Challenges and opportunities for biogeography—What can we still learn from von Humboldt?

Alexander von Humboldt was arguably the most influential scientist of his day. Although his fame has since lessened relative to some of his contemporaries, we argue that his influence remains strong—mainly because his approach to science inspired others and was instrumental in furthering other scientific disciplines (such as evolution, through Darwin, and conservation science, through Muir)—and that he changed the way that large areas of science are done and communicated. Indeed, he has been called the father of a range of fields, including environmental science, earth system science, plant geography, ecology and conservation. His approach was characterized by making connections between non‐living and living nature (including humans), based on interdisciplinary thinking and informed by large amounts of data from systematic, accurate measurements in a geographical framework. Although his approach largely lacked an evolutionary perspective, he was fundamental to creating the circumstances for Darwin and Wallace to advance evolutionary science. He devoted considerable effort illustrating, communicating and popularizing science, centred on the excitement of pure science. In biogeography, his influence remains strong, including in relating climate to species distributions (e.g. biomes and latitudinal and elevational gradients) and in the use of remote sensing and species distribution modelling in macroecology. However, some key aspects of his approach have faded, particularly as science fragmented into specific disciplines and became more reductionist. We argue that asking questions in a more Humboldtian way is important for addressing current global challenges. This is well‐exemplified by researching links between geodiversity and biodiversity. Progress on this can be made by (a) systematic data collection to improve our knowledge of biodiversity and geodiversity around the world; (b) improving our understanding of the linkages between biodiversity and geodiversity; and (c) developing our understanding of the interactions of geological, biological, ecological, environmental and evolutionary processes in biogeography.     

Speciation in the highlands of Mexico: genetic and phenotypic divergence in the Mexican jay (Aphelocoma ultramarina)

The pine-oak woodlands of the Mexican highlands harbour significant biological diversity, yet little is known about the evolutionary history of organisms inhabiting this region. We assessed genetic and phenotypic differentiation in 482 individuals representing 27 populations of the Mexican jay ( Aphelocoma ultramarina ) — a widespread bird species of the Mexican highlands — to test whether populations in the central and northern Mexican sierras display discrete breaks between groups, which would be consistent with a role for the different mountain chains in divergence and speciation. We found abrupt breaks in mitochondrial DNA (mtDNA; ND2 and control region) delineating four major genetic groups found in the Sierra Madre Occidental, Sierra Madre Oriental, southern Central Plateau (Bajio), and Transvolcanic Belt. These mtDNA groups were largely corroborated by data from nuclear microsatellites and phenotypic data, except that clades from the Central Plateau and Sierra Madre Oriental showed clinal change in these data sets. Uncertainty about the mutation rate for our mitochondrial markers warrants considerable caution with regard to estimating divergence times, but the major genetic groups appear to have split before the most extreme period of glacial cycling that marked the last 0.7 million years and after Mexico's period of major mountain formation. The fact that some genetic breaks do not coincide with well-known geographic barriers suggests a role for ecology in divergence and speciation, and we discuss implications for taxonomy and conservation.     

Darwin’s Galapagos gourd: providing new insights 175 years after his visit

The year 2010 marks the 175th anniversary of Charles Darwin’s visit to the Galapagos Islands. A recent paper by J. C. Briggs, ‘Darwin’s biogeography’ (Journal of Biogeography, 2009, 36 , 1011–1017), summarizes Darwin’s contributions to the field of biogeography, stressing the importance of his natural history specimens. Here, we illustrate how a plant collected by Darwin during his visit to Floreana and not collected since can provide insights into dispersal to oceanic islands as well as extinction of island plants, based on ancient DNA from Darwin’s herbarium specimen.     

Phylogeography and molecular systematics of the Peromyscus aztecus species group (Rodentia: Muridae) inferred using parsimony and likelihood

Mice of the Peromyscus aztecus species group occur at mid to high elevations in several mountain ranges in the highlands of Middle America (Mexico and Central America), a region of high endemicity. We examined the biogeography of this group by conducting phylogenetic analyses of 668 bp of the mitochondrial cytochrome b (cyt b) gene. Phylogenetic analyses under both parsimony and likelihood frameworks produced the same topologies, but estimates of nodal support were artificially high in weighted parsimony analyses. This difference is attributed to the inability of parsimony to optimize branch lengths when evaluating topologies. These data indicate that the P. aztecus-like populations from south and east of the Isthmus of Tehuantepec currently assigned to P. a. oaxacensis represent a distinct species, with genetic distances as high as 0.091. In addition, P. hylocetes is strongly divergent from Mexican populations of P. aztecus (genetic distances of 0.044-0.069), supporting the recognition of this taxon as a distinct species. The history of divergence in this group can be explained by a series of apparently early to middle Pleistocene vicariance events associated with glacial cycles. The Sierra Madre Occidental and Cordillera Transvolcanica each appear to be faunistically isolated, the Isthmus of Tehuantepec appears to have been a strong Pleistocene barrier, and the Sierra Madre Oriental has affinities with the Sierra Madre del Sur and the highlands of central Oaxaca.     

Halffter's Mexican transition zone (1962–2014), cenocrons and evolutionary biogeography

The Mexican transition zone is the complex and varied area in which the Neotropical and Nearctic biotas overlap. In a series of contributions, Gonzalo Halffter provided a coherent theory that explains how sets of taxa that evolved in different geographical areas assembled in this transition zone. Halffter's theory developed gradually, being refined and clarified in successive contributions from him and other authors. After a review of the historical development of the Mexican transition zone, including the characterization of the dispersal or distributional patterns recognized by Halffter, its relevance for evolutionary biogeography is discussed briefly. The Mexican transition zone in the strict sense includes the highlands of Mexico and Guatemala (Sierra Madre Occidental, Sierra Madre Oriental, Transmexican Volcanic Belt, Sierra Madre del Sur and Chiapas Highlands provinces), whereas northern Mexico and the southern United States are clearly Nearctic, and the lowlands of southern Mexico and Central America are clearly Neotropical. The distributional patterns recognized by Halffter are considered to represent cenocrons (sets of taxa that share the same biogeographical history, constituting identifiable subsets within a biota by their common biotic origin and evolutionary history). The development of the Mexican transition zone is summarized into the following stages: (1) Jurassic–Cretaceous: the four Paleoamerican cenocrons extend in Mexico; (2) Late Cretaceous–Palaeocene: dispersal from South America of the Plateau cenocron; (3) Oligocene–Miocene: dispersal from the Central American Nucleus of the Mountain Mesoamerican cenocron; (4) Miocene–Pliocene: dispersal from North America of the Nearctic cenocron; and (5) Pleistocene: dispersal from South America of the Typical Neotropical cenocron.     

Multilocus phylogeography of the flightless darkling beetle Nyctoporis carinata (Coleoptera: Tenebrionidae) in the California Floristic Province: deciphering an evolutionary mosaic

The California Floristic Province (CFP) is considered a global biodiversity hotspot because of its confluence of high species diversity across a wide range of threatened habitats. To understand how biodiversity hotspots such as the CFP maintain and generate diversity, we conducted a phylogeographic analysis of the flightless darkling beetle, Nyctoporis carinata, using multiple genetic markers. Analyses of both nuclear and mitochondrial loci revealed an east–west genetic break through the Transverse Ranges and high genetic diversity and isolation of the southern Sierra Nevada Mountains. Overall, the results obtained suggest that this species has a deep evolutionary history whose current distribution resulted from migration out of a glacial refugium in the southern Sierra Nevada via the Transverse Ranges. This finding is discussed in light of similar genetic patterns found in other taxa to develop a foundation for understanding the biodiversity patterns of this dynamic area. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 424–444.     

Long‐distance movements and evidence of post‐breeding elevational movements by Cassin’s Vireos

Bird migration is often framed as a straightforward journey between one breeding site and one wintering site, but recent research has shown that the reality is often more complex. Many species of birds undertake short‐distance movements separate from long‐distance migration. Such movements appear to be common in species that breed in western North America, where mountainous terrain creates a mosaic of habitats and climatic conditions. However, individual‐based tracking studies have disproportionately focused elsewhere, leaving gaps in our understanding of the year‐round movements of western species. I used tracking data from light‐level geolocators and citizen science data from eBird to study the movements of Cassin’s Vireos (Vireo cassinii) breeding in the Sierra Nevada mountains of California, USA. During three breeding seasons (2013–2015), my observations suggested that Cassin’s Vireos vacate their breeding territories during the post‐breeding period in July and August. In April and May 2016, I tagged 22 Cassin’s Vireos with light‐level geolocators and, in April and May 2017, recaptured four that had retained their geolocators. Geolocator data showed that these four birds remained in the same geographic region as their breeding territories (likely the same mountain range) during the post‐breeding period in July and August 2016, ruling out the possibility of long‐distance movements during this time. Analysis of eBird citizen science data suggested that Cassin’s Vireos undertake short‐distance molt‐migration to higher elevations in the Sierra Nevada Mountains during the post‐breeding period. Geolocator data revealed that long‐distance fall migration took place in September and spring migration in April or May, and the four birds spent the winter in different parts of the Mexican winter range of Cassin’s Vireos. These results add to the body of literature on the complex movements of migratory songbirds breeding in the mountains of western North America, an understanding that will be important for effective conservation in the future.     

The Life and Work of Marquis Robert de Wavrin,an Early Visual Anthropologist

This article traces the life and work of Marquis Robert de Wavrin de Villers au Tertre (1888–1971), a Belgian explorer and ethnographer. While fragments of his oeuvre are familiar to scholars of South America, he is almost completely unknown in historical studies, and largely forgotten within anthropology too. Here we will explore his filmic work as well as its contribution to the history of visual anthropology. While de Wavrin’s work cannot be divorced from the discipline’s colonial and Eurocentric heritage, we show that his visual record provides notable historic insights and merits further scholarly attention.     

The history of the Purine Club: a tribute to Prof. Geoffrey Burnstock

The international purinergic scientific community has lost its pioneer. Geoffrey Burnstock, born on the 10th of May 1929 in London, died on the 2nd of June 2020, aged 91, in Melbourne (Australia). Geoff was one of the most highly regarded scientists of his generation. In the 1960s and 1970s, he developed a radical and somehow heretical new theory and opened an entire new field of science, signalling via extracellular nucleotides (the “purinergic theory”), which revolutionized our understanding of how cells communicate between each other. Initially, his unconventional theory found a lot of resistance in the scientific community. Once, one scientist even threatened to devote his entire life to disproving Burnstock’s theory. Undeterred, Geoff went further on, and continued to accumulate evidence in favour of his hypothesis, and led the field ever since. He struggled to attract new scientists to this new field of research and, in the early 1990s, due to new molecular biology techniques making it possible to isolate and identify cell surface receptors for ATP and its breakdown product adenosine, did evidence emerge that eventually convinced the doubters. The number of spontaneous obituaries and messages honouring Geoff’s memory that have appeared on specialized Journals and in the public press throughout the world since last June indicates that many people are clearly affected by his death. Besides being a rigorous, ethical and extremely brilliant scientist, Geoff was an extraordinary human being, always eager to collaborate and share data, never jealous of his findings and capable of learning things even from young people. He was known for his enthusiasm, empathy and ability to motivate young scientists and promote their careers. After the establishment of the Purine Club back in the 1990s, numerous Purine Club Chapters have been formed around the world with Geoff’s help and encouragement. He has obviously also been the inspirator and founder of our Journal, Purinergic Signalling (PUSI). For this reason, Charles Kennedy, the current Editor of the Journal, and myself thought that it might be nice to invite representatives from all known Purine Clubs to send a few notes to be published in PUSI on the history of their club and how Geoff inspired, aided or supported them. Here, I have collected all their contributions and I share with the entire purinergic community my personal memories on how the Purine Club was born and developed thanks to the invaluable mentoring of Geoffrey Burnstock. I apologize in advance if I am missing some information or forgot to mention somebody, and I strongly encourage all readers to submit memories and additional information that I shall gather for future writing. Keeping alive the history of how the field developed will be the best tribute that we can play to celebrate Geoff’s work along the years.

     

Deriving artificial models of visitors from dispersed patterns of use in the Sierra Nevada Wilderness, California

Natural resource managers are faced with a complex problem of understanding human use patterns and associated impacts in dispersed recreation wilderness settings. While conventional approaches to modelling have limited use in acquiring and understanding such complex associations, spatial simulation models have been proposed as an alternative. The purpose of this paper is to describe a project whose focus is on a dispersed recreation context of backpacking trips and commercial packstock operations in the John Muir Wilderness in the Sierra Nevada Mountains in California. This paper will discuss the data collection and synthesis to derive agent profiles and rules as a precursor to the development of a dynamic, agent based model that represents the spatial distribution of visitation patterns.     

Inferring Geographic Isolation of Wolverines in California Using Historical DNA

ABSTRACT Delineating a species' geographic range using the spatial distribution of museum specimens or even contemporary detection-non-detection data can be difficult. This is particularly true at the periphery of a species range where species' distributions are often disjunct. Wolverines (Gulo gulo) are wide-ranging mammals with discontinuous and potentially isolated populations at the periphery of their range. One potentially disjunct population occurred in the Sierra Nevada Mountains, California, USA, and appears to have been extirpated by the 1930s. Many early 20th century naturalists believed that this population was connected to other populations occurring in the Cascade Range of northern California, Oregon, and Washington, USA, but a recent analysis of historical records suggests that California wolverines were isolated from other populations in North America. We used DNA extracted from museum specimens to examine whether California wolverines were isolated. Both nuclear and mitochondrial DNA data indicate that California wolverines were genetically distinct from extant populations, suggesting long-term isolation. We identified 2 new control region (mitochondrial DNA) haplotypes located only within California. We used these data and referenced sequences from the Rocky Mountains, USA, to make inferences regarding potential wolverine translocations into California. In addition, we used these genetic data to make inferences about wolverine conservation throughout western North America.     

Panbiogeography: Its origin,metamorphosis and decline

From the viewpoint of 2007, one can trace the history of an interesting and contentious trend in biogeography and evolution that began with Croizat’s concept of panbiogeography in 1958. After a quiescent period of about 16 years, some young biologists in New York and in New Zealand read Croizat’s books and became enthusiastic supporters of his ideas. In New York, in the early 1970s, panbiogeography was combined with a part of Hennig’s phylogenetic method to create vicariance biogeography. In 1986, the name of the latter was changed to cladistic biogeography. In the meantime, Croizat’s followers in New Zealand sought to maintain panbiogeography in its original form without reference to phylogeny. This idea reached its peak of popularity in 1989–1990 and then began to fade. In comparison, cladistic biogeography became much more widespread, especially when its followers began publishing laudatory books and papers. Its decline became noticeable after the turn of the century as the dispersal counterrevolution began to have its effect. It served a useful purpose by engaging the interest of young biologists who otherwise may not have become aware of biogeography.     

A critique of 'countryside biogeography' as a guide to research in human‐dominated landscapes

Proliferation of redundant terms in ecology and conservation slows progress and creates confusion. ‘Countryside biogeography’ has been promoted as a new framework for conservation in production landscapes, so may offer a replacement for other concepts used by landscape ecologists. We conducted a systematic review to assess whether the 'countryside biogeography' concept provides a distinctive framing for conservation in human‐dominated landscapes relative to existing concepts. We reviewed 147 papers referring to countryside biogeography and 81 papers that did not. These papers were divided into categories representing three levels of use of countryside biogeography concepts (strong, weak, cited only) and two categories that did not use countryside biogeography at all but used similar concepts including fragmentation and matrix. We revealed few distinctions among groups of papers. Countryside biogeography papers made more frequent use of the terms 'ecosystem services', 'intensification' and 'land sparing' compared with non‐countryside biogeography papers. Papers that did not refer to countryside biogeography sampled production areas (e.g. farms) less often, and this related to their focus on habitat specialist species for which patch‐matrix assumptions were reasonable. Countryside biogeography offers a conceptual wrapper rather than a distinctive framework for advancing research in human‐modified landscapes. This and similar wrappers such as ‘conservation biogeography’ and ‘agricultural biogeography’ risk creating confusion among new researchers, and can prevent clear communication about research. To improve communication, we recommend using the suite of well‐established terms already applied to conservation in human‐modified landscapes rather than through an interceding conceptual wrapper.     

Species formation and geographical range evolution in a genus of Central American cloud forest salamanders (Dendrotriton)

Aim Montane Central America offers an ideal system for testing geographical hypotheses of species diversification. We examined how the complex geological history of Nuclear Central America has shaped the diversification of a genus of cloud‐forest‐inhabiting salamanders (Dendrotriton). We applied parametric models of geographical range evolution to determine the predominant mode of species formation within the genus and to test existing hypotheses of geographical species formation in the region. Location Montane cloud forests of Nuclear Central America. Methods We estimated a species tree for Dendrotriton using a multi‐locus DNA sequence data set and several coalescent methods, and performed molecular dating for divergence events within the genus. We then applied the species‐tree estimate to a likelihood‐based time‐stratified model of geographical range evolution, based on current species distributions and available geological information for Central America. Results Species trees from all methods contain two groups, one corresponding to species from the Sierra de los Cuchumatanes and the other containing all remaining species. In most cases, species formation within the genus involved an even division of the geographical range of the ancestral species between descendant species. The ancestor of extant Dendrotriton species was estimated to have occurred in either the Sierra de los Cuchumatanes or the Sierra Madre de Chiapas, and both of these areas appear to have been important for diversification within the genus. The single species found in the Quaternary‐age Guatemalan volcanic cordillera dispersed to the volcanoes from an older highland area. Main conclusions Models of geographical range evolution, when combined with robust species‐tree estimates, provide insight into the historical biogeography of taxa not available from phylogenies or distributional data alone. Vicariant species formation, rather than peripatric or gradient speciation, appears to have been the dominant process of diversification, with most divergence events occurring within or between ancient highland areas. The apparent dispersal of Dendrotriton to the Quaternary‐age volcanoes raises the possibility that the rich salamander community there is composed of species that dispersed from geologically older areas. The Motagua Valley appears not to have been as important in vicariant species formation within Dendrotriton as it is within other groups.     

Shifts in fisher (Pekania pennanti) diet in response to climate-induced tree mortality in California assessed with DNA metabarcoding

A recent climate-induced tree mortality event in California, USA has led to dramatic landscape-level changes in the southern Sierra Nevada. Wide-spread conifer mortality was documented in habitat occupied by fisher (Pekania pennanti), a mature-forest associated species of conservation concern in this region. We analyzed fisher scats collected on the Sierra National Forest from the pre-tree-mortality period (PreTM, 2011 – 2013) and post-tree-mortality period (PostTM, 2017 – 2018). We used DNA metabarcoding to successfully identify taxa and summarized diet composition for 109 PreTM and 102 PostTM fisher scats. We observed 48 different diet items (33 assigned to species and 15 to genus). Mammals, birds, and plants of the Ribes genus (gooseberries and currents) comprised the highest proportions of diet items, although scats also contained DNA from reptiles, insects, arachnids, snails, and fungi. The frequency of occurrence of mammalian prey items was lower in PostTM (49.0%) versus PreTM scats (81.7%) with a reduction in the occurrence of two tree squirrels (Douglas squirrel and Humboldt’s flying squirrel). A higher proportion of scats collected PostTM (46.1%) contained DNA from Ribes spp. versus scats collected PreTM (19.3%). Our data reveal potential cascading effects of climate-induced tree mortality on fisher diet in the southern Sierra Nevada. Flexibility in fisher diet, however, may allow resilience to ecological change though future studies should consider the behavioral, energetic, demographic or fitness consequences from a shift away from medium-sized mammalian prey to plants.     

Historical demography and phylogeography of a specialist bark beetle, Dendroctonus pseudotsugae Hopkins (Curculionidae: Scolytinae)

Contemporary distribution of North American species has been shaped by past glaciation events during the Quaternary period. However, their effects were not as severe in the southern Rocky Mountains and Northern Mexico as elsewhere in North America. In this context, we test hypotheses about the historical demography of Dendroctonus pseudotsugae, based on 136 haplotypes of mitochondrial cytochrome oxidase I. The phylogenetic analysis yielded four haplogroups corresponding to northwestern United States and southwestern Canada (NUS), southwestern United States (Arizona, SUS), northwestern Mexico (Sierra Madre Occidental, SMOC), and northeastern Mexico (Sierra Madre Oriental, SMOR). Predictions of demographic expansion were examined through neutrality tests against population growth and mismatch distribution. Results showed that the NUS and SMOC haplogroups have experienced demographic expansion events, whereas the SUS and SMOR haplogroups have not. Divergence times between pairs of haplogroups were estimated from early to middle Pleistocene. The longer divergence time between NUS and all other haplogroups could be the result of refugia within the Pacific Northwest and northern Rocky Mountains and long-term isolation from southernmost populations in Mexico. The results obtained in this study are in agreement with the evolutionary history of the host Douglas-fir, as the warmer climates of interglacial periods pushed conifers northward of Colorado, New Mexico, and Arizona, whereas environmental changes reduced the population size of Douglas-fir and forced fragmentation of distribution range southward into northern Mexico.     

Life in proto‐Amazonia: Middle Miocene mammals from the Fitzcarrald Arch (Peruvian Amazonia)

The Middle Miocene has been identified as a time of great diversification in modern lineages now distributed in tropical South America, and when basic archetypal traits defining Amazonia appear, including climatic humid conditions, basic floral physiognomy and phylogenetic composition of modern rainforests. Nonetheless, Middle Miocene localities in South America are poorly known, especially at low latitudes where only one species‐rich locality, La Venta in Colombia, has been extensively studied. The present contribution describes the mammal fauna of Fitzcarrald, a new Middle Miocene local fauna from western Amazonia in Peru. Fitzcarrald is correlated with the Laventan South American Land Mammal Age based on the presence of taxa defining the ‘Miocochilius assemblage zone’ in La Venta. The mammalian fauna of Fitzcarrald comprises 24 taxa among cingulates, folivores, astrapotheres, notoungulates, litopterns, rodents, odontocetes and a possible marsupial. At this time, tropical South America was characterized by the presence of the Pebas megawetland, a huge lacustrine complex that provided unique ecological and environmental conditions most likely isolating northern South America from southern South America. These isolating conditions might have come to an end with its disappearance in the Late Miocene and the establishment of the subsequent Acre system, the predecessor fluvial system of modern Amazonia. Results of faunistic similarity between Fitzcarrald and other Miocene faunas throughout South America support these scenarios. The Fitzcarrald mammal fauna exhibits first appearance datums and last appearance datums of various taxa, showing that tropical South America has played a crucial role in the evolutionary history and biogeography of major clades, and revealing a more complex biological history than previously proposed, based on the record from the southern cone of the continent.     

Long‐term demographic decline and late glacial divergence in a Californian paleoendemic: Sequoiadendron giganteum (giant sequoia)

Mediterranean ecosystems comprise a high proportion of endemic taxa whose response to climate change will depend on their evolutionary origins. In the California flora, relatively little attention has been given to the evolutionary history of paleoendemics from a molecular perspective, yet they number among some of the world's most iconic plant species. Here, we address questions of demographic change in Sequoiadendron giganteum (giant sequoia) that is restricted to a narrow belt of groves in the Sierra Nevada Mountains. We ask whether the current distribution is a result of northward colonization since the last glacial maximum (LGM), restriction of a broader range in the recent past (LGM) or independent colonizations in the deeper past. Genetic diversity at eleven microsatellite loci decreased with increasing latitude, but partial regressions suggested this was a function of smaller population sizes in the north. Disjunct populations north of the Kings River were divergent from those south of the Kings River that formed a single cluster in Bayesian assignment tests. Demographic inferences supported a demographic contraction just prior to the LGM as the most likely scenario for the current disjunct range of the species. This contraction appeared to be superimposed upon a long‐term decline in giant sequoia over the last 2 million years, associated with increasing aridity due to the Mediterranean climate. Overall, low genetic diversity, together with competition in an environment to which giant sequoia is likely already poorly adapted, will pose major constraints on its success in the face of increasing aridity.     

Phylogeography of the Northern Alligator Lizard (Squamata,Anguidae): Hidden diversity in a western endemic

Western North America includes the California Floristic Province and the Pacific Northwest, biologically diverse regions highlighted by a complex topography, geology, climate and history. A number of animals span these regions and show distinctive patterns of dispersal, vicariance and lineage diversification. Examining phylogeographic patterns in the fauna of this area aids in our understanding of the forces that have contributed to the generation and maintenance of regional biodiversity. Here, we investigate the biogeography and population structure of the Northern Alligator Lizard (Elgaria coerulea), a wide‐ranging anguid endemic to western North America. We sequenced two mtDNA fragments (ND2 and ND4) for 181 individuals across the range of the species and analysed these data with phylogenetic approaches to infer population and biogeographic history, and date major divergences within the taxon. We further used Bayesian clustering methods to assess major patterns of population structure and performed ecological niche modelling (ENM) to aid in our interpretation of geographic structure and diversification of E. coerulea lineages. Our phylogeographic examination of E. coerulea uncovered surprising diversity and structure, recovering 10 major lineages, each with substantial geographic substructure. While some divergences within the species are relatively old (Pliocene, 5.3–2.6 mya), most intraspecific variation appears to be of more recent origin (Pleistocene, 2.6 mya‐11,700 ya). Current diversity appears to have arisen in the Sierra Nevada Mountains and spread west and north since the Pliocene. Finally, our ENMs suggest that much of the Coast Ranges in California provided ideal habitat during the Last Glacial Maxima (LGM) that has since contracted dramatically and shifted northwards, whereas significant portions of the Sierra Nevada were unsuitable during the LGM and have since become more suitable. Interestingly, E. coerulea shares a number of genetic boundaries with other sympatric taxa, suggesting common historical events and geomorphological features have shaped the biota of this region.     

A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation

As humans continue to explore the last uncharted regions of the planet, discoveries of previously unknown species of large vertebrates have become infrequent. Here, we report on the discovery of a spectacular new species of giant, secretive, frugivorous, forest monitor lizard (Genus: Varanus) from the forests of the northern Philippines. Using data from morphology and mitochondrial and nuclear DNA sequences, we demonstrate the taxonomic distinctiveness of this new 2 m long species and provide insight into its historical biogeography and systematic affinities. Our molecular phylogenetic analyses indicate that the new species is closely related to Varanus olivaceus (from southern Luzon and nearby islands), but it differs from this and other varanids with respect to characteristics of scalation, colour pattern, body size, anatomy of the reproductive organs and genetic divergence. The new species appears to be restricted to forests of the central and northern Sierra Madre mountain range; it is separated from the range of V. olivaceus by a more than 150 km stretch that includes at least three low-elevation river valley barriers to dispersal. This discovery identifies a seldom-perceived biogeographic boundary and emphasizes the need for continued biodiversity research in the megadiverse conservation hotspot of the Philippines. It is anticipated that the new species will serve as an important flagship species for conservation efforts aimed at preserving the remaining forests of northern Luzon.