In the coming years and decades, advanced space- and ground-based observatories will allow an unprecedented opportunity to probe the atmospheres and surfaces of potentially habitable exoplanets for signatures of life. Life on Earth, through its gaseous products and reflectance and scattering properties, has left its fingerprint on the spectrum of our planet. Aided by the universality of the laws of physics and chemistry, we turn to Earth's biosphere, both in the present and through geologic time, for analog signatures that will aid in the search for life elsewhere. Considering the insights gained from modern and ancient Earth, and the broader array of hypothetical exoplanet possibilities, we have compiled a comprehensive overview of our current understanding of potential exoplanet biosignatures, including gaseous, surface, and temporal biosignatures. We additionally survey biogenic spectral features that are well known in the specialist literature but have not yet been robustly vetted in the context of exoplanet biosignatures. We briefly review advances in assessing biosignature plausibility, including novel methods for determining chemical disequilibrium from remotely obtainable data and assessment tools for determining the minimum biomass required to maintain short-lived biogenic gases as atmospheric signatures. We focus particularly on advances made since the seminal review by Des Marais et al. The purpose of this work is not to propose new biosignature strategies, a goal left to companion articles in this series, but to review the current literature, draw meaningful connections between seemingly disparate areas, and clear the way for a path forward. Key Words: Exoplanets—Biosignatures—Habitability markers—Photosynthesis—Planetary surfaces—Atmospheres—Spectroscopy—Cryptic biospheres—False positives. Astrobiology 18, 663–708.

Ground-and space-based planet searches employing radial velocity techniques and transit photometry have detected thousands of planet-hosting stars in the Milky Way. With so many planets discovered, the next step toward identifying potentially habitable planets is atmospheric characterization. While the Sun-Earth system provides a good framework for understanding the atmospheric chemistry of Earth-like planets around solar-type stars, the observational and theoretical constraints on the atmospheres of rocky planets in the habitable zones (HZs) around low-mass stars (K and M dwarfs) are relatively few. The chemistry of these atmospheres is controlled by the shape and absolute flux of the stellar spectral energy distribution (SED), however, flux distributions of relatively inactive low-mass stars are poorly understood at present. To address this issue, we have executed a panchromatic (X-ray to mid-IR) study of the SEDs of 11 nearby planet-hosting stars, the Measurements of the Ultraviolet Spectral Characteristics of Low-mass Exoplanetary Systems (MUSCLES) Treasury Survey. The MUSCLES program consists visible observations from Hubble and ground-based observatories. Infrared and astrophysically inaccessible wavelengths (EUV and Lyα) are reconstructed using stellar model spectra to fill in gaps in the observational data. In this overview and the companion papers describing the MUSCLES survey, we show that energetic radiation (X-ray and ultraviolet) is present from magnetically active stellar atmospheres at all times for stars as late as M6. The emission line luminosities of C IV and Mg II are strongly correlated with band-integrated luminosities and we present empirical relations that can be used to estimate broadband FUV and XUV (≡X-ray + EUV) fluxes from individual stellar emission line measurements. We find that while the slope of the SED, FUV/NUV, increases by approximately two orders of magnitude form early K to late M dwarfs (≈0.01-1), the absolute FUV and XUV flux levels at their corresponding HZ distances are constant to within factors of a few, spanning the range 10-70 erg cm −2 s −1 in the HZ. Despite the lack of strong stellar activity indicators in their optical spectra, several of the M dwarfs in our sample show spectacular UV flare emission in their light curves. We present an example with flare/quiescent ultraviolet flux ratios of the order of 100:1 where the transition region energy output during the flare is comparable to the total quiescent luminosity of the star E flare (UV) ∼ 0.3 L * Δt (Δt = 1 s). Finally, we interpret enhanced L(line)/L Bol ratios for C IV and N V as tentative observational evidence for the interaction of planets with large planetary mass-to-orbital distance ratios (M plan /a plan ) with the transition regions of their host stars.

The ultraviolet (UV) spectral energy distributions (SEDs) of low-mass (K-and M-type) stars play a critical role in the heating and chemistry of exoplanet atmospheres, but are not observationally well-constrained. Direct observations of the intrinsic flux of the Lyα line (the dominant source of UV photons from low-mass stars) are challenging, as interstellar H I absorbs the entire line core for even the closest stars. To address the existing gap in empirical constraints on the UV flux of K and M dwarfs, the MUSCLES Hubble Space Telescope Treasury Survey has obtained UV observations of 11 nearby M and K dwarfs hosting exoplanets. This paper presents the Lyα and extreme-UV spectral reconstructions for the MUSCLES targets. Most targets are optically inactive, but all exhibit significant UV activity. We use a Markov Chain Monte Carlo technique to correct the observed Lyα profiles for interstellar absorption, and we employ empirical relations to compute the extreme-UV SED from the intrinsic Lyα flux in ∼100 Å bins from 100-1170 Å. The reconstructed Lyα profiles have 300 km s −1 broad cores, while >1% of the total intrinsic Lyα flux is measured in extended wings between 300 and 1200 km s −1 . The Lyα surface flux positively correlates with the Mg II surface flux and negatively correlates with the stellar rotation period. Stars with larger Lyα surface flux also tend to have larger surface flux in ions formed at higher temperatures, but these correlations remain statistically insignificant in our sample of 11 stars. We also present H I column density measurements for 10 new sightlines through the local interstellar medium.

The UV environment of a host star affects the photochemistry in the atmosphere, and ultimately the surface UV environment for terrestrial planets and therefore the conditions for the origin and evolution of life. We model the surface UV radiation environment for Earth-sized planets orbiting FGKM stars at the 1AU equivalent distance for Earth through its geological evolution. We explore four different types of atmospheres corresponding to an early Earth atmosphere at 3.9 Gyr ago and three atmospheres covering the rise of oxygen to present day levels at 2.0 Gyr ago, 0.8 Gyr ago and modern Earth (following Kaltenegger et al. 2007). In addition to calculating the UV flux on the surface of the planet, we model the biologically effective irradiance, using DNA damage as a proxy for biological damage. We find that a pre-biotic Earth (3.9 Gyr ago) orbiting an F0V star receives 6 times the biologically effective radiation as around the early Sun and 3520 times the modern EarthSun levels. A pre-biotic Earth orbiting GJ 581 (M3.5V) receives 300 times less biologically effective radiation, about 2 times modern Earth-Sun levels. The UV fluxes calculated here provide a grid of model UV environments during the evolution of an Earth-like planet orbiting a range of stars. These models can be used as inputs into photo-biological experiments and for pre-biotic chemistry and early life evolution experiments.