Evolutionary Refugia Research Articles

The ecological and evolutionary dynamics of inselbergs.

Islands are fundamental model systems in ecology, biogeography, and evolutionary biology. However, terrestrial islands, unlike their aquatic counterparts, have received comparatively less attention. Among these land islands, inselbergs (i.e. isolated rock outcrops with diverse lithologies and a modest topographical prominence) stand out as iconic examples distributed worldwide across global biomes. Due to their durable lithology, inselbergs change slowly, persisting for tens of millions of years. In this review, we propose a biological definition for inselbergs that captures three fundamental characteristics of inselbergs from the perspective of biota. These are old age, isolation and the presence of unique microhabitats that are rare or absent in the surrounding matrix, fostering distinct communities often with unique and endemic biota. We synthesise the state of the art and formulate a set of testable hypotheses to deepen our understanding of the origins and maintenance of diversity on inselbergs, which are increasingly exposed to anthropogenic threats. By offering different habitats compared to the surrounding habitat matrix (e.g. moist microhabitats in dryland landscapes and xeric environments in humid tropical landscapes), inselbergs may allow specific lineages to thrive beyond their typical geographical limits. Particularly in drylands and degraded landscapes, inselbergs may not just provide different habitats but also act as ecological refuges or evolutionary refugia by providing a wider range of potential microhabitats than the surrounding matrix, enhancing resilience and promoting regional biodiversity. The central role of the matrix ensures that the ecological and evolutionary dynamics of inselbergs differ from those of true islands such as oceanic islands. Given that inselberg biota coexist within a terrestrial matrix, interactions between inselberg and matrix populations impact each other significantly. Over evolutionary timescales, matrix species may contract to inselberg refugia, preserving lineages while cycles of isolation and reconnection may drive speciation via a species pump. Although inselberg biodiversity has been studied predominantly from an island biogeography perspective, we argue that depending on the spatial scale, habitat specificity and mobility of the organisms considered, a range of different theories and paradigms can help explain the biogeography and local distribution patterns of different taxonomic and functional groups of inselberg species.

Die on this hill? A new monotypic, microendemic and montane vertebrate genus from the Australian Wet Tropics

AbstractThe upland rainforests of the Australian Wet Tropics (AWT) are renowned for their diverse and highly relictual biota, and are amongst the best‐studied areas of tropical rainforest in the world. Nonetheless, the evolutionary history of many endemic AWT taxa remains poorly resolved. Here we use molecular data to show that a small thigmothermic and heat‐sensitive lizard endemic to the Thornton Uplands in the northern portion of the AWT is a palaeoendemic that likely diverged from the nearest living taxa around the early Miocene. On the basis of genetic and morphological data, we remove this taxon from the genus Calyptotis where it was formerly placed, redefine Calyptotis and describing a new monotypic genus, Calorodius gen. n. This taxon is the second microendemic, monotypic and montane vertebrate genus recognised from the AWT, and highlights the potential for small areas of upland habitat in AWT to function as evolutionary refugia. However, despite the deep divergence of this lineage, there is no strong evidence that cool‐mesic environments are ancestral to the broader radiation of sphenomorphin skinks that it sits within. The impacts of future anthropogenic climate change on this new genus are difficult to predict, but a tiny range and apparent sensitivity to even relatively moderate temperatures raise concerns, especially if accelerating climate change leads to more regular drought events and heatwaves in the mountain ranges where it occurs.

Identifying groundwater‐fed climate refugia in remote arid regions with citizen science and isotope hydrology

Abstract Groundwater and surface water‐fed systems act as biodiversity hotspots and ecological refuges and evolutionary refugia in arid regions. Groundwater‐dominated systems are sustained by underground aquifers that are recharged by rain that has fallen in the distant past, while surface water‐dominated systems are fed by recent local rain or floods. Some waterbodies are fed by a mixture of these sources. Perennial, groundwater‐dominated systems will act as refuges and refugia under future rainfall declines associated with global warming. We sought to identify climate refugia, based on groundwater dominance, by using isotope hydrology to characterise water samples collected by citizen scientists across arid central Australia. There is a linear relationship between hydrogen isotopes (2H/1H, δ2H) and oxygen isotopes (18O/16O, δ18O) in rainfall. This relationship is known as the meteoric water line (MWL). By comparing our samples with the Australian MWL, and developing a local evaporation line, we were able to test the hypotheses that groundwater‐dominated systems will follow the Australian MWL while temporary systems follow the local evaporation line, and, accordingly, distinguish between groundwater and surface water‐dominated systems. The isotopic composition of samples collected over a 36‐month period was determined using isotope ratio infrared spectrometry. The electrical conductivity of each sample was recorded to determine where freshwater is available for biota within this arid region. Over 240 water samples were collected from 62 waterbodies and seven bores (groundwater wells) spanning an area of more than 250,000 km2. Approximately 75% of the samples were collected by citizen scientists and 25% by research scientists. Twenty groundwater‐dominated waterbodies, characterised by a small range of δ2H and δ18O values (c. −55 to −20‰ and c. −9 to −3‰, respectively) clustered around the long‐term mean composition of rainfall (δ2H = −37.5‰, δ18O = −6.4‰), were identified as future evolutionary refugia. These sites are likely to contain water through the most severe of droughts and will be critically important for the persistence of water‐dependent species. Based on their isotopic composition, we identified 45 waterbodies (rockholes/waterholes) as temporary or ephemeral (δ2H c. −40 to −100‰ and δ18O c. −4 to +25‰), that is, with no evidence of groundwater inflow. These, together with waterbodies supported by a mix of groundwater and surface water, can act as stepping stones and form part of the aquatic mosaic that is critical to supporting species in arid regions. Over two‐thirds of the waterholes sampled were very fresh (electrical conductivity <0.8 mS/cm), indicating that they provide the freshwater needed to support much of the regional aquatic and terrestrial fauna. All evolutionary refugia are located within protected areas (i.e. national parks or Indigenous Protected Areas), but some are subject to the impacts of feral animal species and invasive plants. Our findings indicate where control programmes and restoration actions can be prioritised to support biodiversity conservation and climate change adaptation. Our approach, combining citizen science and isotope hydrology, can be used to identify future refugia in other remote and arid regions where water scarcity is likely to increase under global climate change.

Persistence ofEtheostoma parvipinne(Goldstripe Darter) in a Single Tributary on the Periphery of Its Range

Abstract We report the occurrence of a single population of Etheostoma parvipinne (Goldstripe Darter) in a small, acidic, spring-fed stream within the Colorado River drainage following an extended period of below-average precipitation (January–September 2011), a wildland fire (September 2011), and subsequent debris and sediment flows (November 2011–May 2012). Goldstripe Darters were taken from Alum Creek (Bastrop County, TX) along with a cyprinid, a poeciliid, and several species of centrarchids in June 2012. Occurrence of Goldstripe Darters in Alum Creek provides a regional example of fish-community responses to wildland fires. In additio n, understanding the mechanism of long-term persistence in a single tributary on the periphery of a species range might offer insight into the origin of endemic fishes and persistence of disjunct fish populations within aquatic evolutionary refugia of central Texas.

Systematic revision of the marbled velvet geckos (Oedura marmorata species complex, Diplodactylidae) from the Australian arid and semi-arid zones.

Lizards restricted to rocky habitats often comprise numerous deeply divergent lineages, reflecting the disjunct nature of their preferred habitat and the capacity of rocky habitats to function as evolutionary refugia. Here we review the systematics and diversity of the predominantly saxicoline Australian marbled velvet geckos (genus Oedura) in the Australian arid and semi-arid zones using newly-gathered morphological data and previously published genetic data. Earlier work showed that four largely allopatric and genetically divergent lineages are present: Western (Pilbara and Gascoyne regions), Gulf (west and south of the Gulf of Carpentaria), Central (central ranges) and Eastern (Cooper and Darling Basins). None of these four populations are conspecific with true O. marmorata, a seperate species complex that is restricted to the Top End region of the Northern Territory. Top End forms share a short, bulbous tail whereas the other four lineages treated here possess a long, tapering tail. Morphological differences among the arid and semi-arid lineages include smaller body size, tapering lamellae and a shorter tail for the Gulf population, and a partially divided rostral scale in the Western population compared to the Central and Eastern populations. Accordingly, we resurrect O. cincta de Vis from synonymy for the Central and Eastern lineages, and regard this species as being comprised of two evolutionary significant units. We also describe the Gulf and Western lineages as new species: Oedura bella sp. nov. and O. fimbria sp. nov., respectively. We note that a predominantly arboreal lineage (the Eastern lineage of O. cincta) is more widely distributed than the other lineages and is phylogenetically nested within a saxicoline clade, but tends to have a deeper head and shorter limbs, consistent with morphological variation observed in other lizard radiations including both saxicoline and arboreal taxa.

Lineage Range Estimation Method Reveals Fine-Scale Endemism Linked to Pleistocene Stability in Australian Rainforest Herpetofauna

Areas of suitable habitat for species and communities have arisen, shifted, and disappeared with Pleistocene climate cycles, and through this shifting landscape, current biodiversity has found paths to the present. Evolutionary refugia, areas of relative habitat stability in this shifting landscape, support persistence of lineages through time, and are thus crucial to the accumulation and maintenance of biodiversity. Areas of endemism are indicative of refugial areas where diversity has persisted, and endemism of intraspecific lineages in particular is strongly associated with late-Pleistocene habitat stability. However, it remains a challenge to consistently estimate the geographic ranges of intraspecific lineages and thus infer phylogeographic endemism, because spatial sampling for genetic analyses is typically sparse relative to species records. We present a novel technique to model the geographic distribution of intraspecific lineages, which is informed by the ecological niche of a species and known locations of its constituent lineages. Our approach allows for the effects of isolation by unsuitable habitat, and captures uncertainty in the extent of lineage ranges. Applying this method to the arc of rainforest areas spanning 3500 km in eastern Australia, we estimated lineage endemism for 53 species of rainforest dependent herpetofauna with available phylogeographic data. We related endemism to the stability of rainforest habitat over the past 120,000 years and identified distinct concentrations of lineage endemism that can be considered putative refugia. These areas of lineage endemism are strongly related to historical stability of rainforest habitat, after controlling for the effects of current environment. In fact, a dynamic stability model that allows movement to track suitable habitat over time was the most important factor in explaining current patterns of endemism. The techniques presented here provide an objective, practical method for estimating geographic ranges below the species level, and including them in spatial analyses of biodiversity.

Evolutionary refugia and ecological refuges: key concepts for conserving Australian arid zone freshwater biodiversity under climate change

Refugia have been suggested as priority sites for conservation under climate change because of their ability to facilitate survival of biota under adverse conditions. Here, we review the likely role of refugial habitats in conserving freshwater biota in arid Australian aquatic systems where the major long-term climatic influence has been aridification. We introduce a conceptual model that characterizes evolutionary refugia and ecological refuges based on our review of the attributes of aquatic habitats and freshwater taxa (fishes and aquatic invertebrates) in arid Australia. We also identify methods of recognizing likely future refugia and approaches to assessing the vulnerability of arid-adapted freshwater biota to a warming and drying climate. Evolutionary refugia in arid areas are characterized as permanent, groundwater-dependent habitats (subterranean aquifers and springs) supporting vicariant relicts and short-range endemics. Ecological refuges can vary across space and time, depending on the dispersal abilities of aquatic taxa and the geographical proximity and hydrological connectivity of aquatic habitats. The most important are the perennial waterbodies (both groundwater and surface water fed) that support obligate aquatic organisms. These species will persist where suitable habitats are available and dispersal pathways are maintained. For very mobile species (invertebrates with an aerial dispersal phase) evolutionary refugia may also act as ecological refuges. Evolutionary refugia are likely future refugia because their water source (groundwater) is decoupled from local precipitation. However, their biota is extremely vulnerable to changes in local conditions because population extinction risks cannot be abated by the dispersal of individuals from other sites. Conservation planning must incorporate a high level of protection for aquifers that support refugial sites. Ecological refuges are vulnerable to changes in regional climate because they have little thermal or hydrological buffering. Accordingly, conservation planning must focus on maintaining meta-population processes, especially through dynamic connectivity between aquatic habitats at a landscape scale.