Juan Carlos Rando

What are We Learning about Speciation and Extinction from the Canary Islands

Summary. Oceanic islands are excellent systems for allowing biologists to test evolutionary hypotheses due to their relative simplicity of habitats, naturally replicated study design and high levels of endemic taxa with conspicuous variation in form, colour and behaviour. Over the last two decades the Canary Islands archipelago has proved an ideal system for evolutionary biologists who seek to unravel how biodiversity arises and disappears. In this review we have evaluated the contribution of the study of Canarian birds to our understanding of how and why species occur and change over time. We focus our attention on both extant and extinct Canarian taxa, and describe how research on these species has filled gaps in our understanding of avian speciation and extinction. In addition, we discuss the necessity of revising the current taxonomy in the Canarian avian taxa, especially the status of the endemic subspecies, some of which might be better treated as full species. An accurate classification of Canarian birds is not only necessary for testing evolutionary, biogeographic and ecological hypotheses, but also for effective decision making about conservation and environmental management. Finally we introduce future avenues of research that we feel will yield the most exciting and promising findings on island evolution in the coming years.

Disentangling Ancient Interactions: A New Extinct Passerine Provides Insights on Character Displacement among Extinct and Extant Island Finches

Background Evolutionary studies of insular biotas are based mainly on extant taxa, although such biotas represent artificial subsets of original faunas because of human-caused extinctions of indigenous species augmented by introduced exotic taxa. This makes it difficult to obtain a full understanding of the history of ecological interactions between extant sympatric species. Morphological bill variation of Fringilla coelebs and F. teydea (common and blue chaffinches) has been previously studied in the North Atlantic Macaronesian archipelagos. Character displacement between both species has been argued to explain bill sizes in sympatry. However, this explanation is incomplete, as similar patterns of bill size have been recorded in F. coelebs populations from islands with and without F. teydea. Methodology/Principal Findings The discovery of a new extinct species in Tenerife (Canary Islands), here named Carduelis aurelioi n. sp. (slender-billed greenfinch), provides the opportunity to study ancient ecological interactions among Macaronesian finches. To help understand the evolutionary histories of forest granivores in space and time, we have performed a multidisciplinary study combining: (1) morphological analyses and radiocarbon dating (11,460±60 yr BP) of the new taxon and, (2) molecular divergence among the extant finch species and populations in order to infer colonization times (1.99 and 1.09 My for F. teydea and F. coelebs respectively). Conclusion/Significance C. aurelioi, F. coelebs and F. teydea co-habited in Tenerife for at least one million years. The unique anatomical trends of the new species, namely chaffinch-like beak and modified hind and forelimbs, reveal that there was a process of divergence of resource competition traits among the three sympatric finches. The results of our study, combined with the presence of more extinct greenfinches in other Macaronesian islands with significant variation in their beak sizes, suggests that the character displacement has influenced patterns of divergence in bill size and shape on other Macaronesian islands as well.

Ancient DNA of the Extinct Lava Shearwater (Puffinus olsoni) from the Canary Islands Reveals Incipient Differentiation within the P. puffinus Complex

Background The loss of species during the Holocene was, dramatically more important on islands than on continents. Seabirds from islands are very vulnerable to human-induced alterations such as habitat destruction, hunting and exotic predators. For example, in the genus Puffinus (family Procellariidae) the extinction of at least five species has been recorded during the Holocene, two of them coming from the Canary Islands. Methodology/Principal Findings We used bones of the two extinct Canary shearwaters (P. olsoni and P. holeae) to obtain genetic data, for use in providing insights into the differentiation process within the genus Puffinus. Although mitochondrial DNA (mtDNA) cytochrome b sequences were successfully retrieved from four Holocene specimens of the extinct Lava shearwater (P. olsoni) from Fuerteventura (Canary Islands), the P. holeae specimens yielded no DNA. Only one haplotype was detected in P. olsoni, suggesting a low genetic diversity within this species. Conclusions The phylogenetic analyses based on the DNA data reveal that: (i) the “Puffinus puffinus complex”, an assemblage of species defined using osteological characteristics (P. puffinus, P. olsoni, P. mauretanicus, P. yelkouan and probably P. holeae), shows unresolved phylogenetic relationships; (ii) despite the differences in body size and proportions, P. olsoni and the extant P. puffinus are sister species. Several hypotheses can be considered to explain the incipient differentiation between P. olsoni and P. puffinus.

Status, conservation and habitat selection of the Houbara Bustard Chlamydotis undulata fuertaventurae on Lanzarote (Canary Islands)

We would like to thank Keith Emmerson for his invaluable information on previous censuses. Pedro Jordano assisted during the dta analysis. Nigel J. Collar and an anonymous referee made useful comments on the manuscript. This study was supported by the Viceconsejeria de Medio Ambiente of the Canary Government.

Five new extinct species of rails (Aves: Gruiformes: Rallidae) from the Macaronesian Islands (North Atlantic Ocean).

Five new species of recently extinct rails from two Macaronesian archipelagoes (Madeira and Azores) are described. All the species are smaller in size than their presumed ancestor, the European rail Rallus aquaticus. Two species inhabited the Madeira archipelago: (1) Rallus lowei n. sp., the stouter of the species described herein, was a flightless rail with a robust tarsometatarsus and reduced wings that lived on Madeira Island; (2) Rallus adolfocaesaris n. sp., a flightless and more gracile species than its Madeiran counterpart, inhabited Porto Santo. So far, six Azorean islands have been paleontologically explored, and the remains of fossil rails have been found on all of them. Here we formally describe the best-preserved remains from three islands (Pico, São Miguel and São Jorge): (1) Rallus montivagorum n. sp., a rail smaller than R. aquaticus with a somewhat reduced flying capability, inhabited Pico; (2) Rallus carvaoensis n. sp., a small flightless rail with short and stout legs and a bill apparently more curved than in R. aquaticus, was restricted to São Miguel; (3) Rallus minutus n. sp., a very small (approaching Atlantisia rogersi in size) flightless rail with a shortened robust tarsometatarsus, lived in São Jorge. We note also the presence of rail fossils on three other Azorean islands (Terceira, Graciosa and Santa Maria). In addition, we describe an extraordinarily complete fossil of an unnamed Rallus preserved in silica from the locality of Algar do Carvão on Terceira.

Population connectivity buffers genetic diversity loss in a seabird

BackgroundAncient DNA has revolutionized conservation genetic studies as it allows monitoring of the genetic variability of species through time and predicting the impact of ecosystems’ threats on future population dynamics and viability. Meanwhile, the consequences of anthropogenic activities and climate change to island faunas, particularly seabirds, remain largely unknown. In this study, we examined temporal changes in the genetic diversity of a threatened seabird, the Cory’s shearwater (Calonectris borealis).FindingsWe analysed the mitochondrial DNA control region of ancient bone samples from the late-Holocene retrieved from the Canary archipelago (NE Atlantic) together with modern DNA sequences representative of the entire breeding range of the species. Our results show high levels of ancient genetic diversity in the Canaries comparable to that of the extant population. The temporal haplotype network further revealed rare but recurrent long-distance dispersal between ocean basins. The Bayesian demographic analyses reveal both regional and local population size expansion events, and this is in spite of the demographic decline experienced by the species over the last millennia.ConclusionsOur findings suggest that population connectivity of the species has acted as a buffer of genetic losses and illustrate the use of ancient DNA to uncover such cryptic genetic events.

Diet of the extinct Lava mouse Malpaisomys insularis from the Canary Islands: insights from dental microwear

Malpaisomys insularis is a mouse-like rodent endemic to the eastern Canary Islands. It became extinct during the fourteenth century. It was a remarkable species living under hyperarid conditions. A dental microwear analysis was performed in order to determine its former diet. The elevated number of fine scratches found in Malpaisomys molars suggests that it consumed a significant part of Poaceae, grass consumption leaving the most distinctive features on dental wear facets. A graminivorous diet with a high amount of abrasive items is in agreement with the broad teeth of Malpaisomys, considered as adapted to grass consumption. However, in the absence of potential competitors over its native range, it is likely that Malpaisomys also foraged on dicots to meet higher nutrient and energetic requirements. The ecology of Malpaisomys is discussed from these results in the context of the desertic climatic conditions of the eastern Canary Islands and with a special concern on its small body size in contrast to other large-sized island murine species such as the giant rats of the central Canary Islands.

Island extinctions: processes, patterns, and potential for ecosystem restoration

Extinctions have altered island ecosystems throughout the late Quaternary. Here, we review the main historic drivers of extinctions on islands, patterns in extinction chronologies between islands, and the potential for restoring ecosystems through reintroducing extirpated species. While some extinctions have been caused by climatic and environmental change, most have been caused by anthropogenic impacts. We propose a general model to describe patterns in these anthropogenic island extinctions. Hunting, habitat loss and the introduction of invasive predators accompanied prehistoric settlement and caused declines of endemic island species. Later settlement by European colonists brought further land development, a different suite of predators and new drivers, leading to more extinctions. Extinctions alter ecological networks, causing ripple effects for islands through the loss of ecosystem processes, functions and interactions between species. Reintroduction of extirpated species can help restore ecosystem function and processes, and can be guided by palaeoecology. However, reintroduction projects must also consider the cultural, social and economic needs of humans now inhabiting the islands and ensure resilience against future environmental and climate change.

Radiocarbon evidence for the presence of mice on Madeira Island (North Atlantic) one millennium ago

Owing to the catastrophic extinction events that occurred following the Holocene arrival of alien species, extant oceanic island biotas are a mixture of recently incorporated alien fauna and remnants of the original fauna. Knowledge of the Late Quaternary pristine island faunas and a reliable chronology of the earliest presence of alien species on each archipelago are critical in understanding the magnitude and tempo of Quaternary island extinctions. Until now, two successive waves of human arrivals have been identified in the North Atlantic Macaronesian archipelagos (Azores, Madeira, Selvagens, Canary and Cape Verde Islands): ‘aboriginal’, which is limited to the Canary Islands around two millennia ago, and ‘colonial’, from the fourteenth century onwards. New surveys in Ponta de São Lourenço (Madeira Island) have allowed us to obtain and date ancient bones of mice. The date obtained (1033 ± 28 BP) documents the earliest evidence for the presence of mice on the island. This date extends the time frame in which the most significant ecological changes occurred on the island. It also suggests that humans could have reached Madeira before 1036 cal AD, around four centuries before Portugal officially took possession of the island.

Late-Holocene asynchronous extinction of endemic mammals on the eastern Canary Islands

The Lava mouse (Malpaisomys insularis), and the Canarian shrew (Crocidura canariensis) are endemic of the Eastern Canary Islands and islets. The former is extinct while Canarian shrew survives in the two main islands and two islets. In order to provide insights regarding causes and processes contributing to the extinction of these endemic mammals: (i) we established last occurrence dates for Lava mouse, and first records for two exotic species – House mouse (Mus musculus) and Black rat (Rattus rattus) – through direct 14C AMS dating of collagen from bones; (ii) we analysed recent material from Barn owl (Tyto alba gracilirostris) roosting sites to evaluate its impact on Canarian shrew in the presence of introduced rodents. The new data strongly suggest that the extinction of Lava mouse was the result of an accumulative process of independent disappearances (or ‘local extinctions’) affecting the isolated populations. The timing of the introduction of the Black rat on the main islands (before Middle Age European contact in Lanzarote and after Middle Age European contact in Fuerteventura) matches with the last occurrence dates for the presence of Lava mouse on these islands, and are very probably their cause. The losses of these Lava mouse populations occurred in an asynchronous way, spreading across at least six centuries. On small islands, hyperpredation emerges as the most plausible process to explain the disappearance of the Lava mouse in the absence of rat populations, although stochastic processes can not be definitively excluded.