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On two collections of reptiles and amphibians from the Cape Verde Islands, with descriptions of three new taxa
... The radiation of the geckos after the single colonization event gave origin to four currently accepted endemic species with several subspecies, some of them exclusive to one of these island groups: T. darwini Joger 1984b, T. caboverdiana Schleich 1984, T. rudis Boulenger, 1906, and T. gigas (Bocage, 1875. However, the most exhaustive recent revision regarding the genetic vari- ability of Tarentola from the Cape Verde Islands using mitochondrial markers recovered 15 evolutionary sig- nificant units (ESUs) arranged into four main groups ( Fig. 2; Vasconcelos et al., 2010) not completely congru- ent with the current taxonomy (Schleich, 1987;Joger, 1993). The first group included all T. darwini plus T. rudis from Boavista, although both the bootstrap and posterior probability (PP) values were low; the second one grouped T. caboverdiana from S?o Vicente, Santa Luzia, Branco, Raso, and Santo Ant?o; the third one was exclusively formed by T. caboverdiana nico- lauensis from S?o Nicolau; finally, the fourth group included the remaining T. rudis populations. ...
... A total of 135 sequences of Cape Verdean Tarentola were included in the genetic analyses of the nuclear data and 92 in the multivariate morphological analy- sis. All specimens were identified in the field using diagnostic characters published by Joger (1984bJoger ( , 1993 and Schleich (1987) and a piece of tail was removed and stored in 96% ethanol. Before the animals were released, digital photographs (from dorsal, ventral, and lateral parts) were taken to quali- tatively analyse the colour pattern characteristics that may disappear in preserved specimens and to perform pholidotic counts a posteriori. ...
... Appendix 2); eye/ear opening ratio averages 1.59; ear-eye/eye- snout distance ratio averages 0.83. Eight to 11 supralabials; seven to nine infralabials; nine to ten enlarged lamellae under the 4th finger; 112-143 midbody scales (Joger, 1993); narrow central keeled dorsal tubercles ( Fig. 5A1) with 20-24 midbody lon- gitudinal lines and 14-18 transverse rows; prominent tubercle above and anterior to the ear opening. Light orangey or yellowish to pinkish grey dorsal coloration slightly translucent with reduced pattern in adults (Figs 6A1, 7A1) and whitish below. ...
Recent phylogeographical analyses using mitochondrial DNA (mtDNA) sequences indicate that the Tarentola geckos from the Cape Verde archipelago originated from a propagule that dispersed from the Canary Islands approximately 7.7 Mya and that underwent a fast evolutionary radiation. Molecular analyses carried out to date clearly show some incongruences with the current taxonomy of Tarentola from the Cape Verde Islands, with some species being paraphyletic or polyphyletic, and several independently evolving lineages needing formal taxonomic recognition. The aim of this study was to clarify the systematics of this group to unravel its taxonomy by applying an integrative approach based on information from three independent sources: mtDNA, nuclear genes, and morphology. As a result of this taxonomic revision, two novel species for the islands of S. Nicolau and Fogo are described and eight subspecies are upgraded to species level. Moreover, an identification key for the genus Tarentola from the Cape Verde archipelago is presented. This study reconciles taxonomy and phylogeny in this group, provides a better understanding of diversity patterns, new insights on evolutionary hypotheses, and supports the basic framework for the future management and conservation of this unique reptile radiation.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 328–360.
... It is a volcanic archipelago with 10 islands and various islets, ranging from 26 to 6 Myr old (Fig. 1). Before this study, 13 extant native reptile species were recognised (see Joger 1993;Arnold et al. 2008), all endemic to the archipelago. These belong to three genera: the Hemidactylus and Tarentola geckos and the Chioninia skinks. ...
... These belong to three genera: the Hemidactylus and Tarentola geckos and the Chioninia skinks. Within the latter, six extant species were recognised by Joger (1993): Chioninia delalandii (Duméril & Bibron, 1839), Chioninia vaillanti (Boulenger, 1887), Chioninia fogoensis (O'Shaughnessy, 1874), Chioninia geisthardti (Joger, 1993), Chioninia stangeri (Gray, 1845), Chioninia spinalis (Boulenger, 1906) and the extinct Chioninia coctei (Duméril & Bibron, 1839). Although the phylogenetic relationships within Chioninia have been investigated previously (Brehm et al. 2001;Brown et al. 2001;Carranza et al. 2001), all these studies stressed that a review of the systematics of the Cape Verdean skinks was needed. ...
... These belong to three genera: the Hemidactylus and Tarentola geckos and the Chioninia skinks. Within the latter, six extant species were recognised by Joger (1993): Chioninia delalandii (Duméril & Bibron, 1839), Chioninia vaillanti (Boulenger, 1887), Chioninia fogoensis (O'Shaughnessy, 1874), Chioninia geisthardti (Joger, 1993), Chioninia stangeri (Gray, 1845), Chioninia spinalis (Boulenger, 1906) and the extinct Chioninia coctei (Duméril & Bibron, 1839). Although the phylogenetic relationships within Chioninia have been investigated previously (Brehm et al. 2001;Brown et al. 2001;Carranza et al. 2001), all these studies stressed that a review of the systematics of the Cape Verdean skinks was needed. ...
Miralles, A., Vasconcelos, R., Perera, A., Harris, D. J. & Carranza, S. (2010). An integrative taxonomic revision of the Cape Verdean skinks (Squamata, Scincidae). —Zoologica Scripta, 40, 16–44.
A comprehensive taxonomic revision of the Cape Verdean skinks is proposed based on an integrative approach combining (i) a phylogenetic study pooling all the previously published molecular data, (ii) new population genetic analyses using mitochondrial and nuclear data resulting from additional sampling, together with (iii) a morphological study based on an extensive examination of the scalation and colour patterns of 516 live and museum specimens, including most of the types. All Cape Verdean species of skinks presently recognised, formerly regarded as members of the genera Mabuya Fitzinger, 1826 and Macroscincus Bocage, 1873 are considered as members of the Cape Verdean endemic genus Chioninia Gray, 1845. The new phylogeny and networks obtained are congruent with the previously published phylogenetic studies, although suggesting older colonization events (between 11.6 and 0.8 Myr old), and indicate the need for taxonomic changes. Intraspecific diversity has been analysed and points to a very recent expansion of Chioninia delalandii on the southern islands and its introduction on Maio, to a close connection between Chioninia stangeri island populations due to Pleistocene sea-level falls and to a generally low haplotypic diversity due to the ecological and geological characteristics of the archipelago. Three new consistent morphological synapomorphies supporting two of the four main clades of the genus have been identified. The complex taxonomic status of Euprepes fogoensis O’Shaughnessy, 1874 has been resolved and a lectotype has been designated for this species; Chioninia fogoensis nicolauensis (Schleich, 1987) is elevated to species rank, whereas Chioninia fogoensis antaoensis (Schleich, 1987) is now regarded as a junior subjective synonym of C. fogoensis. Additionally, one new subspecies of Chioninia vaillanti and two of Chioninia spinalis are described (Chioninia vaillanti xanthotis ssp. n., Chioninia spinalis santiagoensis ssp. n. and Chioninia spinalis boavistensis ssp. n.) and a lectotype has been designated for Mabuia spinalis Boulenger, 1906. Finally, an identification key for the Chioninia species is presented.
... The giant scincid Macroscincus coctei was another endemic species but it is believed to have been extinct from the beginning of the twentieth century (Schleich, 1982). The taxonomic status of some of these 6 species remains controversial especially at the subspecific level (e.g., Mertens, 1955; Schleich, 1987; Joger, 1993). Based on morphological characters, the two sympatric species M. delalandii and M. vaillanti are considered to form a closely related group (Greer, 1976; Pinheiro, 1989; Joger, 1993). ...
... The taxonomic status of some of these 6 species remains controversial especially at the subspecific level (e.g., Mertens, 1955; Schleich, 1987; Joger, 1993). Based on morphological characters, the two sympatric species M. delalandii and M. vaillanti are considered to form a closely related group (Greer, 1976; Pinheiro, 1989; Joger, 1993). They possess fused interparietals and parietals (unique in skinks) and retain 26 presacral vertebrae, which is typical in Mabuya (Greer et al., 2000). ...
... M. fogoensis is distributed on the windward islands of Santo Antã o and Sã o Nicolau and is divided into two endemic subspecies, M. f. antaoensis and M. f. nicolauensis (Schleich, 1987). The remaining 3 species, M. stangeri, M. salensis, and M. spinalis, share several morphological characteristics and have been the center of most of the taxonomic controversy (Angel, 1935; Mertens, 1955; Schleich, 1987; Joger, 1993). Based on morphological characters Joger (1993) proposed that all Scincidae from the Cape Verde islands, including the extinct giant skink M. coctei, are a monophyletic group presumably from the nearby West African coast. ...
Partial DNA sequences from two mitochondrial (mt) and one nuclear gene (cytochrome b, 12S rRNA, and C-mos) were used to estimate the phylogenetic relationships among the six extant species of skinks endemic to the Cape Verde Archipelago. The species form a monophyletic unit, indicating a single colonization of the islands, probably from West Africa. Mabuya vaillanti and M. delalandii are sister taxa, as indicated by morphological characters. Mabuya fogoensis and M. stangeri are closely related, but the former is probably paraphyletic. Mabuya spinalis and M. salensis are also probably paraphyletic. Within species, samples from separate islands always form monophyletic groups. Some colonization events can be hypothesized, which are in line with the age of the islands. C-mos variation is concordant with the topology derived from mtDNA.
... The Cape Verde Islands are volcanic and comprise nine major inhabited islands and several smaller uninhabited islands and islets. Recent taxonomic revisions have elevated the number of endemic Mabuya species to six ( based on morphology), with two of these ( M. spinalis and M. fogoensis ) being further divided into several subspecies (Schleich 1987;Joger 1993). Prior to this, M. spinalis had been incorporated (as three subspecies) within M. stangeri . ...
... M. delalandii is also widespread and found on at least three major neighbouring islands and several islets in the southern Sotavento group (and possibly Boavista in the north-east) ( Fig. 1). Here, differentiation appears lower and no subspecies have been described, although small interisland differences in scalation have been noted ( Joger 1993). We hypothesized that this lower diversity is a result of more recent colonizations as opposed to similar selection regimes on different islands. ...
... There were 238 variable sites across all Mabuya sequences ( giving rise to 23 variable amino acid sites), 169 variable Schleich (1987) and Joger (1993). Although there are museum records for M. delalandii in Boavista, we were unable to detect it despite 4 days of intensive searching. ...
The Cape Verde Islands are of volcanic origin with most having appeared between the early Miocene and mid-Pleistocene. They contain six known species of Mabuya skinks. Phylogeographical relationships within and among the relatively widespread taxa M. stangeri, M. spinalis and M. delalandii were inferred, based on approximately 1 kbp of the cytochrome b gene (mitochondrial DNA). Reciprocal monophyly of M. spinalis and M. stangeri was established, which may have arisen from an early Pliocene/late Miocene cladogenetic event. Considerable between-island sequence divergence was detected among M. spinalis, which appears to have colonized the older islands (Sal and Boavista) first. Much lower sequence divergence was found in M. delalandii, indicating a more recent range expansion. Here, evidence points to colonization of the younger islands of Brava and Fogo soon after appearance. There are similarities between some of the described patterns and those seen in lizards from the Canary Islands.
... Additional specimens examined are shown in table 1. Specimens collected in the field were identified following Joger(1993) and Schleich (1987), and released after tail tips were taken. Tissues are preserved in the collections of CCBG (University of Madeira). ...
... However, two of these came from the island of Sao Vicente (T23850 and T23851). Previously only 1: c. substituta has been recorded from this island (Joger, 1993). We also included two samples of 1: c. caboverdiana Schleich, 1984 from Santo Anmo. ...
Thirteen specimens of Tarentola from the Cape Verde islands were sequenced for 695 base pairs of 12S rRNA and cytochrome b mitochondrial genes, and analysed with published sequences. Our results support many of the relationships previously proposed. We report the presence of Tarentola gigas Bocage, 1875 on São Nicolau and Tarentola caboverdiana nicolauensis Schleich, 1984 on São Vicente. This increases the number of genetically distinct forms on these islands; hence community structure appears to be more complex than previously understood. We also sequenced seven individuals for 375 base pairs of the nuclear gene, C-mos. Two sites were variable, much less than expected given the high levels of differentiation based on mitochondrial DNA sequences.
... Both species were introduced there (Klemmer 1976 ;Recuero et al. , 2007Recuero et al. , : 1214. Finally, the Cape Verde Islands in the far south of Macaronesia , 500 km from Senegal is known only to one kind of amphibians : it is the widespread there tropical African toad Bufo regularis Reuss, 1833 (Joger 1993). ...
... Segundo dados de campanhas oceanográficas realizadas na Zona Económica Exclusiva de Cabo Verde (ZEE), a produtividade primária das águas varia de 150-500 mg de C/m 2 /dia o que o insere na categoria de águas de alta produtividade (Almada, 1993). Apesar disso, poucos estudos filogenéticos têm sido feitos nas ilhas, e a maior parte baseados em espécies terrestres (répteis e anfíbios) como é o caso dos trabalhos de Greer (1976), Joger (1993), Carranza et al. (2001), Brehm et al. (2001) e Jesus et al., (2002. divididas em 2 grupos principais relacionados com o tamanho da concha: de "concha grande", variando de 37 a 75 mm, e de "concha pequena entre 10 a 29 mm (Cunha et al., 2005). ...
... A total of 405 new specimens of Tarentola were included in the genetic analyses. Specimens were identified in the field using diagnostic characters published by Joger (1984Joger ( , 1993 and Schleich (1987), digital photographs were taken and a piece of tail was removed and stored in 96% ethanol. Sampled animals were released afterwards. ...
Aim To reassess the relationships between Tarentola geckos from the Cape Verde Islands by including specimens from all islands in the range. To determine the variation within forms by sequencing over 400 specimens, thereby allowing the discovery of cryptic forms and resolving some of the issues raised previously. This extensive sampling was also used to shed light on distributions and to explain genetic diversity by comparing the ages and ecological and geological features of the islands (size, elevation and habitat diversity).
Location The Cape Verde Islands: an oceanic archipelago belonging to the Macaronesian biogeographic region, located around 500 km off Senegal.
Methods A total of 405 new specimens of Tarentola geckos were collected from nine islands with very different geological histories, topography, climate and habitats. Mitochondrial cytochrome b (cyt b ) gene and 12S rRNA partial sequences were obtained and analysed using phylogenetic methods and networks to determine molecular diversity, demographic features and phylogeographic patterns.
Results The phylogenetic relationships between all known forms of Cape Verdean Tarentola specimens were estimated for the first time, the relationships between new forms were assessed and previously hypothesized relationships were re‐examined. Despite the large sample size, low intraspecific diversity was found using a 303‐bp cyt b fragment. Star‐like haplotype networks and statistical tests suggest the past occurrence of a rapid demographic and geographical expansion over most of the islands. Genetic variability is positively correlated with size, elevation and habitat diversity of the islands, but is not linearly related to the age of the islands. Biogeographical patterns have, in general, high concordance with phylogenetic breaks and with the three eco‐geographical island groups. Volcanism and habitat diversity, both tightly linked with island ontogeny, as postulated by the general dynamic model of oceanic island biogeography, as well as present and historical size of the islands appear to be the main factors explaining the genetic diversity of this group.
Main conclusions The Tarentola radiation was clarified and is clearly associated with the geological and ecological features of the islands. Two factors may account for the low intraspecific variation: (1) recent volcanic activity and high ecological stress, and (2) poor habitat diversity within some islands. More studies are needed to align taxonomy with phylogenetic relationships, whereas GIS modelling may help to predict precise species distributions.
... Antão, St. Luzia, Raso, S. Nicolau, Santiago; also possibly Brava (Fea 1899b;Angel 1937;Loveridge 1947;Mertens 1955). Although it as also been reported from Fogo by Angel (1935Angel ( , 1937, Loveridge (1947), Mertens (1955) Schleich (1982, 1996, Joger (1993) and López-Jurado et al. (2005); at least some of these reports may actually refer to H. lopezjuradoi. (Bocourt 1870) (Fig. 5E) Emydactylus bouvieri Bocourt (1870) Distribution. ...
A total of 1854 bp of mitochondrial DNA (669 bp of cytochrome b (cyt b) and 386 bp of 12S rRNA), and 804 bp of a nuclear gene (RAG2) were investigated in endemic Hemidactylus from eight Cape Verde Islands, and used to explore their phylogeny, biogeography and evolution. Maximum-likelihood, maximum-parsimony and Bayesian analyses based on mtDNA revealed four well-supported clades with uncorrected genetic divergences of 7.8–12.4% in the cyt b plus 12S rRNA genes, which were also supported by nuclear DNA. A population from the southern island of Fogo is the most divergent in both molecules and morphology and is described as Hemidactylus lopezjuradoi sp. n., and the populations on Sal and Boavista are also assigned species status as H. boavistensis. Although divergent in their DNA, the clade on S. Nicolau and that in the north-western islands are morphologically similar and both are assigned to H. bouvieri for the present. Hemidactylus b. razoensis from Raso is genetically similar to H. b. bouvieri and differs only in its smaller body size. A molecular clock suggests that the ancestor of the endemic Hemidactylus of the Cape Verde Islands colonized the archipelago approximately 10 ± 2.48 Mya, perhaps reaching the north-eastern islands first. The H. lopezjuradoi lineage separated soon after, and the north-western islands were colonized progressively but slowly, S. Nicolau probably being reached first, then S. Vicente and islands on the same bank, and finally Sto. Antão, which is likely to have been colonized less than 1 Mya. Hemidactylus boavistensis is abundant on the arid islands where it occurs, but H. bouvieri appears to have been uncommon at least since it was described 130 years ago, and the same may be true of H. lopezjuradoi sp. n. The impact of introduced H. angulatus and H. mabouia on the endemic Hemidactylus of the Cape Verde Islands is not clear, but the discovery of substantial genetic diversity in endemic Cape Verde Hemidactylus means that the conservation requirements of the group should be reassessed.
Chatogekko amazonicus is a miniaturized gecko from northern South America and is among the smallest of toe pad bearing lizards. The toe pads of C. amazonicus are miniscule, between 18% and 27% of the plantar surface area. We aimed to investigate the relationship between adhesive toe pad morphology, body size, and adhesive capabilities. Using scanning electron microscopy, we determine that the adhesive pads of C. amazonicus exhibit branched setae similar to those of other geckos, but that are generally much smaller. When compared with other gecko taxa, we show that C. amazonicus setae occupy a similar range of seta length: snout–vent length ratio and aspect ratio as other gekkonoid species (i.e. Gekkonidae, Phyllodactylidae, and Sphaerodactylidae). We demonstrate that C. amazonicus—even with its relatively small toe pads—is capable of climbing a smooth glass surface at a nearly vertical angle. We suggest that sphaerodactylids like C. amazonicus offer an excellent system for studying toe pad morphology and function in relation to miniaturization.
See http://www.afriherp.org/gekkota-catalog/ for details
Some general principles of zoogeography are considered. The regionalization based on various animal groups might result in different zoogeographic schemes. Therefore, it would be quite desirable to accumulate reliable data with various taxonomic groups as a subject of comparative zoogeography. Zoogeographic regionalization should be based on animal distribution itself but not on any other external factors. The identification of realm affinities of species should include both the recent distribution of the species and its phylogenetic position (relationships) within higher taxa (species groups, genus, etc.). The distribution of amphibian species in the north, west (Atlantic archipelagos) and east of temperate Eurasia was analysed in order to identify the borders of the Palearctic Realm. Special attention was paid to the southern limits of the Palearctic in North Africa (Sahara Desert), Middle East (Sinai, Arabian Peninsula, Iran), Afghanistan, Pakistan, western Himalaya, as well as in China аnd Japan. Outlined territory of the Palearctic harbors 214 amphibian species from 47 genera, 17 families and two orders. The composition of amphibians of the Palearctic is analysed in taxonomic and zoogeographic aspects. The Palearctic share was evaluated in relation to amphibians of the World, endemism level among species, genera, and families was calculated as well. Differentiation of genera (monotypic and other, in terms of species richess and range’s size) as well as species was discussed. Attached are the description of a new toad genus, Strauchophryne gen. nov., and full list of amphibian species, genera and families of the Palearctic, with designation of realm affinity and endemism for all species.
Cape Verde has a higher number of reptile taxa and endemics than any of the five archipelagos in the Macaronesian region. Mapping the precise distributions and assessing the conservation status of reptiles is the first step towards effective conservation. Presence/absence and abundance data were gathered from extensive fieldwork and post-1980 literature. Evaluation of conservation status was considered at specific and subspecific levels, following IUCN Red List criteria and using RAMAS. Fieldwork confirmed the occurrence of 34 of 37 previously recorded taxa (31 native, three exotic). One taxon continues to be considered Extinct. Three broad distribution and rarity patterns were identified: widespread and abundant taxa occurring on ≥ 2 islands/islets, widespread or abundant taxa restricted to one island, and rare or limited range taxa occurring on small areas of islands or islets. More than a third of taxa have areas of occupancy < 20 km2 and extents of occurrence < 100 km2. Geckos are rarer than skinks because of their high habitat specialization, with 58% occurring on only one island/islet. About half of all taxa are potentially threatened, twice the proportion of those in the Canary Islands, a difference that could be explained by the smaller area and greater aridity of the Cape Verde islands. The criterion used for most threat categorizations is geographical range, and the most pervasive threats are natural disasters, intrinsic factors of the species and introduced species. The importance of applying conservation status at the subspecific level to island endemics is emphasized. Several conservation measures are proposed, including optimized design of protected areas.
We provide the first complete list of the present and lost amphibian and reptile type specimens of the Hessisches Landesmuseum Darmstadt and the Museum Wiesbaden Naturhistorische Landessammlung. The Darmstadt collection currently houses primary types of 5 taxa (holotypes) and secondary types of addi-tional 15 taxa (paratypes). The Wiesbaden collection includes primary types of 13 taxa (8 represented by holotypes, 4 by syntypes and 1 by lectotype). Furthermore, the primary types of 6 taxa formerly housed at the Museum Wiesbaden are consid-ered lost. In several cases, we comment on the current status of the taxa present in the collections. Bufo spinulosus var. arapensis Andersson, 1908 was obviously overlooked and neglected in the literature. It is here considered a junior synonym of Chaunus spinulosus (Wiegmann, 1834). The largest specimen of the type series is designated as lectotype (MWNH 153/1). The current status of Hylambates rufus var. aubryioides Andersson, 1908, as a junior synonym of Leptopelis modestus (Werner, 1898) is rejected.
Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk.
Location World‐wide.
Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions.
Results I obtained body size data for 99% of the world's lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non‐aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed.
Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.
Morphological systematics makes it clear that many non-volant animal groups have undergone extensive transmarine dispersal with subsequent radiation in new, often island, areas. However, details of such events are often lacking. Here we use partial DNA sequences derived from the mitochondrial cytochrome b and 12S rRNA genes (up to 684 and 320 bp, respectively) to trace migration and speciation in Tarentola geckos, a primarily North African clade which has invaded many of the warmer islands in the North Atlantic Ocean. There were four main invasions of archipelagos presumably by rafting. (i) The subgenus Neotarentola reached Cuba up to 23 million years (Myr) ago, apparently via the North Equatorial current, a journey of at least 6000 km. (ii) The subgenus Tarentola invaded the eastern Canary Islands relatively recently covering a minimum of 120 km. (iii) The subgenus Makariogecko got to Gran Canaria and the western Canary Islands 7-17.5 Myr ago, either directly from the mainland or via the Selvages or the archipelago of Madeira, an excursion of 200-1200 km. (iv) A single species of Makariogecko from Gomera or Tenerife in the western Canaries made the 1400 km journey to the Cape Verde Islands tip to 7 Myr ago by way of the south-running Canary current. Many journeys have also occurred within archipelagos, a minimum of five taking place in the Canaries and perhaps 16 in the Cape Verde Islands. Occupation of the Cape Verde archipelago first involved an island in the northern group, perhaps São Nicolau, with subsequent spread to its close neighbours. The eastern and southern islands were colonized from these northern islands, at least two invasions widely separated in time being involved. While there are just three allopatric species of Makariogecko in the Canaries, the single invader of the Cape Verde Islands radiated into five, most of the islands being inhabited by two of these which differ in size. While size difference may possibly be a product of character displacement in the northern islands, taxa of different sizes reached the southern islands independently.
The scincid lizards of the Cape Verde islands comprise the extinct endemic giant Macroscincus coctei and at least five species of Mabuya, one of which, Mabuya vaillanti, also had populations with large body size. Phylogenetic analysis based on DNA sequences derived from the mitochondrial cytochrome b, cytochrome oxidase I and 12S rRNA genes (711, 498 and 378 base pairs (bp), respectively) corroborates morphological evidence that these species constitute a clade and that Macroscincus is unrelated to very large skinks in other areas. The relationships are ((M. vaillanti and Mabuya delalandii) (Mabuya spinalis and Macroscincus coctei (Mabuya fogoensis nicolauensis (Mabuya fogoensis antaoensis and Mabuya stangeri)))). The Cape Verde archipelago was colonized from West Africa, probably in the Late Miocene or Early Pliocene period. The north-eastern islands were probably occupied first, after which the ancestor of M. vaillanti and M. delalandii may have originated on Boavista, the ancestor of the latter species arriving on Santiago or Fogo later. The M. fogoensis--M. stangeri clade colonized the islands of Branco, Razo, Santa Luzia and São Vicente from São Nicolau and reached Santo Antão after this. Colonization of these northeastern islands was slow, perhaps because the recipient islands had not developed earlier or because colonization cut across the path of the Canary Current and the Northeast Trade Winds, the main dispersing agents in the region. Rapid extension of range into the southwestern islands occurred later in M. spinalis and then in M. vaillanti and M. delalandii. The long apparent delay between the origin of these species and their southwestern dispersal may have been because there were earlier colonizations of the southern islands which excluded later ones until the earlier inhabitants were exterminated by volcanic or climatic events. The evolution of large size in Macroscincus occurred in the northwestern islands and was paralleled in the eastern and southern islands by populations of M. vaillanti. Both cases of size increase in Cape Verde skinks were accompanied by the development of herbivory.
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