Darwin and the Galápagos

Charles Darwin's historic visit to the Galápagos Islands in 1835 represents a landmark in the annals of science. But contrary to the legend long surrounding Darwin's famous Galápagos visit, he continued to believe that species were immutable for nearly a year and a half after leaving these islands. This delay in Darwin's evolutionary appreciation of the Galápagos evidence is largely owing to numerous misconceptions that he entertained about the islands, and their unique organic inhabitants, during the Beagle voyage. For example, Darwin mistakenly thought that the Galápagos tortoise–adult specimens of which he did not collect for scientific purposes–was not native to these islands. Hence he apparently interpreted reports of island-to-island differences among the tortoises as analogous to changes that are commonly undergone by species removed from their natural habitats. As for Darwin's finches, Darwin initially failed to recognize the closely related nature of the group, mistaking certain species for the forms that they appear, through adaptive radiation, to mimic. Moreover, what locality information he later published for his Galápagos finch specimens was derived almost entirely from the collections of three other Beagle shipmates, following his return to England. Even after he became an evolutionist, in March of 1837 (when he discussed his Galápagos birds with the eminent ornithologist John Gould), Darwin's theoretical understanding of evolution in the Galápagos continued to undergo significant developments for almost as many years as it took him to publish the Origin of Species (1859). The Darwin-Galápagos legend, with its romantic portrait of Darwin's 'eureka-like' insight into the Galápagos as a microcosmic 'laboratory of evolution', masks the complex nature of scientific discovery, and, thereby, the real nature of Darwin's genius.     

Identity and taxonomic affinity of some members of the Amaranthaceae from the Galápagos Islands

ELIASSON, U. H. 1985. Identity and taxonomic affinity of some members of the Amaranthaceae from the Galápagos Islands. The type species of Pleuropetalum, P. DarwiniI, is endemic to the Galápagos and is morphologically closer to P. pleiogynum than to P. sprucei. Pleuropetalum pleiogynum has a wider distribution than previously assumed, ranging from Costa Rica to Peru. Amaranlhus anderssonii is closely related to A. urceolatus, a species of the Pacific coast of S America, and to A. berlandieri, a species of Texas and NE Mexico. Amaranlhus squamulatus, previously regarded as endemic to the Galápagos, is reported from the Guayas province of Ecuador. The genus Galápagosus, proposed for Amaranlhus sclerantoides, is rejected. Lithophila scirpoides, an overlooked taxon from the Galápagos, is believed to be an aberrant specimen of L. radicala. Lithophila subscaposa from the islands of Santiago and Pinzon differs morphologically from the specimens of Floreana. Iresine edmonstonii, probably erroneously reported from the Galápagos, is conspecific with J. angusti/olia. Blutaparon rigidum is morphologically divergent from other members of the genus, being adapted to arid conditions, and is assumed to be extinct.     

Darwin was astonished

Darwin did not approach the Galápagos with the same enthusiasm and energy as he showed at earlier places visited by the Beagle. Notwithstanding, he looked back on the five weeks the Beagle spent in the Galápagos as a time when he made observations important for the development of his evolutionary ideas. In retrospect, he was astonished at what he saw there.     

Scalesia gordilloi sp. nov. (Asteraceae) from the Galápagos Islands, Ecuador

Scalesia gordilloi está descrita desde la isla San Cristóbal, Islas Galápagos, Ecuador. Tiene una distribución limitada y local al lado de la costa sur en la zona vegetal árida. Está relacionada a S. divisa Anderss. y S. incisa Hook. f. Se discute la taxonomía de S. gordilloi.     

A new look at evolution in the Galápagos: evidence from the late Cenozoic marine molluscan fauna

Endemism is not as common in the marine invertebrate fauna of the Galápagos Islands region as in the adjacent terrestrial biota. Marine invertebrates in the Galápagos are largely cosmopolitan species from the Panamic, Indo-Pacific, Californian, or Peruvian faunal provinces. However, an endemic component is also present in the fauna. The observed pattern among marine invertebrate organisms can be accounted for by at least two processes: (1) genetic continuity between mainland and island populations mediated through planktonic larvae; and (2) lower rates of intrinsic evolutionary change. The evolutionary scenario standardly applied to terrestrial organisms in the Galápagos, namely, adaptive radiation and speciation in reproductive isolation from mainland source populations, does not apply to all marine invertebrates. Evidence in support of the alternative scenario for marine invertebrates comes from both published records of species occurring in the islands and recent studies of fossil-bearing deposits on several islands in the archipelago. Two misconceptions–considering the islands and sedimentary deposits to be older than now thought, and equating the rate of evolution of the terrestrial biota with the marine biota–can lead to an incorrect interpretation of evolution in the Galápagos Contrasts between marine invertebrate and terrestrial organisms serve to illustrate some fundamental differences which have important evolutionary implications. Some of these are: endemism; dispersal; taxonomic relationships; island definitions; rates of evolutionary change; and age of fossils. In terms of Darwin's evolutionary scenario, terrestrial organisms represent the paradigm and marine organisms represent the paradox.     

Geology of Galápagos

This paper reviews the origins and geological history of the Galápagos Islands. The islands arose from a 'hot-spot'. The oceanic crust on which the islands are built can be no more than 10 million years old, and the islands themselves have been in existence at least 3.3 million years.
The Galápagos are among the most active volcanic groups in the world, and the physical nature of the islands is dominated by lava structures of various ages, including lava tubes or tunnels, which have been of particular interest to biologists. Weathering of the rock to produce soil has generally been slow, particularly in the drier parts.     

Reproductive ecology of the endemic Lecocarpus pinnatifidus (Asteraceae) in an isolated population in the Galápagos Islands

Lecocarpus pinnatifidus is an endemic member of the Asteraceae occurring on only one island in the Galápagos archipelago. The capitula are large with female ray florets and male disc florets. They are self-compatible but this study suggests fruit set is pollen limited. Visits from Xylocopa darwini and other larger insect pollinators are rare, and small insects seem to be the main pollinators. Small insects carry few pollen grains and most likely mediate self-pollinations. Self-compatibility and seed set after selfing are the most common reproductive strategy in the Galápagos Islands and L. pinnatifidus seemingly fits well into this group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 171–180.     

Limited dispersal by Nazca boobies Sula granti

We documented natal and breeding dispersal at several spatial scales by Galápagos Nazca boobies Sula granti , a wide-ranging pelagic seabird. We found exceptionally low degrees of both types of dispersal despite these birds' vagility. Median natal dispersal distances were 26 m and 105 m for males and females, respectively. Median breeding dispersal distances for both sexes were 0 m. No natal or breeding dispersals occurred from our study site at Punta Cevallos, Isla Española to six other colonies in the Galápagos, but we did document four long-distance natal dispersals from Punta Cevallos to islands near the South American coast. Recaptures and dead recoveries of ringed birds showed long distance non-breeding movements to the Central American coast and elsewhere in the eastern Pacific, contrasting with the very limited dispersal to breeding sites.     

The vascular plants of Joseph Dalton Hooker's An enumeration of the plants of the Galapagos Archipelago; with descriptions of those which are new

The first floristic work on the Galápagos Islands was published by J. D. Hooker in 1847. It was based mainly on the Beagle collections of Charles Darwin, but those of James McRae, John Scouler. Hugh Cuming, David Douglas, Thomas Edmonston, Abel Du Petit-Thouars and John Goodridge were also used, though those of Archibald Menzies, the first botanist to visit the archipelago, were not. The visits of these naturalists and their collections are discussed. Darwin's field notes on Galápagos plants, hitherto unpublished, are given, and most of the species identified. In the taxonomic portion of the paper, each taxon included by Hooker is discussed, the nomenclaturally or-taxonomically correct name is indicated, specimens examined by Hooker are enumerated, and additional specimens presumably seen but not cited by him are given. Hooker wrote before our present type concept had evolved, and consequently 51 of his 114 new names needed to be typified, as did four others based on Darwin Galápagos collections. In addition, examination of these specimens revealed one species not yet reported from the archipelago ( Galium canescens ), one new island record ( Chamaesyce hirta on James Island), and one deletion from the flora ( Malachra capitala )     

Hybridization between the Galápagos land and marine iguana (Canolophus subcristatus and Amblyrhynchus cristatus) on Plaza Sur

Hybridization plays an important role in the evolution of some of the vertebrate taxa on the Galápagos Islands, such as the Darwin finches. Conversely, only a single possible hybrid between the Galápagos marine iguana ( Amblyrhynchus ) and the land iguana ( Conolophus ) has been reported from the island Plaza Sur. In this paper, the hybrid status of a morphologically unusual iguana from this island is confirmed, using restriction fragment length polymorphism (RFLP) analyses of the nuclear ribosomal DNA (rDNA). Sequencing of the hybrid's mitochondrial cytochrome b gene revealed that it was the offspring of a female land iguana and a male marine iguana. Preliminary molecular analyses of morphologically typical marine and land iguanas from Plaza Sur did not detect introgression of nuclear or mitochondrial markers between species. The potential significance of hybridization for the evolution of the Plaza Sur iguana populations is discussed.     

The bark and ambrosia beetles (Curculionidae, Scolytinae) of Cocos Island, Costa Rica and the role of mating systems in island zoogeography

Cocos Island is a small oceanic island midway between Costa Rica and the Galápagos Archipelago; about 2 Myr in age, it is the only tropical oceanic island in the eastern Pacific with tropical wet forest. We identified several hundred bark beetle specimens collected during recent expeditions by INBio, the National Biodiversity Institute of Costa Rica, and re-examined all specimens from earlier collections. We report 19 species in ten genera, seven or eight of which are endemic, making scolytines the largest group of beetles known from the island. We describe as new Pycnarthrum pseudoinsulare , Xyleborinus cocoensis , and Xyleborus sparsegranulosus , resurrect Xyleborus bispinatus as separate from X. ferrugineus , and report six other species as new to Cocos Island. Three-quarters of the scolytines reproduce by brother–sister mating, and we argue that inbreeders are superior island colonists because they are less affected than are outbreeders by problems of mate location and inbreeding depression. The fauna and flora of Cocos Island arrived by dispersal and human transport. We examine natural colonization patterns for the fauna, using the distributions of the relatives of island endemics: most colonization came from the Americas, but the closest relatives to some endemics are found on Caribbean or Galápagos islands. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 729–743.     

The microevolution of the Galápagos marine iguana Amblyrhynchus cristatus assessed by nuclear and mitochondrial genetic analyses

Marine iguanas may have inhabited the Galápagos archipelago and its former, now sunken islands for more than 10 million years (Myr). It is therefore surprising that morphological and immunological data indicate little evolutionary divergence within the genus. We utilized mitochondrial DNA (mtDNA) sequence analyses and nuclear DNA fingerprinting to re-evaluate the level and pattern of genetic differentiation among 22 marine iguana populations from throughout the archipelago. Both genetic marker systems detect a low level of within-genus divergence, but they show contrasting levels of geographical subdivision among the populations. The mitochondrial gene pools of populations from different regions of the archipelago are isolated, and the mtDNA pattern appears to follow the sequence in which the islands were colonized by marine iguanas. Conversely, the nuclear DNA study indicates substantial interpopulational gene exchange, and the geographical distribution of the nuclear markers seems to be determined by isolation by distance among the populations. The natural history of marine iguanas suggests that the contrasting nuclear and mitochondrial DNA patterns result from an asymmetric migration behaviour of the two sexes, with higher (active and passive) interisland dispersal for males than females. Separate genetic analyses for the sexes appear to support this hypophesis. Based on these findings, a scenario is proposed that explains the marine iguanas' low genetic divergence, notwithstanding their long evolutionary history in the Galápagos archipelago.     

Three Variable Arenopontia Species (Crustacea, Copepoda) from Panamá

Three species of the genus Arenopontia (Cylindropsyllidae) from the Pacific and Atlantic coast of Panamá are presented. Two of them are A. trisetosa Mielke and A. peteruxi Mielke, both of which were described for the first time from the Galápagos Islands. The third species is tentatively identified as A. gussoae Cottarelli previously reported from an interstitial habitat in Cuba.     

Chromosome number of Macraea Iaricifolia Hooker fil. (Compositae) and its bearhg on the taxonomic affinity of the genus

ELIASSON, U. H., 1984. Chromosome number of Macraea lariciflia Hooker fil. (Compositae) and its bearing on the taxonomic affinity of the genus. The chromosome number of Macraea laricifolia , a composite endemic to the Galápagos Islands, is reported to be 2 n = 28. Macraea was once regarded as congeneric with Lipochacta , a genus restricted to Hawaii. The chromosome number supports morphological evidence that the two genera arose independently from wedelioid ancestors within the Heliantheae.     

Increasing prevalence of avian poxvirus in Darwin's finches and its effect on male pairing success

Island populations harbour a comparatively species-poor pathogen community, often resulting in naïve host species that experience compromised immunity when faced with novel diseases. Over 95% of the Galápagos avifauna have survived 400 years of human settlement, yet currently face threats due to introduced diseases such as avian poxvirus. On Hawaii, declining populations of birds and even some extinctions have been attributed to avian poxvirus, and hence, identifying the prevalence and fitness costs of avian poxvirus on the Galápagos is a conservation priority. Surveys of avian poxvirus in Darwin's finches on Santa Cruz Island between 2000 and 2004 found a 33% annual increase in the prevalence of pox lesions in ground finches. Comparisons of pox prevalence on three islands (Santa Cruz, Floreana, and Isabela) were made in 2004, which indicated significant variation in pox prevalence across islands (Isabela>Santa Cruz>Floreana). Darwin's finch species were found to be differentially affected by poxvirus, with a higher prevalence in ground finches than in tree finches. There was a significant effect of habitat, even within species, with higher prevalence in the lowlands than highlands. Pox prevalence was not correlated with sex or body condition. However, male small ground finches Geospiza fuliginosa with evidence of pox were less likely to have a mate (16.6% paired) compared with males without pox (77% paired), indicating fitness costs associated with poxvirus infection.     

Islands and evolution: theory and opinion in Darwin's earlier years

Islands were not of special interest to evolutionists before Darwin. It was he who first appreciated their importance for demonstrating evolution in miniature. They were not of special interest because: (a) their peculiar products seemed no more peculiar than those of continents; (b) there was no special category of oceanic islands, but a continuum from such groups as the Canaries, Madeiras and Galápagos through New Zealand and Madagascar to Australia, Britain, and true continents; and (c) the concept of adaptive radiation, if known at all, was applied only to the higher levels of classification, and then very feebly.
When Darwin was young, classification at the lower levels hardly recognized convergence, and at the higher levels was subject to great changes, while only slowly separating out the major groups. In consequence, many of the facts of geographical distribution were misinterpreted, and numerous theories of the origination of species, groups, and biogeographical provinces were still plausible. It was largely the need for a historical, not ecological, explanation of the distribution of some mammals and plants, plus what he saw for himself in the Galápagos Islands, that convinced Darwin that evolution had occurred. His was a remarkable achievement in recognizing through all this 'noise' the meaning of adaptive radiation.     

Dynamics of a stand of Scalesia pedunculata Hooker fil., Santa Cruz Island, Galapagos

A seven-year study of a stand of the endemic Scalesia pedunculata (Asteraceae), which is the dominant tier ill the humid forest on the island oS Santa Cruz, Galápagos, reveals a characteristic pattern of growth and survival. The Scalesia trees initially show a rapid increase in both height and girth. After approximately 34–56 years the trees may reach 7–8 m in height and start producing flowers; the relative increase in girth then becomes more pronounced. The mortality rate is very high during the first 4–5 years, decreasing thereafter as the individuals age. It is suggested that the growth and survival pattern of S. pedunculata is an expression of an adaptive strategy fitted for a pioneer or early-successional tree. This tree is simultaneously able to persist in the community by the means of a large reproductive effort and a rapid population turnover. The method of seed-dispersal in the genus discussed in relation to the growth pattern.     

Population size and trends of the Galápagos Penguin Spheniscus mendiculus

We applied a capture–mark–resight (CMR) method to estimate the population size of the Galápagos Penguin Spheniscus mendiculus . In 1999, we estimated 1198 individuals, with lower and upper 95% confidence limits of 1054 and 1403 individuals, respectively, and estimated that approximately 57% of the total population was counted in an annual census. Applying this estimate to the 2003 census, we estimated that the population size for the whole archipelago was 1351 individuals. We also applied the correction factor derived from the 1999 CMR data to other censuses carried out between 1970 and 2003 and estimated a maximum population of about 4000 individuals in 1971 when the highest numbers of Penguins were counted. Although the Penguin population size has fluctuated in the last 33 years, the overall trend shows larger populations in 1970–80 followed by relatively smaller populations and a slow recovery rate in 1983–2003. The data set also shows two major population declines (in 1983 and 1998), which are coincident with El Niño episodes.     

THE DIET OF GALÁPAGOS SPERM WHALES PHYSETER MACROCEPHALUS AS INDICATED BY FECAL SAMPLE ANALYSIS

Fecal samples were collected while following sperm whales ( Physeter macrocephalus ) off the Galápagos Islands, Ecuador. They contained 133 upper beaks and 164 lower beaks of cephalopods. Analysis of the lower beaks suggests that the sperm whales fed primarily on three genera of cephalopods; Histioteutbis (62%), Ancistrocbeirus (16%), and Octopoteutbis (7%). The beak dimensions indicate that the cephalopods ranged in mantle length from 5 to 54 cm and in mass from 12 to 650 g. Fecal samples varied significantly between five study years and over different parts of the study area, but the number of beaks collected per sample did not correlate significantly with defecation rate (a measure of feeding success). Using beak material from fecal samples gives a biased estimate of sperm whale diet, reducing the frequencies of very small and very large cephalopods. However, all other available methods of assessing sperm whale diet also possess biases.     

Partial self‐incompatibility in the polyploid endemic species Scalesia affinis (Asteraceae) from the Galápagos: remnants of a self‐incompatibility system?

Different pollination treatments of capitula were used to examine the breeding system of individuals of the tetraploid endemic species Scalesia affinis from the Galápagos Islands. All types of crossings resulted in approximately 35 achenes per capitulum, but in actively and passively self-pollinated capitula these were mostly without embryos. Among self-pollinated individuals a large variation was found in the production of embryos. Some individuals failed completely to set filled achenes while others produced few or many. Additionally, we found a clear difference in female phenology of florets in self-pollinated capitula compared with florets in cross-pollinated capitula. Female florets in self-pollinated capitula remained receptive for longer time. These results suggest that Scalesia affinis is partly self-incompatible. Outcrossing is assured by the endemic carpenter bee, Xylocopa darwini , which proved to be an important pollinator of Scalesia affinis . Nevertheless, geitonogamy seemed to be considerable in the focal population, for which reason open pollinated capitula produced fewer achenes than controlled outcrossed capitula.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 93–101.