why imperfect mimicry exists should provide critical insights into the limits of natural selection in producing complex adaptations

Notes and Comments Predator Cognition Permits Imperfect Coral Snake Mimicry

abstract: Batesian mimicry is often imprecise. An underexplored explanation for imperfect mimicry is that predators might not be able to use all dimensions of prey phenotype to distinguish mimics from models and thus permit imperfect mimicry to persist. We conducted a field experiment to test whether or not predators can distinguish deadly coral snakes (Micrurus fulvius) from nonvenomous scarlet kingsnakes (Lampropeltis elapsoides). Although the two species closely resemble one another, the order of colored rings that encircle their bodies differs. Despite this imprecise mimicry, we found that L. elapsoides that match coral snakes in other respects are not under selection to match the ring order of their model. We suggest that L. elapsoides have evolved only those signals necessary to deceive predators. Generally, imperfect mimicry might suffice if it exploits limitations in predator cognitive abilities

A Batesian mimic and its model share color production mechanisms

Batesian mimics are harmless prey species that resemble dangerous ones (models), and thus receive protection from predators. How such adaptive resemblances evolve is a classical problem in evolutionary biology. Mimicry is typically thought to be difficult to evolve, especially if the model and mimic produce the convergent phenotype through different proximate mechanisms. However, mimicry may evolve more readily if mimic and model share similar pathways for producing the convergent phenotype. In such cases, these pathways can be co-opted in ancestral mimic populations to produce high-fidelity mimicry without the need for major evolutionary innovations. Here, we show that a Batesian mimic, the scarlet kingsnake Lampropeltis elapsoides, produces its coloration using the same physiological mechanisms as does its model, the eastern coral snake Micrurus fulvius. Therefore, precise color mimicry may have been able to evolve easily in this system. Generally, we know relatively little about the proximate mechanisms underlying mimicry

Endless forms most hidden: katydids that masquerade as moss

In the cloud forests of the central range of the Colombian Andes, we discovered a species of katydid (Orthoptera: Tettigoniidae) that imitates mosses to an uncanny degree and is exceedingly difficult to detect. The camouflage exhibited by this particular katydid seems quite specific. We discuss the evolutionary consequences of this sort of specialization. Selection to maintain effective disguises can result in reproductive isolation between populations specialized for different microhabitats, which makes it reasonable to speculate that camouflage may increasing diversification rates. Camouflage could also come at the price of elevated extinction risk. This possibility must be considered because although antipredator defenses are often thought of as leading to “escape-and-radiate” dynamics where diversification follows innovation that allows expansion into new niches, recent work has shown unexpected extinction risk associated with some antipredator adaptations. Highly specialized camouflage would seem an ambiguous case because of its obvious benefits, but also potential costs such as inhabiting habitats with low carrying capacities, vulnerability to predators at high densities if predators form search images, or metabolic trade-offs with thermoregulation. Groups such as the Tettigoniidae provide a tantalizing opportunity for their exceptional diversity, wide geographic distribution, and striking array of disguises suggest that many independent evolutionary experiments have already taken place

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

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

Evaluating the utility of camera traps in field studies of predation

Artificial prey techniques—wherein synthetic replicas of real organisms are placed in natural habitats—are widely used to study predation in the field. We investigated the extent to which videography could provide additional information to such studies. As a part of studies on aposematism and mimicry of coral snakes (Micrurus) and their mimics, observational data from 109 artificial snake prey were collected from video-recording camera traps in three locations in the Americas (terra firme forest, Tiputini Biodiversity Station, Ecuador; premontane wet forest, Nahá Reserve, Mexico; longleaf pine forest, Southeastern Coastal Plain, North Carolina, USA). During 1,536 camera days, a total of 268 observations of 20 putative snake predator species were recorded in the vicinity of artificial prey. Predators were observed to detect artificial prey 52 times, but only 21 attacks were recorded. Mammals were the most commonly recorded group of predators near replicas (243) and were responsible for most detections (48) and attacks (20). There was no difference between avian or mammalian predators in their probability of detecting replicas nor in their probability of attacking replicas after detecting them. Bite and beak marks left on clay replicas registered a higher ratio of avian:mammalian attacks than videos registered. Approximately 61.5% of artificial prey monitored with cameras remained undetected by predators throughout the duration of the experiments. Observational data collected from videos could provide more robust inferences on the relative fitness of different prey phenotypes, predator behavior, and the relative contribution of different predator species to selection on prey. However, we estimate that the level of predator activity necessary for the benefit of additional information that videos provide to be worth their financial costs is achieved in fewer than 20% of published artificial prey studies. Although we suggest future predation studies employing artificial prey to consider using videography as a tool to inspire new, more focused inquiry, the investment in camera traps is unlikely to be worth the expense for most artificial prey studies until the cost:benefit ratio decreases

Dose response of the 16p11.2 distal copy number variant on intracranial volume and basal ganglia.

Carriers of large recurrent copy number variants (CNVs) have a higher risk of developing neurodevelopmental disorders. The 16p11.2 distal CNV predisposes carriers to e.g., autism spectrum disorder and schizophrenia. We compared subcortical brain volumes of 12 16p11.2 distal deletion and 12 duplication carriers to 6882 non-carriers from the large-scale brain Magnetic Resonance Imaging collaboration, ENIGMA-CNV. After stringent CNV calling procedures, and standardized FreeSurfer image analysis, we found negative dose-response associations with copy number on intracranial volume and on regional caudate, pallidum and putamen volumes (β = -0.71 to -1.37; P < 0.0005). In an independent sample, consistent results were obtained, with significant effects in the pallidum (β = -0.95, P = 0.0042). The two data sets combined showed significant negative dose-response for the accumbens, caudate, pallidum, putamen and ICV (P = 0.0032, 8.9 × 10-6, 1.7 × 10-9, 3.5 × 10-12 and 1.0 × 10-4, respectively). Full scale IQ was lower in both deletion and duplication carriers compared to non-carriers. This is the first brain MRI study of the impact of the 16p11.2 distal CNV, and we demonstrate a specific effect on subcortical brain structures, suggesting a neuropathological pattern underlying the neurodevelopmental syndromes

Curated genome annotation of Oryza sativa ssp. japonica and comparative genome analysis with Arabidopsis thaliana

We present here the annotation of the complete genome of rice Oryza sativa L. ssp. japonica cultivar Nipponbare. All functional annotations for proteins and non-protein-coding RNA (npRNA) candidates were manually curated. Functions were identified or inferred in 19,969 (70%) of the proteins, and 131 possible npRNAs (including 58 antisense transcripts) were found. Almost 5000 annotated protein-coding genes were found to be disrupted in insertional mutant lines, which will accelerate future experimental validation of the annotations. The rice loci were determined by using cDNA sequences obtained from rice and other representative cereals. Our conservative estimate based on these loci and an extrapolation suggested that the gene number of rice is ~32,000, which is smaller than previous estimates. We conducted comparative analyses between rice and Arabidopsis thaliana and found that both genomes possessed several lineage-specific genes, which might account for the observed differences between these species, while they had similar sets of predicted functional domains among the protein sequences. A system to control translational efficiency seems to be conserved across large evolutionary distances. Moreover, the evolutionary process of protein-coding genes was examined. Our results suggest that natural selection may have played a role for duplicated genes in both species, so that duplication was suppressed or favored in a manner that depended on the function of a gene

Measurement of the mid-rapidity transverse energy distribution from sNN=130\sqrt{s_{NN}}=130 GeV Au+Au collisions at RHIC

The first measurement of energy produced transverse to the beam direction at RHIC is presented. The mid-rapidity transverse energy density per participating nucleon rises steadily with the number of participants, closely paralleling the rise in charged-particle density, such that E_T / N_ch remains relatively constant as a function of centrality. The energy density calculated via Bjorken's prescription for the 2% most central Au+Au collisions at sqrt(s_NN)=130 GeV is at least epsilon_Bj = 4.6 GeV/fm^3 which is a factor of 1.6 larger than found at sqrt(s_NN)=17.2 GeV (Pb+Pb at CERN).Comment: 307 authors, 6 pages, 4 figures, 1 table, submitted to PRL 4/18/2001; revised version submitted to PRL 5/24/200

Event-by-event fluctuations in Mean pTp_T and Mean eTe_T in sqrt(s_NN) = 130 GeV Au+Au Collisions

Distributions of event-by-event fluctuations of the mean transverse momentum and mean transverse energy near mid-rapidity have been measured in Au+Au collisions at sqrt(s_NN) = 130 GeV at RHIC. By comparing the distributions to what is expected for statistically independent particle emission, the magnitude of non-statistical fluctuations in mean transverse momentum is determined to be consistent with zero. Also, no significant non-random fluctuations in mean transverse energy are observed. By constructing a fluctuation model with two event classes that preserve the mean and variance of the semi-inclusive p_T or e_T spectra, we exclude a region of fluctuations in sqrt(s_NN) = 130 GeV Au+Au collisions.Comment: 10 pages, RevTeX 3, 7 figures, 4 tables, 307 authors, submitted to Phys. Rev. C on 22 March 2002. Plain text data tables for the points plotted in figures for this and previous PHENIX publications are (will be made) publicly available at http://www.phenix.bnl.gov/phenix/WWW/run/phenix/papers.htm