Exploring Paleogene Tibet's warm temperate environments through target enrichment and phylogenetic niche modelling of Himalayan spiny frogs (Paini, Dicroglossidae).

The Cenozoic topographic development of the Himalaya-Tibet orogen (HTO) substantially affected the paleoenvironment and biodiversity patterns of High Asia. However, concepts on the evolution and paleoenvironmental history of the HTO differ massively in timing, elevational increase and sequence of surface uplift of the different elements of the orogen. Using target enrichment of a large set of transcriptome-derived markers, ancestral range estimation and paleoclimatic niche modelling, we assess a recently proposed concept of a warm temperate paleo-Tibet in Asian spiny frogs of the tribe Paini and reconstruct their historical biogeography. That concept was previously developed in invertebrates. Because of their early evolutionary origin, low dispersal capacity, high degree of local endemism, and strict dependence on temperature and humidity, the cladogenesis of spiny frogs may echo the evolution of the HTO paleoenvironment. We show that diversification of main lineages occurred during the early to Mid-Miocene, while the evolution of alpine taxa started during the late Miocene/early Pliocene. Our distribution and niche modelling results indicate range shifts and niche stability that may explain the modern disjunct distributions of spiny frogs. They probably maintained their (sub)tropical or (warm)temperate preferences and moved out of the ancestral paleo-Tibetan area into the Himalaya as the climate shifted, as opposed to adapting in situ. Based on ancestral range estimation, we assume the existence of low-elevation, climatically suitable corridors across paleo-Tibet during the Miocene along the Kunlun, Qiangtang and/or Gangdese Shan. Our results contribute to a deeper understanding of the mechanisms and processes of faunal evolution in the HTO.

Influences of ski-runs, meadow management and climate on the occupancy of reptiles and amphibians in a high-altitude environment of Italy.

Alpine ecosystems harbour a rich and highly specialised biodiversity, which is particularly susceptible to anthropogenic disturbances such as habitat loss and fragmentation as well as to climate change. Combined with other forms of land-use conversion, construction and maintenance of ski resorts can have severe consequences on alpine biodiversity. In this study, we show how one amphibian and two reptile species, namely Rana temporaria, Zootoca vivipara and Vipera berus, respond to such impacts by means of a multi-season occupancy analysis. We found all three species both in and outside ski-runs, showing that these habitats do not necessarily preclude their occurrence. Contrarily, this is influenced more by microhabitat availability, such as ground vegetation, humid areas and rock cover, rather than by macro-characteristics like elevation or habitat type. Moreover, we found a climatic influence on the year-to-year occupancy change of the species, with activity-month conditions being more relevant than overwintering ones. Our results demonstrate how, in the specific case of reptiles and amphibians, ski resorts do not necessarily limit species' occurrence and that a mild series of management actions might secure the species' persistence in the area.

Variation in claw morphology among the digits of Bent-toed geckos (Cyrtodactylus: Gekkota: Gekkonidae).

BACKGROUND: Ecomorphological studies of lizards have increasingly employed comparison of claw morphology among species in relation to spatial niche use. Typically, such studies focus on digit IV of the autopodia, especially the pes. Uniformity of claw morphology among digits is more often implicitly assumed than tested. RESULTS: Using four species of Cyrtodactylus, comprising two generalist and two scansorial taxa that use different substrates, we examined whether claw morphology is uniform among digits and among species. We found that, within each species, ventral claw curvature is uniform across all digits whereas there are small but insignificant differences in ventral claw length and claw depth. The claws of the pes of each species are longer and deeper than those of the corresponding digits of the manus. The claw of digit I of each species is significantly shorter and shallower on both autopodia compared to those on digits IV and V (digit I, including its claw, is idiosyncratically variable among lizards in general). CONCLUSIONS: We conclude that digit IV is an adequate representative of claw form in each species and exhibits variation among species, thereby serving as an exemplar for use in studies of potential discrimination between ecomorphological types in studies of Cyrtodactylus.

Captive breeding, embryonic and larval development of Ranitomeyavariabilis (Zimmermann & Zimmermann, 1988), (Anura, Dendrobatidae).

A solid basis to address the conservation challenges of amphibians requires an increased knowledge on their natural history and biology. Recent data on reproductive modes in amphibians suggest that they are much more complex and variable than previously thought but understudied. However, detailed information on the reproductive history is especially important to fill the current knowledge gaps. Following recent taxonomic changes in Ranitomeyavariabilis, information about captive-breeding management, image-based measurements of total length and surface area of the silhouette for individuals from embryonic to metamorphic development, and detailed larval staging for captive-bred specimens are provided from French Guiana. The development of R.variabilis from the stage eight (Gosner 1960) through metamorphosis took 79 to 91 days (n = 6) with a survival rate of 46%. The developmental stages largely matched those of the generalized staging system of Gosner (1960), with differences in the stages when labia and teeth differentiation and atrophy of the oral apparatus occurred. Compared with other studies the total length of R.variabilis tadpoles was greater at given stages than those of R.variabilis from a Peruvian population and those of the sister species, R.amazonica. Other studies concerning growth curves based on surface area data revealed that R.variabilis tadpoles at peak size were larger than those of R.amazonica, R.imitator, R.reticulata, R.sirensis, and R.vanzolinii, but smaller than R.benedicta. Our results represent the first embryonic and larval staging for R.variabilis, and detailed information is provided on their initial life phases. These data may facilitate the identification of R.variabilis tadpoles in the wild, as well helping to clarify the biogeographical distribution and taxonomic arrangement of the species. In addition, knowledge is added to the captive-breeding methodology for the species.

Global diversity patterns of larger benthic foraminifera under future climate change.

Global warming threatens the viability of tropical coral reefs and associated marine calcifiers, including symbiont-bearing larger benthic foraminifera (LBF). The impacts of current climate change on LBF are debated because they were particularly diverse and abundant during past warm periods. Studies on the responses of selected LBF species to changing environmental conditions reveal varying results. Based on a comprehensive review of the scientific literature on LBF species occurrences, we applied species distribution modeling using Maxent to estimate present-day and future species richness patterns on a global scale for the time periods 2040-2050 and 2090-2100. For our future projections, we focus on Representative Concentration Pathway 6.0 from the Intergovernmental Panel on Climate Change, which projects mean surface temperature changes of +2.2°C by the year 2100. Our results suggest that species richness in the Central Indo-Pacific is two to three times higher than in the Bahamian ecoregion, which we have identified as the present-day center of LBF diversity in the Atlantic. Our future predictions project a dramatic temperature-driven decline in low-latitude species richness and an increasing widening bimodal latitudinal pattern of species diversity. While the central Indo-Pacific, now the stronghold of LBF diversity, is expected to be most pushed outside of the currently realized niches of most species, refugia may be largely preserved in the Atlantic. LBF species will face large-scale non-analogous climatic conditions compared to currently realized climate space in the near future, as reflected in the extensive areas of extrapolation, particularly in the Indo-Pacific. Our study supports hypotheses that species richness and biogeographic patterns of LBF will fundamentally change under future climate conditions, possibly initiating a faunal turnover by the late 21st century.

Larval description of Theloderma albopunctatum (Liu & Hu, 1962) (Anura: Rhacophoridae) from Northern Vietnam, with comparison between the North-Vietnamese and North-eastern Thai clades.

The mossy frogs of the genus Theloderma Tschudi comprise 28 described taxa (Sivongxay et al. 2016; Frost 2022), which are distributed from north-eastern India and Myanmar to southern China, across the peninsula of Indochina and Malaysia, to Indonesia (Poyarkov et al. 2015; Frost 2022). Theloderma albopunctatum is a small-sized taxon that is assigned to the T.-asperum species complex (Poyarkov et al. 2015, 2018; Sivongxay et al. 2016; Dever 2018). For long time, it has been believed to be a synonym of T. asperum. However, genetic analyses revealed that both taxa show significant differences. Currently, populations south of the Isthmus of Kra (southern Thailand, Malayan peninsular) are assigned to T. asperum, while populations north of it (southern China, northern and central Vietnam, adjacent Laos, south-eastern Cambodia) are assigned to T. albopunctatum. In addition, this species complex might contain further cryptic species (cf. Nguyen et al. 2015; Poyarkov et al. 2015) and according to Chunskul et al. (2021) four genetic groups do exist: group A comprises T. albopunctatum from southern and central Vietnam, Laos, central and north-eastern Thailand; group B is composed of populations from northern Vietnam and China; group C ranges from north-western Vietnam to northern Thailand and Myanmar; and group D is distributed in northern Vietnam (Thanh Hoa).

Phylogenetic and morphological influence on habitat choice in moisture-harvesting horned lizards (Phrynosoma spp.).

In previous studies, the superhydrophilic skin of moisture-harvesting lizards has been linked to the morphological traits of the lizards' integument, that is, the occurrence of honeycomb-shaped microstructures. Interestingly, these structures can also cover the skin of lizards inhabiting wet habitats. We therefore tested the influence of the microstructures' main features on the habitat choice and wettability in the genus Phrynosoma. The genus Phrynosoma comprises moisture-harvesting species as well as nonspecialists. Lizards of this genus inhabit large areas of North America with diverse climatic conditions. Remarkably, the differences in the manifestation of microstructures are just as versatile as their surroundings. The phylogeny of the lizards as well as the depth of their ventral microstructures, though independent of each other, correlated with the precipitation in their respective habitat. All other morphological traits, as well as the skin's wettability itself, could not predict the habitat of Phrynosoma species. Hence, it is unlikely that the microstructure influences the wettability, at least directly. Hence, we presume an indirect influence for the following reasons: (a) As the ventral side cannot get wet by rain, but the belly could easily interact with a wet surface, the microstructure might facilitate water absorption from wet soil following precipitation. (b) We found the number of dorsal microstructures to be linked to the occurrence of silt in the habitat. In our study, we observed scales being heavily contaminated, most likely with a mixture of dead skin (after shedding) and silt. As many lizards burrow themselves or even shovel sand onto their backs, deploying the substrate might be a mechanism to increase the skin's wettability.

Climate change drives mountain butterflies towards the summits.

Climate change impacts biodiversity and is driving range shifts of species and populations across the globe. To understand the effects of climate warming on biota, long-term observations of the occurrence of species and detailed knowledge on their ecology and life-history is crucial. Mountain species particularly suffer under climate warming and often respond to environmental changes by altitudinal range shifts. We assessed long-term distribution trends of mountain butterflies across the eastern Alps and calculated species' specific annual range shifts based on field observations and species distribution models, counterbalancing the potential drawbacks of both approaches. We also compiled details on the ecology, behaviour and life-history, and the climate niche of each species assessed. We found that the highest altitudinal maxima were observed recently in the majority of cases, while the lowest altitudes of observations were recorded before 1980. Mobile and generalist species with a broad ecological amplitude tended to move uphill more than specialist and sedentary species. As main drivers we identified climatic conditions and topographic variables, such as insolation and solar irradiation. This study provides important evidence for responses of high mountain taxa to rapid climate change. Our study underlines the advantage of combining historical surveys and museum collection data with cutting-edge analyses.

Ecophysiological models for global invaders: Is Europe a big playground for the African clawed frog?

One principle threat prompting the worldwide decline of amphibians is the introduction of nonindigenous amphibians. The African Clawed Frog, Xenopus laevis, is now one of the widest distributed amphibians occurring on four continents with ongoing range expansion including large parts of Europe. Species distribution models (SDMs) are essential tools to predict the invasive risk of these species. Previous efforts have focused on correlative approaches but these can be vulnerable to extrapolation errors when projecting species' distributions in nonnative ranges. Recent developments emphasise more robust process-based models, which use physiological data like critical thermal limits and performance, or hybrid models using both approaches. Previous correlative SDMs predict different patterns in the potential future distribution of X. laevis in Europe, but it is likely that these models do not assess its full invasive potential. Based on physiological performance trials, we calculate size and temperature-dependent response surfaces, which are scaled to geographic performance layers matching the critical thermal limits. We then use these ecophysiological performance layers in a standard correlative SDM framework to predict the potential distribution in southern Africa and Europe. Physiological performance traits (standard metabolic rate and endurance time of adult frogs) are the main drivers for the predicted distribution, while the locomotor performance (maximum velocity and distance moved in 200 ms) of adults and tadpoles have low contributions.

The thermal ecology and physiology of reptiles and amphibians: A user's guide.

Research on the thermal ecology and physiology of free-living organisms is accelerating as scientists and managers recognize the urgency of the global biodiversity crisis brought on by climate change. As ectotherms, temperature fundamentally affects most aspects of the lives of amphibians and reptiles, making them excellent models for studying how animals are impacted by changing temperatures. As research on this group of organisms accelerates, it is essential to maintain consistent and optimal methodology so that results can be compared across groups and over time. This review addresses the utility of reptiles and amphibians as model organisms for thermal studies by reviewing the best practices for research on their thermal ecology and physiology, and by highlighting key studies that have advanced the field with new and improved methods. We end by presenting several areas where reptiles and amphibians show great promise for further advancing our understanding of how temperature relations between organisms and their environments are impacted by global climate change.

Snakes of the Pernambuco Endemism Center, Brazil: diversity, natural history and conservation.

The Atlantic Forest is one of the largest and richest tropical rainforests on the planet, being one of the 25 world priorities for conservation. The Atlantic Forest portion located north of the São Francisco River corresponds to the Pernambuco Endemism Center (PEC). We describe the snake composition of the PEC, providing information about the diversity, natural history and geographical distribution of the species, based on records from five scientific collections and additional information from the literature. A total of 78 species of snakes distributed in eight families was registered in the Pernambuco Endemism Center. The Caatinga is the Brazilian biome that most shares species with the PEC, followed by Cerrado. On the other hand, seven species are considered endemic of this region. Most of the snake species in the PEC have been registered in forest (94.8%), followed by "Brejos Nordestinos" (46.1%), Tabuleiros (43.5%), Restingas (14.1%) and Mangroves (5.1%). The PEC snake fauna includes mainly terrestrial species (60.2%) and cryptozoic and/or fossorial species (21.7%), but also presents a high richness of semi-arboreal and arboreal species (29.5%). Vertebrates are the main food item consumed by the species (78% of species), among the main prey are mammals, lizards, and amphibians. Most species show a strictly nocturnal activity period (50%), followed by strictly diurnal (38%). The PEC is the most degraded and least known region of the Atlantic Forest, yet it has revealed a high richness of snake species, including seven endemic species. It is emphasized that regional conservation efforts need to be intensified, because few forests in the region are formally protected, and the majority consist of small and poorly protected fragments, which means that many species in the region may be in risk of extinction.

Continuous expansion of the geographic range linked to realized niche expansion in the invasive Mourning gecko Lepidodactylus lugubris (Duméril & Bibron, 1836).

Lepidodactylus lugubris is a parthenogenetic gecko which has been increasingly expanding its range during the last century. This invasive species has been reported from multiple tropical and subtropical countries in five continents, most of which were colonized in recent times. In order to understand how the realized niche of the species was affected by this dramatic geographic range expansion, we reconstructed the history of the geographic range expansion. We built models of the realized niche of the species at different points in time during the invasion process. This was achieved through the implementation of modern hypervolume construction methods, based on the Hutchinson's niche concept. The models were then compared to detect possible realized climatic niche expansion over time. Furthermore, we investigated possible pathways used by the species to spread. A progressive expansion of the realized niche was identified. As the species spread into new areas, we observed a tendency to colonize regions with warmer temperatures and higher precipitation rates. Finally, we found evidence for cargo shipping being the major pathway through which the species expands its range. Further studies on this topic should aim to investigate the role of biological interactions, and how they shape the distribution of L. lugubris on a global scale. A deeper understanding of this kind of processes will help us tackle the issue of invasive species, which has become a major challenge in conservation biology.

Integrative taxonomy reveals three new taxa within the Tylototriton asperrimus complex (Caudata, Salamandridae) from Vietnam.

The Tylototriton asperrimus complex from northern Vietnam is reviewed based on morphological comparisons and analysis of the mitochondrial marker NADH dehydrogenase subunit 2 (ND2). Based on molecular divergences, which were revealed to be higher than in other congeners, in concert with morphological differences, two new species and one subspecies are described herein: Tylototriton pasmansi sp. nov. differs from T. asperrimus sensu stricto by 3.2 to 3.6 % genetic divergence and a combination of distinct morphological characters, such as head slightly longer than wide, distinct mid-dorsal ridge, relatively wide distance between the eyes, tips of fingers reaching the eye when foreleg is laid forward, labial and gular folds present, central belly skin with tubercles shaped like transverse wrinkles and distinct, pointy to round rib nodules. The population of T. pasmansi sp. nov. consists of two subclades, the nominotypic one occurring on the eastern side of the Da River (or Black River, including Hoa Binh and Phu Tho provinces), and another occurring on the western side (including Son La and Thanh Hoa provinces). These two subclades differ by 2.5 to 3.1 % genetic divergence and distinct morphological characters. The western subclade is herein described as Tylototriton pasmansi obsti ssp. nov., which differs from the nominotypic form by a wider head, longer and narrower snout, shorter femur length, and an overall less granulose skin, without an increased concentration of warts on the body sides. A second new species, Tylototriton sparreboomi sp. nov. is described from Lai Chau Province. It differs from T. asperrimus sensu stricto by 4.1 to 4.2 % and from Tylototriton pasmansi sp. nov. by 3.6 to 4.5 % genetic divergences as well as by a combination of distinct morphological characters, such as head longer than wide, tips of fingers reaching nostril when foreleg adpressed along head, rib nodules distinct, round and relatively enlarged, and wide distance between the eyes.

Differential effects of habitat loss on occupancy patterns of the eastern green lizard Lacerta viridis at the core and periphery of its distribution range.

The effects of habitat loss on the distribution of populations are often linked with species specialization degree. Specialist species can be more affected by changes in landscape structure and local patch characteristics compared to generalist species. Moreover, the spatial scale at which different land covers (eg. habitat, cropland, urban areas) affect specialist species can be smaller. Specialization is usually assumed as a constant trait along the distribution range of species. However, for several taxa, there is evidence of higher specialization degree in peripheral populations compared with populations in the core. Hence, peripheral populations should have a higher sensitivity to habitat loss, and strongest effects should be found at a smaller spatial scale. To test these expectations, we implemented a patch-landscape approach at different spatial scales, and compared effects of landscape structure and patch characteristics on occupancy probability among northern peripheral, more specialized populations (Czech Republic) and core populations (Bulgaria) of the eastern green lizard Lacerta viridis. We found that landscape structure and patch characteristics affect differently the occupancy probability of Lacerta viridis in each region. Strongest effects of habitat loss were found at a spatial scale of 150m around patches in the periphery, but at a scale of 500m in the core. In the periphery occupancy probability of populations was principally affected by landscape composition, and the effect of habitat quality was stronger compared to core populations. In the core, persistence of populations was mainly explained by characteristics of the spatial configuration of habitat patches. We discuss possible ecological mechanisms behind the relationship between sensitivity to habitat loss, populations' specialization degree and position in the distribution range, and suggest conservation measures for L. viridis.

Realized niche and microhabitat selection of the eastern green lizard (Lacerta viridis) at the core and periphery of its distribution range.

The available range of habitats and suitable abiotic conditions like temperature and radiation tends to be narrower toward the periphery of the distribution range of species. Peripheral populations of generalist species could then be more specialized and have a smaller and differentiated realized niche (habitat niche in our study) compared to populations at the core. Likewise, patterns of microhabitat selection can differ between periphery and core. In our study, we compared niche size and microhabitat selection among core (Bulgaria) and northern peripheral (Germany, Czech Republic) populations of Lacerta viridis and estimated niche differentiation among regions. We collected data on vegetation structure and abiotic parameters at the microhabitat scale in each region. In order to compare niche size among regions and estimate niche differentiation, we built multidimensional niche hypervolumes. We applied generalized linear mixed models and model averaging, accounting for spatial autocorrelation when necessary, to analyze microhabitat differences among regions and microhabitat selection in each region. Peripheral populations were more specialized, having a smaller niche than core ones, and their niche differed from that in the core (Sørensen overlap in all comparisons <0.3). Microhabitats at the periphery had lower radiation and soil compaction and less structured vegetation. Microhabitat selection at the core depended solely on abiotic parameters, while at the periphery it was defined by only vegetation structure (Czech Republic) or a combination of both, vegetation structure, and abiotic factors (Germany). Thus, peripheral populations seem to compensate for overall harsher climatic conditions by responding to different parameters of the microhabitat compared to core populations. We suggest specific conservation measures for L. virids in each studied region and point out the general implications of a higher specialization degree of peripheral populations in relation to climate change and habitat fragmentation.

Mark-release-recapture meets Species Distribution Models: Identifying micro-habitats of grassland butterflies in agricultural landscapes.

Habitat demands and species mobility strongly determine the occurrence of species. Sedentary species with specific habitat requirements are assumed to occur more patchy than mobile habitat generalist species, and thus suffer stronger under habitat fragmentation and habitat deterioration. In this study we measured dispersal and habitat preference of three selected butterfly species using mark-release-recapture technique. We used data on species abundance to calculate Species Distribution Models based on high-resolution aerial photographs taken using RGB / NIR cameras mounted on a UAV. We found that microhabitats for species with specific habitat requirements occur spatially restricted. In contrast, suitable habitats are more interconnected and widespread for mobile habitat generalists. Our models indicate that even managed grassland sites have comparatively little habitat quality, while road verges provide high quality micro-habitats. In addition, dispersal was more restricted for specialist butterfly species, and higher for the two other butterfly species with less ecological specialisation. This study shows synergies arising when combining ecological data with high precision aerial pictures and Species Distribution Models, to identify micro-habitats for butterflies. This approach might be suitable to identify and conserve high quality habitats, and to improve nature conservation at the ground.

Patterns of species richness and the center of diversity in modern Indo-Pacific larger foraminifera.

Symbiont-bearing Larger Benthic Foraminifera (LBF) are ubiquitous components of shallow tropical and subtropical environments and contribute substantially to carbonaceous reef and shelf sediments. Climate change is dramatically affecting carbonate producing organisms and threatens the diversity and structural integrity of coral reef ecosystems. Recent invertebrate and vertebrate surveys have identified the Coral Triangle as the planet's richest center of marine life delineating the region as a top priority for conservation. We compiled and analyzed extensive occurrence records for 68 validly recognized species of LBF from the Indian and Pacific Ocean, established individual range maps and applied Minimum Convex Polygon (MCP) and Species Distribution Model (SDM) methodologies to create the first ocean-wide species richness maps. SDM output was further used for visualizing latitudinal and longitudinal diversity gradients. Our findings provide strong support for assigning the tropical Central Indo-Pacific as the world's species-richest marine region with the Central Philippines emerging as the bullseye of LBF diversity. Sea surface temperature and nutrient content were identified as the most influential environmental constraints exerting control over the distribution of LBF. Our findings contribute to the completion of worldwide research on tropical marine biodiversity patterns and the identification of targeting centers for conservation efforts.

The ontogeny of developmental buffering in lizard head shape.

Canalization and developmental stability (DS) are important organismal properties involved in determining the level of phenotypic variation. Ontogenetic patterns of phenotypic variance components can shed light on the mechanistic basis of developmental buffering (DB). Here, we analyze how individual FA and among-individual variation in head shape change in ontogenetic series of three lizard species raised in laboratory. The degree of asymmetry increased slightly with size, suggesting that developmental mechanisms hypothesized to correct for deviations either do not exist, or that their efficiency is truncated with increasing size. Alternatively, they may need the disturbance as a trigger. The relationship between asymmetry and age was complex, with asymmetry being stable across the age range in two species but increased with age in the third. Lack of congruence in ontogenetic patterns of asymmetry might be due to intrinsic differences in buffering mechanisms or a result of species-specific growth patterns. Head shape was shown to be equally canalized across both size and age range in all species, probably as a result of a balance between the buffering mechanisms and mechanisms generating variance. The patterns of symmetric and asymmetric head shape variation were highly correlated across species meaning that DS and canalization may rely on similar mechanisms.

Snails in the desert: Species diversification of Theba (Gastropoda: Helicidae) along the Atlantic coast of NW Africa.

The spatial subdivision of species often plays a pivotal role in speciation. Across their entire range, species are rarely panmictic and crucial consequences of spatial subdivision are (1) random genetic drift including historical factors, (2) uniform selection, and (3) divergent selection. Each of these consequences may result in geographic variation and eventually reproductive isolation, but their relative importance in speciation is still unclear. In this study, we used a combination of genetic, morphological, and climatic data to obtain a comprehensive picture of differentiation among three closely related, parapatrically distributed taxa of the land snail genus Theba occurring along the Atlantic coasts of South Morocco and Western Sahara. We conducted Mantel and partial Mantel tests to relate phenotypic and genotypic variation of these species to geography and/or climate. As null hypothesis for an evolutionary scenario, we assumed nonadaptive speciation and expected a pattern of isolation by distance among taxa. Rejection of the null hypothesis would indicate isolation by environment due to adaptation. Generally, genetic drift plays an important role but is rarely considered as sole driver of speciation. It is the combination of drift and selection that predominantly drives speciation. This study, however, provides a potential example, in which nonadaptive speciation, that is, genetic drift, is apparently the main driver of shaping the diversity of Theba in NW Africa. Restriction of gene flow between populations caused by geographic isolation probably has played an important role. Climate oscillations during the Plio- and Pleistocene may have led to repeated ecological changes in NW Africa and disruptions of habitats promoting differentiation by geographic isolation. The inferred evolutionary scenario, however, did not fully explain the incongruence between the AFLP- and mtDNA-tree topologies. This incongruence might indicate past hybridization among the studied Theba forms.