Camarhynchus Parvulus Research Articles

A beak size locus in Darwin's finches facilitated character displacement during a drought.

Ecological character displacement is a process of morphological divergence that reduces competition for limited resources. We used genomic analysis to investigate the genetic basis of a documented character displacement event in Darwin's finches on Daphne Major in the Galápagos Islands: The medium ground finch diverged from its competitor, the large ground finch, during a severe drought. We discovered a genomic region containing the HMGA2 gene that varies systematically among Darwin's finch species with different beak sizes. Two haplotypes that diverged early in the radiation were involved in the character displacement event: Genotypes associated with large beak size were at a strong selective disadvantage in medium ground finches (selection coefficient s = 0.59). Thus, a major locus has apparently facilitated a rapid ecological diversification in the adaptive radiation of Darwin's finches.

Divergent foraging behavior in a hybrid zone: Darwin’s tree finches (Camarhynchus spp.) on Floreana Island

Abstract Hybrid speciation is increasingly recognized as a mechanism for novel evolutionary trajectories. However, we know very little about the ecology of a contact zone that has arisen in sympatry. This study examines the foraging behavior and fitness of two species of Darwin’s tree finches (Camarhynchus parvulus, C. pauper) and hybrid offspring on Floreana Island. Previous study showed that the percentage of hybrids in the tree finch population increased from 19% in 2005 to 41% in 2010, and their body and beak size increased by ~5% (parental phenotype did not change). In 2005–2006, all three tree finch groups (two parental species and hybrid birds) used the same foraging substrate, technique, and height. By 2010–2013, the small tree finch C. par-vulus had changed its foraging technique and the medium tree finch C. pauper had changed its foraging height. Both parental species had higher body condition when foraging at (divergent) mean foraging heights per species but hybrid birds did not. We discuss the implications of conserving forest to facilitate vertical niche expansion and the role of hybridization for genetic persistence.

Invasive Parasites, Habitat Change and Heavy Rainfall Reduce Breeding Success in Darwin's Finches

Invasive alien parasites and pathogens are a growing threat to biodiversity worldwide, which can contribute to the extinction of endemic species. On the Galápagos Islands, the invasive parasitic fly Philornis downsi poses a major threat to the endemic avifauna. Here, we investigated the influence of this parasite on the breeding success of two Darwin's finch species, the warbler finch (Certhidea olivacea) and the sympatric small tree finch (Camarhynchus parvulus), on Santa Cruz Island in 2010 and 2012. While the population of the small tree finch appeared to be stable, the warbler finch has experienced a dramatic decline in population size on Santa Cruz Island since 1997. We aimed to identify whether warbler finches are particularly vulnerable during different stages of the breeding cycle. Contrary to our prediction, breeding success was lower in the small tree finch than in the warbler finch. In both species P. downsi had a strong negative impact on breeding success and our data suggest that heavy rain events also lowered the fledging success. On the one hand parents might be less efficient in compensating their chicks' energy loss due to parasitism as they might be less efficient in foraging on days of heavy rain. On the other hand, intense rainfalls might lead to increased humidity and more rapid cooling of the nests. In the case of the warbler finch we found that the control of invasive plant species with herbicides had a significant additive negative impact on the breeding success. It is very likely that the availability of insects (i.e. food abundance)is lower in such controlled areas, as herbicide usage led to the removal of the entire understory. Predation seems to be a minor factor in brood loss.

Species Collapse via Hybridization in Darwin’s Tree Finches

Species hybridization can lead to fitness costs, species collapse, and novel evolutionary trajectories in changing environments. Hybridization is predicted to be more common when environmental conditions change rapidly. Here, we test patterns of hybridization in three sympatric tree finch species (small tree finch Camarhynchus parvulus, medium tree finch Camarhynchus pauper, and large tree finch: Camarhynchus psittacula) that are currently recognized on Floreana Island, Galápagos Archipelago. Genetic analysis of microsatellite data from contemporary samples showed two genetic populations and one hybrid cluster in both 2005 and 2010; hybrid individuals were derived from genetic population 1 (small morph) and genetic population 2 (large morph). Females of the large and rare species were more likely to pair with males of the small common species. Finch populations differed in morphology in 1852-1906 compared with 2005/2010. An unsupervised clustering method showed (a) support for three morphological clusters in the historical tree finch sample (1852-1906), which is consistent with current species recognition; (b) support for two or three morphological clusters in 2005 with some (19%) hybridization; and (c) support for just two morphological clusters in 2010 with frequent (41%) hybridization. We discuss these findings in relation to species demarcations of Camarhynchus tree finches on Floreana Island.

Did tool-use evolve with enhanced physical cognitive abilities?

The use and manufacture of tools have been considered to be cognitively demanding and thus a possible driving factor in the evolution of intelligence. In this study, we tested the hypothesis that enhanced physical cognitive abilities evolved in conjunction with the use of tools, by comparing the performance of naturally tool-using and non-tool-using species in a suite of physical and general learning tasks. We predicted that the habitually tool-using species, New Caledonian crows and Galápagos woodpecker finches, should outperform their non-tool-using relatives, the small tree finches and the carrion crows in a physical problem but not in general learning tasks. We only found a divergence in the predicted direction for corvids. That only one of our comparisons supports the predictions under this hypothesis might be attributable to different complexities of tool-use in the two tool-using species. A critical evaluation is offered of the conceptual and methodological problems inherent in comparative studies on tool-related cognitive abilities.

Increasing avian pox prevalence varies by species, and with immune function, in Galápagos finches

Avian pox virus (APV), a pathogen implicated as a major factor in avian declines and extinctions in Hawaii, was introduced to the Galápagos in the late 1890s. While APV is thought to have increased in prevalence in recent years, no study has carefully evaluated the threat this pathogen poses to the Galápagos avifauna. In this paper, we examine the course of the APV epidemic in seven species of Galápagos finch on Santa Cruz Island (Geospiza fuliginosa, G. fortis, G. magnirostris, G. scandens, Camarhynchus parvulus, Cactospiza pallida, and Certhidea olivacea). We describe temporal changes in the prevalence of the avian pox disease (AP) caused by APV and the proportion of individuals that have recovered from AP from 2000 to 2009. Then we examine species differences in susceptibility to AP and how this variation correlates with differences in innate immune function. We show that AP prevalence has increased dramatically from 2000 to 2009. However, this increase in prevalence varied by species; specifically, we found that prevalence increased rapidly in G. fuliginosa, C. parvulus, G. scandens, and C. olivacea, but not at all in G. fortis. Furthermore, innate immune function varies between years and species, and this variation correlates with increased prevalence and species variation in susceptibility to APV. To our knowledge, this is the first study to demonstrate significant interannual variation in innate immune function in wild birds, and to show that this immune variation correlates with susceptibility to an introduced disease.

The Relationship Between Foraging, Learning Abilities and Neophobia in Two Species of Darwin’s Finches

AbstractThe ability to unlearn a previously established association is an important component of behavioural flexibility and may vary according to species ecology. Previously, two closely related sympatric Darwin’s finches were found to differ in their learning abilities. Small tree finches (Camarhynchus parvulus) outperformed woodpecker finches (Cactospiza pallida) in reversal learning but performed worse in an operant task. We attributed this difference to the habit of woodpecker finches to engage in long bouts of energetic pecking during extractive foraging. Persistently repeating one action without reward could favour performance in operant tasks but also limit behavioural flexibility. Here, we tested whether perseverance is the reason for woodpecker finches’ depressed reversal learning performance. Two new reversal conditions allowed the disentanglement of two sources of error in reversal learning: perseverant choice of the previously rewarded stimulus and failure to respond to the previously non‐rewarded stimulus. For the within‐species comparison, we predicted that woodpecker finches should find it more difficult to learn to avoid the previously rewarded stimulus than learning to choose the previously non‐rewarded stimulus. For the species comparison, we predicted the woodpecker finches should make more errors of perseverance than small tree finches. As performance could also be influenced by reaction to novelty, we compared neophobic responses between species and related them to reversal learning proficiency. We found no significant difference in reversal learning in the predicted direction, but found a negative correlation between neophobia and reversal learning at the inter‐ and the intraspecific level, which points towards a general relationship between reaction to novelty and flexibility.

Sometimes tool use is not the key: no evidence for cognitive adaptive specializations in tool-using woodpecker finches

The use and manufacture of tools has been considered to be cognitively demanding and thus a possible evolutionary driving factor of intelligence. Animal tool use provides the opportunity to investigate whether the use of tools evolved in conjunction with enhanced physical cognitive abilities. However, success in physical tasks may simply reflect enhanced general learning abilities and not cognitive adaptations to tool use. To distinguish between these possibilities, we compared general learning and physical cognitive abilities between the tool-using woodpecker finch, Cactospiza pallida , and its close relative, the small tree finch, Camarhynchus parvulus . Since not all woodpecker finches use tools, we also compared tool-using and nontool-using individuals, predicting that domain-specific experience should lead tool-using woodpecker finches to outperform nontool-users in a task that is similar to their natural tool use. Contrary to our predictions, woodpecker finches did not outperform small tree finches in either of the physical tasks and excelled in only one of the general learning tasks, and tool-using woodpecker finches did not outperform nontool-using woodpecker finches in the physical task closely resembling tool use. Our results provide no evidence that tool use in woodpecker finches has evolved in conjunction with enhanced physical cognition or that domain-specific experience hones domain-specific skills. This is an important contribution to a growing body of evidence indicating that animal tool use, even that which seems complex, does not necessitate specialized cognitive adaptations. ► Tool use may have evolved with enhanced cognitive abilities. ► Woodpecker finches did not outperform their nontool-using relatives in physical tasks. ► Nontool-using small tree finches made fewer errors and needed fewer trials in these tasks. ► Woodpecker finches were only better in an operant task that required perseverance. ► No indication that tool use evolved with enhanced cognitive abilities in this species.

Physical cognition and tool-use: performance of Darwin’s finches in the two-trap tube task

The trap tube is a classic test of causal reasoning abilities in animals in the physical domain. Recently, a modified version of this task improved its diagnostic capacity and allowed testing of non-tool-using animals. We used this modified two-trap tube task to compare the cognition of two Darwin's finch species: the woodpecker finch, Cactospiza pallida, a tool-using species, and the small tree finch, Camarhynchus parvulus, a closely related non-tool-using species. Not all woodpecker finches use tools in nature, and we therefore also tested non-tool-using individuals to assess the effect of experience on trap tube performance. No small tree finches and only two non-tool-using woodpecker finches solved the initial task which was operated using a pre-inserted piston. One tool-using woodpecker finch solved the task when allowed to use its own tool instead of the pre-inserted piston. The fact that none of these subjects transferred their knowledge when the features of the task changed, suggests that in this species, neither experience using tools nor the genetic composition of a tool-user are associated with the general physical cognitive skills required to solve the trap tube task.

Evaluating the safety of Rodolia cardinalis to two species of Galapagos finch; Camarhynchus parvulus and Geospiza fuliginosa

Some species of coccinellids exude a fluid that contains an alkaloid which can be toxic to vertebrates that ingest them. Because of this, before introducing Rodolia cardinalis to the Galapagos Islands as a biological control agent for the invasive scale insect, Icerya purchasi, it was considered necessary to determine whether it is toxic to birds. In this study, experiments were carried out with two common, insectivorous finches that are endemic to the Galapagos Islands; Camarhynchus parvulus (12 individuals) and Geospiza fuliginosa (16 individuals). Finches were maintained in captivity for 12–18 days. This period was divided into three phases: adaptation, experimental, and observation. During the experimental phase, test individuals, in addition to their regular diet, were fed 0.15 g portions of larvae or adults of R. cardinalis on alternate days over four days. The control group were fed 0.15 g of Asciodes quietalis abdomens in place of R. cardinalis. Symptoms of toxicity (mortality, deterioration, critical weight loss) from feeding on R. cardinalis were not observed in either species while in captivity, but birds avoided or rejected beetles indicating that they were distasteful. Finches regurgitated a higher number of larvae than adult beetles suggesting that larvae were less palatable than adults. Although long-term monitoring was not possible, results suggest that at least two species of finch will be unaffected by the release of R. cardinalis into the Galapagos Islands.

Male response to intruders is related to song characteristics in Darwin’s small tree finch (Camarhynchus parvulus)

Bird song functions in mate choice and species recognition, and hence variation in song can contribute to divergence and reproductive isolation. We used playback experiments to examine male response to conspecific song in Darwin’s small tree finch. Song is a reliable signal of bill morphology in this species, and individuals displayed stronger response to songs of males with similar bill size. These findings suggest that, in the context of territorial defence, males discriminate between intruders on the basis of song characteristics. Given that male response to song may be examined as a proxy for female response, this study also implies that females could discriminate between males on the basis of song. The findings suggest that: (1) perceived threat of intruders is related to reproductive competition and not fighting assessment, and (2) geographical isolation is not required for biologically meaningful song variation in Darwin’s finches.

Darwin's Finches as Seed Predators and Dispersers

We investigated seed predation and dispersal by nine Darwin's finch (Geospizinae) species, including species previously regarded as seed predators or insectivores. All nine ate fruit. Eight commonly discarded seeds, acting as short-distance dispersers. Seven defecated viable seeds, being therefore potential longer-distance dispersers. Fruit comprised the majority of dietary items in all four Geospiza species and Small Tree Finch (Camarhynchus parvulus). All except Common Cactus Finch (G. scandens) defecated intact seeds. The highest proportions of feces containing viable seeds were of Small Ground Finch (G. fuliginosa) and the “insectivorous” species Woodpecker Finch (Camarhynchus pallidus) and Warbler Finch (Certhidea olivacea). These two may be more important endochorous dispersers than other species that eat more fruit but are better seed predators. Intact seeds were found in 23% of fecal samples; 50% of the samples with intact seeds had viable seeds. The most frequently encountered intact seeds were of the tiny-seeded, fleshy-fruited Miconia robinsoniana and a sedge, Kyllinga brevifolia. Dry fruit, such as sedges and grasses, were usually crushed but occasionally found intact in feces, and might frequently be dispersed long distances. Seeds of larger-seeded, fleshy-fruited species were often discarded, but may be commonly dispersed short distances. Dispersal of plants by finches is probably more common than suggested by their predominant feeding patterns of seed predation and insectivory, and rare long-distance dispersal could be important for plant evolution and spread of introduced plants in the Galapagos Islands.

Philornis downsi parasitism is the primary cause of nestling mortality in the critically endangered Darwin’s medium tree finch (Camarhynchus pauper)

Darwin’s medium tree finch (Camarhynchus pauper) meets the 2009 International Union for Conservation of Nature Red List criteria for a critically endangered species because it has “a very small range on a single island” and is “declining rapidly owing to the effects of the parasite Philornis downsi”, habitat degradation, and introduced predators. The medium tree finch is only found in patches of remnant highland forest on Floreana Island, where it co-exists with breeding populations of small and large tree finches (C. parvulus and C. psittacula). Here, we examine the intensity of P. downsi in nests of small, medium, and large tree finches on Floreana. We expected that parasite intensity would increase with finch body size, and with greater rainfall, and would also correlate with increased nestling mortality. We found a trend in the expected direction for parasite intensity and rainfall. Combined meta-analytically with data from a previous study, the overall trend for the two studies was significant. We also found a significant linear trend in parasite intensity with finch body size. In addition, the medium tree finch exhibited a somewhat higher parasite intensity than would be expected based on body mass alone. Of 63 active medium tree finch nests, 17 nests had nestlings: all of which were infested with P. downsi. Only 25% of medium tree finch nestlings fledged, 28% were depredated, 41% died due to P. downsi parasitism, and 6% died for other reasons.

Mixed species nesting associations in Darwin's tree finches: nesting pattern predicts predation outcome

Predation is the main cause of passerine nesting failure. Traditionally, large intraspecific group size is thought to accrue individuals with fitness benefits from increased predator vigilance and hence lower predation risk. To date, few studies have investigated interspecific group size in relation to predation risk. In the present study, we examined predation outcome in Darwin's small tree finch, Camarhynchus parvulus, in nests with many or few interspecific neighbours. We tested the predictions: (1) nests in mixed associations have lower predation than do more solitary nests; (2) mixed species nesting associations covary with nest site vegetation characteristics; (3) older (i.e. presumably experienced) males more commonly nest in mixed associations than younger males; (4) older males select more concealed nesting sites; and (5) controlling male age, females prefer to pair with males in mixed associations than at solitary nests. Almost half of all nests occurred in mixed associations (46%) compared to solitary nests (54%), and the overall distribution of nests was decidedly nonrandom, displaying a bimodal pattern. Nest site vegetation characteristics of the focal species were inconsistently associated with nesting pattern, but older males did select more concealed nesting sites. Controlling differences in surrounding vegetation characteristics, mixed nesting associations experienced markedly lower predation than solitary nests, and females showed a preference for males in mixed associations, as demonstrated by higher male pairing success. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98, 313–324.

Bill Morphology Does Not Influence Vocal Performance in Darwin's Small Tree Finch on Floreana Island

Bird song is used for mate attraction and is important for establishing reproductive isolation. Current research highlights performance constraints on song production that may be influenced by variation in bill morphology. Darwin’s finches are a model system for studying the relationship between morphology and song performance, with previous studies suggesting that lower vocal performance is correlated with larger bill size. Here, we tested for a relationship between bill morphology and vocal performance in Darwin’s Small Tree Finch (Camarhynchus parvulus) on Floreana Island. We found no evidence of a correlation between bill morphology and vocal performance. This finding is in agreement with prior study of the Small Tree Finch, but contrasts a greater body of work addressing song in Darwin’s Finches. We discussed our findings in the context of ecological divergence, and ecological variation across species.

Love thy neighbour? Social nesting pattern, host mass and nest size affect ectoparasite intensity in Darwin’s tree finches

Social nesting behaviour is commonly associated with high prevalence and intensity of parasites in intraspecific comparisons. Little is known about the effects of interspecific host breeding density for parasite intensity in generalist host–parasite systems. Darwin’s small tree finch (Camarhynchus parvulus) on Santa Cruz Island, Galapagos Islands, nests in both heterospecific aggregations and at solitary sites. All Darwin finch species on Santa Cruz Island are infested with larvae of the invasive blood-sucking fly Philornis downsi. In this study, we test the prediction that total P. downsi intensity (the number of parasites per nest) is higher for nests in heterospecific aggregations than at solitary nests. We also examine variation in P. downsi intensity in relation to three predictor variables: (1) nest size, (2) nest bottom thickness and (3) host adult body mass, both within and across finch species. The results show that (1) total P. downsi intensity was significantly higher for small tree finch nests with many close neighbours; (2) finches with increased adult body mass built larger nests (inter- and intraspecific comparison); (3) parasite intensity increased significantly with nest size across species and in the small tree finch alone; and (4) nest bottom thickness did not vary with nest size or parasite intensity. These results provide evidence for an interaction between social nesting behaviour, nest characteristics and host mass that influences the distribution and potential impact of mobile ectoparasites in birds.

Jack-of-all-trades or master of one? Variation in foraging specialisation across years in Darwin’s Tree Finches (Camarhynchus spp.)

Theory predicts that species should have wider foraging niches during conditions of resource scarcity. However, empirical evidence to date has shown mixed patterns, including studies of Darwin’s finches that have found narrower foraging niches during conditions of resource scarcity. Here, we compare foraging behaviour in three species of Darwin’s Tree Finches (Camarhynchus spp.) in a dry versus wet year on Floreana Island to examine the change in foraging breadth under conditions of resource scarcity. We provide descriptive data on diet, foraging substrate, technique, height, and foraging time across the Small Tree Finch (C. parvulus), Medium Tree Finch (C. pauper), and Large Tree Finch (C. psittacula). During dry versus wet years, we made the following predictions: (1) lower intraspecific niche breadth (that is, a higher level of specialisation), (2) lower interspecific overlap in all foraging parameters, and (3) longer foraging times due to relative resource scarcity. Our findings showed that the Small and Medium Tree Finches were generalist foragers, while the Large Tree Finch was a specialist. Resource specialisation varied across years of differing rainfall: both generalist species were less specialised (higher Shannon diversity index) during the dry year, while the specialist species was more specialised (lower Shannon diversity index).

Nesting success in Darwin's small tree finch, Camarhynchus parvulus: evidence of female preference for older males and more concealed nests

Age-assortative mating has been documented among birds, but there is little evidence of direct benefits to females that pair with older males. In this study, I examined nest concealment in Darwin's small tree finch and tested the following predictions: (1) nest concealment increases with male age, (2) females prefer males that build well-concealed nests, and (3) females receive direct benefits through reduced predation at concealed nests. Darwin's small tree finch has a pattern of annual moult that makes it easy to observe age-related variation in reproductive success. In this species, males build a display nest that is sometimes used by the female for nesting. Controlling for male age, females preferred display nests for nesting that were well concealed by surrounding vegetation, whereas exposed display nests were rarely used for nesting (although occasionally a female paired with the male that built the unused display nest). Using cross-sectional and longitudinal data, this study found evidence of a female pairing preference for older males, as well as increased nest concealment among older males. Females that paired with older males had higher fledging success (using the longitudinal data), mostly due to reduced predation.

Assortative pairing and divergent evolution in Darwin’s Small Tree Finch, Camarhynchus parvulus

Size assortative mating has received increasing attention due to its potential to drive divergence and perhaps speciation. In this study, we examined assortative pairing at 17 nests of Darwin’s Small Tree Finches, Camarhynchus parvulus. We found positive assortative pairing for two traits: bill length and tarsus length, and these traits showed a significant positive correlation to each other. Assortative pairing could be driven by female choice for similar phenotypes because male–male competition has rarely been observed in the Small Tree Finch, nor have males been observed to reject potential mates. Given the high heritability of bill morphology in Darwin’s finches, it is possible that female preference for male bill length, a trait that is known to be important for foraging, will influence offspring bill size to maximise efficient exploitation of resources. The finding of size assortative pairing on the basis of tarsus length requires more research, but suggests different trait utilities for different foraging niches. Interestingly, the highland distribution of tarsus length across the population showed a unimodal distribution, but a bimodal distribution after pairing. While not significant, we found comparatively large differences across study plots in tarsus length, which suggests the possibility of phenotype–habitat matching at a small spatial scale in this species. Our findings are significant in the context of the adaptive radiation of Darwin’s finches as they are consistent with the allopatric model of speciation but also show potential for adaptive divergence in sympatry in Darwin’s tree finches.

Song is a reliable signal of bill morphology in Darwin's small tree finch Camarhynchus parvulus, and vocal performance predicts male pairing success

The shape and movement of the vocal tract are known to influence bird song. Current theory predicts that large bill and body size are correlated with low frequency song and slow trill rate. It is also widely accepted that song characteristics are important for mate choice by females. We investigated the relationship between bill morphology, song characteristics, and pairing success in Darwin's small tree finch Camarhynchus parvulus, on the Galapagos Islands. Contrary to predictions from a previous cross‐species study on Darwin's finches, we found that individuals with larger bill size produced songs with slow trill rate, high dominant frequency, and broad frequency bandwidth, indicating that song is a reliable signal of bill morphology. Vocal performance as indicated by the deviation from an upper performance limit was higher in paired than unpaired males. Pairing was not skewed in favour of a particular bill size, and both small and large billed males that sang high performance song had high pairing success. The reliable signalling function of song has implications for female choice and territorial defence, given that both females and conspecific competitors can assess the relative size of males’ bills through song, while females may use vocal performance as a signal of male quality.