The role of urban greenspaces in fauna conservation: the case of the iguana Ctenosaura similis in the ‘patios’ of León city, Nicaragua

Private urban greenspaces, called ‘patios’ in Latin America, can act as important refuges for wildlife in scattered growing cities of the tropics. We studied the presence and abundance of the black spiny-tailed iguana (Ctenosaura similis) in patios of León city (Nicaragua). Forty patios were structurally characterized and abundances of iguanas were determined through surveys of inhabitants supported by observations of specimens and burrows. Patio area and maximum tree height were variables positively related with presence and abundance of iguanas and presence of burrows. The permeability of fences for iguana movements and the presence of preferred trees for food were also related to presence and abundance of iguanas. Stepwise selection model for abundance of iguanas included number of preferred trees, maximum tree height and permeability of fences. The presence of iguanas was only explained by maximum tree height. Our results show that the indigenous-root types of patios offer the most adequate conditions for the establishment and maintenance of iguana populations among the studied cases. In the light of these results, management practices to preserve the Ctenosaura similis populations in urban patios are suggested.     

Anti-predator adaptations in nesting Marsh Tits Parus palustris: the role of nest-site security

A proposal that nest predation is the most important selective pressure shaping nest-site use of Marsh Tits Parus palustris was checked in a long-term study (almost 500 nests) carried out in the Białowieża National Park (eastern Poland) in primeval conditions, where the birds breed in natural holes and their nests are at risk from a variety of predators. It was predicted that predation rates would depend on hole attributes, so that Marsh Tits should use the most secure holes. Predation was responsible for 70% of total nest losses. Predation risk depended on hole attributes. Nests in dead wood were predated significantly more often than those in live wood. Nests in old woodpecker holes were predated more frequently than nests in holes of other origin, and nests situated closer to the entrance were more at risk than ones further from it. The entrance size did not influence overall predation risk, but small entrance size was important in preventing access by larger predators. These differences can account for the very rare use of holes in dead wood or of woodpecker holes, the fact that they nest in holes with small entrances and relatively far from the entrance. It is concluded that the patterns of nest-site use found in this species are best explained as anti-predator adaptations, which have evolved and are maintained by the pressure of nest predators. This study also indicates the possible limits of, and constraints on, these adaptations.     

Ultraviolet light and heat source selection in captive spiny-tailed iguanas (Oplurus cuvieri)

Three experimental manipulations were conducted to assess the influence of heat source selection and active thermoregulation on ultraviolet (UV) light exposure in captive spiny-tailed iguanas (Oplurus cuvieri) at the Jersey Wildlife Preservation Trust. Four replicates per manipulation were conducted on six individual lizards. All animals were tested in a separate enclosure to which they were acclimated before observations. Data on choice of thermal sources were collected during the first 2 hr of light, when lizards were actively thermoregulating. Animals were allowed to choose between incandescent light, UV light and a non-light heat source (thermotube) in different combinations. Recorded temperatures close to the incandescent light (37°C) were always significantly higher than at the thermotube (33°C) and at the UV light (29°C). Manipulation 1 offered the animals a choice of an UV light and an incandescent light as thermal sources. Manipulation 2 presented animals with the thermal choices in Manipulation 1, but substrates under each source in Manipulation 1 were switched. In Manipulation 3, animals could choose between an incandescent light and the thermotube. All studied lizards were significantly more attracted to the incandescent light than to the UV light or thermotube. Incandescent light elicited a significantly higher proportion of basking behaviors in all individuals than the other sources. A high proportion of time basking was also spent in front of the thermotube but fewer individuals and less time were spent basking under the UV light. Heat source selection was generally found to be independent of substrate. Management applications of this preference are suggested for juvenile diurnal heliothermic iguanids. Zoo Biol 16:391–401, 1997. © 1997 Wiley-Liss, Inc.     

Microhabitat Selection in Diet and Trophic Ecology of a Spiny-Tailed Iguana Ctenosaura hemilopha

Annual variation in the diet of adult spiny-tailed iguanas was studied by analyzing feces collected at 38 known iguana refuges in the desert of the southern Baja California peninsula. Iguanas consumed mostly plants, a small percentage of arthropods, their own skin, and remains from conspecifics. They consumed at least 22 of 83 plant species known to be present in the area. Leaves of three perennial Leguminosae species formed the year-round basis of their diet. Iguanas ate flowers and fruit when available. We propose that iguanas could be efficient seed dispersers for Cactaceae, but that they crush many of the seeds of leafy tree species. No significant differences in food consumption pattern were found between dry and rainy seasons. Iguanas apparently moved to neighboring trees only when searching for food. We propose that predation risk could have more influence than the nutritional value of plants on the species' foraging behavior and microhabitat selection in this area.     

Escape Behavior and Predation Risk of Mainland and Island Spiny-tailed Iguanas (Ctenosaura hemilopha)

We investigated the relationships between predator avoidance behavior and predation risk by comparing the wariness of iguanas (Ctenosaura hemilopha) belonging to an island population with few predators with that of iguanas belonging to a mainland population under high predation pressure. We predicted that island iguanas would be less wary than mainland ones. Island iguanas allowed the closer approach of potential predators before their first reaction and fleeing. The responses of both sexes did not differ statistically, except for the frequent displays shown by males as the first reaction. These results suggest that island iguanas, which were probably introduced onto the island by man, have modified their behavior in response to a decline in predation risk, supporting the hypothesis that predation pressure can contribute to intraspecific variation in antipredator behavior. The results also quantify the tameness of prey on islands without terrestrial predators, and their consequent vulnerability if foraneous predators are introduced. No significant difference was found between the populations in the time needed by frightened iguanas to resume their normal activity, suggesting that the island iguanas could learn to recognize predators and develop a suitable antipredator behavior.     

Temporal dynamics of woodpecker predation on emerald ash borer (Agrilus planipennis) in the northeastern U.S.A.

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Predominance of single paternity in the black spiny-tailed iguana: conservation genetic concerns for female-biased hunting

Because of female-biased illegal harvesting, knowledge about the genetic mating system of the black spiny-tailed iguana Ctenosaura pectinata is of primary interest for the conservation of this threatened species. Based on the high levels of multiple paternity found in clutches of many other reptiles, particularly in lizards, it is hypothesised that multiple paternity may also be common in black iguanas. This was investigated by using microsatellite DNA to estimate the number of males siring nine litters (9 mothers, 121 offspring genotyped at ten polymorphic loci) of black iguanas. Contrary to expectations, only 11% of sampled black iguana females produced litters consistent with being sired by multiple males. These data are the first evidence for the predominance of single paternity within an iguanid lizard, and suggest that black iguana may be more susceptible to loss of genetic variation in the face of gender-biased over-hunting pressure than previously thought.     

Behavioral,physiological, and morphological components of dominance and mate attraction in male green iguanas

This study investigated the morphological, physiological, and behavioral components of social dominance important for mate attraction in male green iguanas (Iguana iguana). A group of 9 male and 11 female adult green iguanas was studied in a large semi-natural enclosure during one reproductive season (October–January). Four of the nine males never initiated aggressive encounters; the other five were observed to display aggressively toward each other and were ranked in a linear dominance hierarchy. Head size was the most important factor influencing fighting success. Head size and display frequency were positively correlated with plasma testosterone levels. Dominance rank directly influenced ability to monopolize areas containing resources used by females. The quality of a male's home range, measured as his access to a large basking rock in the enclosure, was related to the proportion of potential mates found within his home range. One male greatly surpassed the others in his ability to defend a home range of high quality and attract potential mates. These data suggest that physiological and morphological factors, through their influence on social behavior, may ultimately affect male reproductive fitness. © 1992 Wiley-Liss, Inc.     

Behavioral and physiological adjustments to new predators in an endemic island species, the Galápagos marine iguana

For the past 5 to 15 million years, marine iguanas (Amblyrhynchus cristatus), endemic to the Galápagos archipelago, experienced relaxed predation pressure and consequently show negligible anti-predator behavior. However, over the past few decades introduced feral cats and dogs started to prey on iguanas on some of the islands. We investigated experimentally whether behavioral and endocrine anti-predator responses changed in response to predator introduction. We hypothesized that flight initiation distances (FID) and corticosterone (CORT) concentrations should increase in affected populations to cope with the novel predators. Populations of marine iguanas reacted differentially to simulated predator approach depending on whether or not they were previously naturally exposed to introduced predators. FIDs were larger at sites with predation than at sites without predation. Furthermore, the occurrence of new predators was associated with increased stress-induced CORT levels in marine iguanas. In addition, age was a strong predictor of variation in FID and CORT levels. Juveniles, which are generally more threatened by predators compared to adults, showed larger FIDs and higher CORT baseline levels as well as higher stress-induced levels than adults. The results demonstrate that this naive island species shows behavioral and physiological plasticity associated with actual predation pressure, a trait that is presumably adaptive. However, the adjustments in FID are not sufficient to cope with the novel predators. We suggest that low behavioral plasticity in the face of introduced predators may drive many island species to extinction.     

Thermal ecology and behaviour of the nomadic social forager Malacosoma disstria

The present study examines whether the nomadic social caterpillar Malacosoma disstria Hübner (Lepidoptera: Lasiocampidae) can thermoregulate despite the lack of a tent, and evaluates the role of thermoregulation in directing the colony's behaviour. The presence of a radiant heat and light source (i.e. a lamp in the laboratory experiments and the sun in the field observations) enables caterpillar colonies to increase body temperature by basking (remaining still under a heat source) and this is only effective when caterpillars cluster in groups. Body temperatures achieved when basking in a group coincide with the temperatures at which the development rate is maximal for this species. Indeed, in the laboratory experiments, the presence of a lamp results in higher growth rates, confirming that thermoregulation is an advantage to group living. When a radiant heat/light source is provided at a distance from the food in the laboratory, caterpillars behave to maximize thermal gains: colonies move away from the food to bivouac (i.e. group together and remain still on a silk mat) under the lamp, spend more time on the bivouac and cluster in a more cohesive group. Thermal needs thus influence habitat selection and colony aggregation. Malacosoma disstria relies on developing rapidly, despite low seasonal temperatures, aiming to benefit from springtime high food quality and low predation rates; however, unlike others in its genus, it does not build a tent but instead exhibits collective nomadic foraging (i.e. the whole colony moves together between temporary resting and feeding sites). In this species, collective thermoregulatory behaviour is not only possible and advantageous, but also drives much of the colony's behaviour, in large part dictating the temporal and spatial patterns of movement. These findings suggest that thermoregulation may be an important selection pressure keeping colonies together.     

Behavioural Response of Bats to Perceived Predation Risk While Foraging

The ability to detect and respond to predation risk while foraging may have important fitness consequences for prey organisms. Anti‐predator behaviours may reduce the probability of mortality because of predation, but they may also be associated with reduced foraging efficiency. Several behaviours of bats have been suggested to serve as anti‐predator responses, and there is evidence that predation, particularly by avian predators such as owls, may be an important cause of bat mortality. Previous studies have attempted to determine whether predator presence affects bat behaviour when emerging from roost sites, but few have examined effects of predator presence on bat behaviour while foraging. In this study, we investigated whether foraging bats respond to predator cues by presenting bats with an acoustic cue simulating the presence of an owl. Within matched trials, which were conducted at different locations each of 18 nights, significantly fewer bat detections were recorded at owl playback stations than at control stations (no auditory cue), suggesting an avoidance response by bats. An acoustic control (i.e. station playing woodpecker calls), however, did not have significantly more detections than the stations playing the owl calls, suggesting that bats may simply be avoiding noise and more detailed investigation is warranted. Although evidence for owl predation on bats is minimal in North America, results of this study may indicate that the perceived presence of owls may represent a factor influencing the behaviour of bats while foraging.     

Predation‐associated divergence of male genital morphology in a livebearing fish

Male genital morphology is remarkably diverse across internally fertilizing animals, a phenomenon largely attributed to sexual selection. Ecological differences across environments can alter the context of sexual selection, yet little research has addressed how this may influence the rapid, divergent evolution of male genitalia. Using the model system of Bahamas mosquitofish (Gambusia hubbsi) undergoing ecological speciation across blue holes, we used geometric morphometric methods to test (i) whether male genital shape (the small, approximately 1 mm long, distal tip of the sperm‐transfer organ, the gonopodium) has diverged between populations with and without predatory fish and (ii) whether any observed divergence has a genetic basis. We additionally examined the effects of genetic relatedness and employed model selection to investigate other environmental factors (i.e. interspecific competition, adult sex ratio and resource availability) that could potentially influence genital shape via changes in sexual selection. Predation regime comprised the most important factor associated with male genital divergence in this system, although sex ratio and some aspects of resource availability had suggestive effects. We found consistent, heritable differences in male genital morphology between predation regimes: Bahamas mosquitofish coexisting with predatory fish possessed more elongate genital tips with reduced soft tissue compared with counterparts inhabiting blue holes without predatory fish. We suggest this may reflect selection for greater efficiency of sperm transfer and fertilization during rapid and often forced copulations in high‐predation populations or differences in sexual conflict between predation regimes. Our study highlights the potential importance of ecological variation, particularly predation risk, in indirectly generating genital diversity.     

The effect of the thermal environment on the ability of hatchling Galapagos land iguanas to avoid predation during dispersal

Summary During the month of February 1979, several hundred hatchling land iguanas (Conolophus pallidus) were observed emerging from their natal burrows in a 2 ha communal nesting area on Isla Santa Fe, Galapagos Islands. During this emergence, as many as nine Galapagos hawks were observed to patrol the nesting area and attack hatchling iguanas.The hypothesis that the ability of hatchling land iguanas to escape predation could be influenced by the interaction of the physiological state of the lizards and the thermal environment was analyzed using (1) empirical data on the effect of body temperature (T _b) on locomotory ability of iguanas and (2) biophysical modeling of the T _b's of hatchlings under natural conditions. This hypothesis was tested by assessing the success of natural hawk attacks on lizards exposed to different thermal environments.During those periods when predicted T _b's of hatchlings were always <32°C, (at which temperatures land iguanas were shown to have less than maximal ability to sprint rapidly) hawks were successful in 67% of the observed attacks. However, when T _b's of hatchlings were always 32° C, hawks were successful on only 19% of observed attacks. During periods when hatchling T _b's could be <32° C or 32–40° C (depending upon which microhabitat the hatchling occupied before the attack), the hawks were successful in 46% of the observed attacks.These data indicate that the physical environment, as mediated through the physiological state of the lizards and to correlated locomotary abilities, significantly affects the ability of hatchling land iguanas to escape predation.     

On the phylogenetic position of the Okinawa woodpecker (Sapheopipo noguchii)

The Okinawa woodpecker Sapheopipo noguchii is the rarest extant woodpecker species. The monotypic genus Sapheopipo was considered to be a representative of an old lineage of woodpeckers that led to the Eurasian genera Picus and the Blythipicus–Gecinulus species. This view, based on similarities in color patterns, external morphology and foraging behavior, has been adopted in all major accounts of the family. The alternative view, that this woodpecker may be related to the widespread white-backed woodpecker Dendrocopos leucotos, which evolved distinctive subspecies on other East Asian islands, has not been generally accepted. We analyzed partial sequences of the mitochondrial cytochrome b (cyt b) gene to test these hypotheses. The data suggest that the Okinawa woodpecker is a member of the genus Dendrocopos, with white-backed woodpecker and great spotted woodpecker D. major as close relatives. Color patterns support the genetic results and indicate a closer relationship with the white-backed woodpecker. Consequently, the correct taxonomic designation of the Okinawa woodpecker would be Dendrocopos noguchii (Seebohm in Ibis 5(5):173–182, 1887) in the tribe Campetherini rather than Picini.This revised version was published online in February 2005 with corrections to tables 1 and 2.     

Effects of intraguild predators on nest-site selection by prey

Nest-site selection involves tradeoffs between the risk of predation (on females and/or nests) and nest-site quality (microenvironment), and consequently suitable nesting sites are often in limited supply. Interactions with “classical” predators (e.g., those not competing for shared resources) can strongly influence nest-site selection, but whether intraguild predation also influences this behavior is unknown. We tested whether risk of predation from an intraguild predator [the diurnal scincid lizard Eutropis (Mabuya) longicaudata] influences nest-site selection by its prey (the nocturnal gecko Gekko hokouensis) on Orchid Island, Taiwan. These two species putatively compete for shared resources, including invertebrate prey and nesting microhabitat, but the larger E. longicaudata also predates G. hokouensis (but not its hard-shelled eggs). Both species nested within a concrete wall containing a series of drainage holes that have either one (“closed-in”) or two openings (“open”). In allopatry, E. longicaudata preferred to nest within holes that were plugged by debris (thereby protecting eggs from water intrusion), whereas G. hokouensis selected holes that were open at both ends (facilitating escape from predators). When we experimentally excluded E. longicaudata from its preferred nesting area, G. hokouensis not only nested in higher abundances, but also modified its nest-site selection, such that communal nesting was more prevalent and both open and closed-in holes were used equally. Egg viability was unaffected by the choice of hole type, but was reduced slightly (by 7%) in the predator exclusion area (presumably due to higher local incubation temperatures). Our field experiment demonstrates that intraguild predators can directly influence the nest density of prey by altering maternal nest-site selection behavior, even when the predator and prey are active at different times of day and the eggs are not at risk of predation.     

Vigilance against predators induced by eavesdropping on heterospecific alarm calls in a non-vocal lizard Oplurus cuvieri cuvieri (Reptilia: Iguania)

Prey animals can reduce their risk of predation by detecting potential predators before encounters occur. Some animals gain information about nearby predators by eavesdropping on heterospecific alarm calls. Despite having well-developed ears, most lizards do not use vocal information for intraspecific communication, and few studies have shown practical use of the ears in wild lizards. Here, we show that the Madagascan spiny-tailed iguana (Oplurus cuvieri cuvieri) obtains auditory signals for predator detection. The Madagascan spiny-tailed iguana and the Madagascar paradise flycatcher (Terpsiphone mutata) are syntopic inhabitants of the Ampijoroa dry deciduous forest of Madagascar. The iguana and the flycatcher have neither a predator–prey relationship nor resource competition, but they have shared predators such as raptors and snakes. Using playback experiments, we demonstrated that the iguana discriminates mobbing alarm calls of the flycatcher from its songs and then enhances its vigilance behaviour. Our results demonstrate the occurrence of an asymmetrical ecological relationship between the Madagascan spiny-tailed iguana and the paradise flycatcher through eavesdropping on information about the presence of predators. This implies that indirect interspecific interactions through information recognition may be more common than generally thought in an animal community.     

CONSISTENCY AND VARIATION IN PHENOTYPIC SELECTION EXERTED BY A COMMUNITY OF SEED PREDATORS

Phenotypic selection that is sustained over time underlies both anagenesis and cladogenesis, but the conditions that lead to such selection and what causes variation in selection are not well known. We measured the selection exerted by three species of predispersal seed predators of lodgepole pine (Pinus contorta latifolia) in the South Hills, Idaho, and found that net selection on different cone and seed traits exerted by red crossbills (Loxia curvirostra) and cone borer moths (Eucosma recissoriana) over 10 years of seed crops was similar to that measured in another mountain range. We also found that the strength of selection increased as seed predation increased, which provides a mechanism for the correlation between the escalation of seed defenses and the density of seed predators. Red crossbills consume the most seeds and selection they exert accounts for much of the selection experienced by lodgepole pine, providing additional support for a coevolutionary arms race between crossbills and lodgepole pine in the South Hills. The third seed predator, hairy woodpeckers (Picoides villosus), consumed less than one‐sixth as many seeds as crossbills. Across the northern Rocky Mountains, woodpecker abundance and therefore selective impact appears limited by the elevated seed defenses of lodgepole pine.     

Behavioral and physiological adjustments to new predators in an endemic island species,the Galápagos marine iguana

For the past 5 to 15 million years, marine iguanas (Amblyrhynchus cristatus), endemic to the Galápagos archipelago, experienced relaxed predation pressure and consequently show negligible anti-predator behavior. However, over the past few decades introduced feral cats and dogs started to prey on iguanas on some of the islands. We investigated experimentally whether behavioral and endocrine anti-predator responses changed in response to predator introduction. We hypothesized that flight initiation distances (FID) and corticosterone (CORT) concentrations should increase in affected populations to cope with the novel predators. Populations of marine iguanas reacted differentially to simulated predator approach depending on whether or not they were previously naturally exposed to introduced predators. FIDs were larger at sites with predation than at sites without predation. Furthermore, the occurrence of new predators was associated with increased stress-induced CORT levels in marine iguanas. In addition, age was a strong predictor of variation in FID and CORT levels. Juveniles, which are generally more threatened by predators compared to adults, showed larger FIDs and higher CORT baseline levels as well as higher stress-induced levels than adults. The results demonstrate that this naive island species shows behavioral and physiological plasticity associated with actual predation pressure, a trait that is presumably adaptive. However, the adjustments in FID are not sufficient to cope with the novel predators. We suggest that low behavioral plasticity in the face of introduced predators may drive many island species to extinction.     

Crown dieback events as key processes creating cavity habitat for magellanic woodpeckers

Abstract Woodpeckers are considered keystone species for webs of cavity nesters and habitat and resource specialists that strongly depend on availability of trees suitable for cavity excavation. Most studies carried out in northern hemisphere temperate coniferous forests emphasize the importance of old growth stages of forests or large dead trees as habitat for cavity builders. We present a study of Nothofagus pumilio tree selection by the magellanic woodpecker (Campephilus magellanicus) that incorporates dendroecological data on long‐term growth trends of trees that provides new insights into the processes that create suitable habitat for cavity excavating species. We analysed 351 cavity and neighbouring control trees in terms of age and radial growth patterns, as well as external tree characteristics. In addition, from a subsample of these trees we developed tree‐ring chronologies for each group using standard methods in order to analyse potential differences in radial growth patterns between cavity and non‐cavity trees. Multivariate models that account for differences between paired cavities versus control trees indicated that growth decline and the degree of crown dieback were the primary variables explaining magellanic woodpecker tree selection for cavity building. In contrast to previous work, neither diameter (above a certain threshold) nor age, were important determinants of selection. Furthermore, trees that became present cavity are those that had synchronously declined in radial growth during the 1943–44 and 1956–57 droughts and the 1985–86 massive caterpillar defoliation. Insect outbreaks and extreme climatic events may episodically reduce vigour, induce partial crown mortality, trigger increased fungal attack and heart rot formation at different tree heights on the bole in a group of trees and thus increase availability of soft substrate and their likelihood of cavity excavation by the magellanic woodpecker. These results underscore the importance of drought/biotically‐induced canopy dieback events in creating habitat for woodpeckers and their dependent cavity users.     

Nest-site selection and its impact on breeding success of the Lesser Kestrel,Falco naumanni,in the North of Iran (Aves: Falconiformes)

The Lesser Kestrel (Falco naumanni) is a small migratory falcon which nests coloni- ally on inland cliffs and ruined buildings. Bandar Turkman city holds one of the most important breeding colonies in Iran. It nests in holes in roofs, walls or on ledges of buildings throughout of the city. We tried to find out whether there is a relationship between nest-site selection and breeding success. Although height from the ground is the most important factor in nest-site selection by F. naumanni, height from the floor and distance to the nearest neighbour are other main factors which affect breeding success. This could be the result of accessibility by human and natural predators such as domestic cats which threaten nests. The number of possible nest-sites in the study area is limited as many old, ruined buildings are being replaced by new construc- tions; this colonially breeding bird species therefore suffers from a lack of highly suitable nest-sites and this directly affect breeding success.