Nest site selection of loggerhead sea turtles: The case of the island of Zakynthos, W Greece

The sandy beaches of Zakynthos Island support the largest single nesting aggregation in the Mediterranean Region of the endangered loggerhead turtle Caretta caretta. The present study attempts to determine possible correlations between a series of habitat variables and nest site selection. Nesting activities, including total and nesting emergences were examined in response to the recorded biotic and abiotic variables. The results of the analysis indicate that beach width is the most critical habitat variable affecting nest site selection. Further analysis of nesting performance implies that sea turtles use multiple environmental cues for nest site selection during the different steps of the nesting processes such as emergence from the surf and nesting. Nevertheless, we caution that a detailed study needs to be conducted over a more extensive period of time to verify these suggestions.     

Dyadic relationships in nesting males of the three-spined stickleback, Gasterosteus aculeatus

Multidimensional scaling analysis of nest site location, nesting interval, and aggressive and mating interactions revealed that male three-spined sticklebacks, Gasterosteus aculeatus, formed nesting clusters comprising five patterns of dyadic relationships. These patterns were defined by intervening variables such as size difference between the two males, nest site locations, synchronization of nesting cycles (egg presence in the nest), distance between two nests, and nest raiding behaviours. Several patterns of dyadic relationships were integrated among nesting males and their constellations indicated an expression of the social structure. Through an understanding the social structure, I described effects of social context or situations on individual reproductive success.     

Testing models of female reproductive migratory behaviour and population structure in the Caribbean hawksbill turtle, Eretmochelys imbricata, with mtDNA sequences

Information on the reproductive behaviour and population structure of female hawksbill turtles, Eretmochelys imbricata , is necessary to define conservation priorities for this highly endangered species. Two hypotheses to explain female nest site choice, natal homing and social facilitation, were tested by analyzing mtDNA control region sequences of 103 individuals from seven nesting colonies in the Caribbean and western Atlantic. Under the social facilitation model, newly mature females follow older females to a nesting location, and subsequently use this site for future nesting. This model generates an expectation that female lineages will be homogenized among regional nesting colonies. Contrary to expectations of the social facilitation model, mtDNA lineages were highly structured among western Atlantic nesting colonies. These analyses identified at least 6 female breeding stocks in the Caribbean and western Atlantic and support a natal homing model for recruitment of breeding females. Reproductive populations are effectively isolated over ecological time scales, and recovery plans for this species should include protection at the level of individual nesting colonies.     

Nest‐site selection by Interior Least Terns and Piping Plovers at managed,off‐channel sites along the Central Platte River in Nebraska,USA

Given the high productivity of Interior Least Terns (Sternula antillarum athalassos) and Piping Plovers (Charadrius melodus) on constructed off‐channel nesting sites along the central Platte River in Nebraska, USA, and the possibility of creating similar habitats at other locations in their breeding range, understanding how these species use off‐channel nesting habitats is important. We used data collected along the central Platte River in Nebraska, USA, over a 15‐year period (2001–2015), and a discrete‐choice modeling framework to assess the effects of physical site attributes and inter‐ and intraspecific associations on off‐channel nest‐site selection by Interior Least Terns and Piping Plovers. We found that Piping Plovers avoided nesting near each other, whereas colonial Interior Least Terns selected nest sites near those of conspecifics. In addition, the relative probability of use for both species was maximized when distance to the nearest predator perch was ≥ 150 m and elevation above the waterline was ≥ 3 m. Probability of use for nesting by Interior Least Terns increased as distance to water increased, whereas the probability of use by Piping Plovers was maximized when distance to water was ~50 m. Our results suggest that important features of constructed, off‐channel nesting sites for both species should include no potential predator perches within 150 m of nesting habitat and nesting areas at least 3 m above the waterline. Efficient site designs for Interior Least Terns would be circular, maximizing the area of nesting habitat away from the shoreline, whereas an effective site design for Piping Plovers would be more linear, maximizing the area of nesting habitat near the waterline. An efficient site design for both species would be lobate, incorporating centralized nesting habitat for Interior Least Terns and increased access to foraging areas for nesting and brood‐rearing Piping Plovers.     

Nesting bird species in sun coffee, pasture, and understory forest in southern Costa Rica

We investigated avian nest distribution and success in understoryforest, sun coffee plantations, and pasture in southern Costa Rica. Nestsearching occurred in plantations and forest in 1999 and 2000 and in pastures in2000. Nests were monitored until they failed or fledged young. Antbirds(Thamnophilidae) were the most common understory forest nesters and were notfound nesting in the plantations or pastures. Common nesting species in theplantations included Turdidae, Tyrannidae, Cardinalidae, and Thraupidae, many ofwhich are typical of forest edge/canopy or open, scrubby habitats. Two speciesassociated with forest interior, Henicorhina leucostictaand Buarremon brunneinucha, were found nesting in theplantations. Pastures supported similar types of nesting species as theplantations, with the exception of the forest-interior species. Daily mortalityrates (DMRs) for above-ground cup-nesting species in plantations and pastureswere similar to those for species nesting in forest at our site and a site inPanama. The results indicate that conversion from forest to pastures and suncoffee plantations diminishes nesting habitat for forest-interior species, whilenumerous forest edge/canopy species and open-country species are able to nest inthese agricultural land-cover types. As a group, species nesting in theplantations and pastures do not have unusually high nest mortality rates,although species-specific studies are lacking. Nesting species distributionsacross habitat types and DMRs at our study site may be influenced by the largeamount of forest in the landscape.     

Variation of mitochondrial DNA D-loop sequences in the endangered Siberian crane Grus leucogeranus Pallas

To evaluate inter- and intra-populational divergence of the endangered Siberian crane a comparative analysis of mtDNA D-loop region (490 bp) sequences was performed for 17 captive cranes including 14 originating from the eastern and 3 from the central nesting area. Thirteen variable sites form 9 haplotypes, 6 belonging to the eastern nesting site individuals, 2 belonging to the central nesting site and one being common for both nesting areas. The haplotype network analysis did not show significant separation of these two groups. The data obtained could provide genetic grounds for the programs of restoration of the central nesting area population by captive breeding and reintroduction to the wild using the eastern nesting site population gene pool. These results could also be a basis for further evaluation of Siberian crane populations genetic divergence needed for its adequate protection.     

Effects of colony nesting of Adrea cinerea and Egretta alba modesta on soil properties and herb layer composition in a Pinus densiflora forest

Effects of colony nesting of herons on soil properties and herb layer composition in Pinus densiflora forest were studied at Pomaeri, Kangwon Province in Korea. Herons have used this habitat as a nesting site from January to October every year. In 1995, more than 500 herons were observed in this habitat. Nutrient content of soil was much higher at the nesting site than that of the non-nesting site (control). Total nitrogen concentration of soil at the nesting site and the control site was 14.8±1.85 mg g^-1 and 2.8±0.35 mg g^-1, respectively. Phosphorus content of soil in the nesting site was 32 times greater than that of the control site. This is evidently due to the addition of feces of the herons, and decomposition of thin twigs and organic debris from the canopy of dead trees and bird nests. Light intensity at herb layer of the nesting site and of the control site was 80% and 20%, respectively, of incident on outside forest. Species diversity of herb layer in the nesting site (9 species) was quite lower than that in the control site (14 species). Similarity index of the herb layer between the two sites was 0.07. The nesting site was dominated by indicator species of soil eutrophication such as Humulus japonicus, Persicaria perfoliata, Persicaria fauriei, Commelina communis, Chelidonium majus var. asiaticum. Changes of herb species composition in the nesting site was evidently due to the eutrophication of the soil and increased light intensity of the herb layer.     

THE BREEDING OF THE LESSER FLAMINGO IN THE MWERU WANTIPA, NORTHERN RHODESIA.

A hitherto unreported breeding site of the Lesser Flamingo Phoeniconaias minor is recorded in the Mweru Wantipa or Mweru Marsh, Northern Rhodesia. The site of the nesting colony, its structure, nest construction and behaviour of the birds is described. The importance of flooding in determining the site of the nesting colony is noted; adverse conditions caused by a rise in the water-level of Lake Mweru Wantipa appear to have completely destroyed the nests. The instability of such environments does not offer much chance of regular and permanent breeding, and may influence local movements on the part of the birds.     

Nest boxes for Cape Parrots Poicephalus robustus in the Hogsback area,Eastern Cape,South Africa

Breeding propensity of tree-cavity nesting bird species are often limited by a shortage of natural nesting sites. Artificial nests can be used to provide alternative nest sites. Cape Parrots Poicephalus robustus are nationally endangered and nest in existing tree-cavities in high-altitude fragmented Afromontane forests in South Africa, assumed to be in short supply due to historic and current logging practices. To increase nest site availability, 179 wooden bird boxes and 28 bee boxes (to ‘pull’ bees) were erected during 2011–2012 in Hogsback, Eastern Cape. In 2016, no bird boxes were occupied by Cape Parrots. A total of 43% were used by other species, 51% were unused and 6% could not be inspected due to tree instability and inaccessibility. Two bird boxes were inspected by two pairs of Cape Parrots, but were never occupied. Occupancy of boxes by birds was not associated with nest, tree or habitat characteristics. However, occupancy of boxes by bees was associated with habitat type and tree species. Future conservation efforts will include locating natural Cape Parrot nesting sites and reforestation efforts to ensure the long-term availability of natural nesting sites.     

Genetic structure at three spatial scales is consistent with limited philopatry in Ricord's Rock Iguanas (Cyclura ricordii)

Cyclura ricordii is an endemic iguana from Hispaniola Island and is threatened on the IUCN Red List. The main threats are predation by introduced mammals, habitat destruction, and hunting pressure. The present study focused on two nesting sites from Pedernales Province in the Dominican Republic. The hypothesis that natal philopatry influences dispersal and nest‐site selection was tested. Monitoring and sampling took place in 2012 and 2013. Polymorphic markers were used to evaluate whether natal philopatry limits dispersal at multiple spatial scales. Ripley's K revealed that nests were significantly clustered at multiple scales, when both nesting sites were considered and within each nesting site. This suggests a patchy, nonrandom distribution of nests within nest sites. Hierarchical AMOVA revealed that nest‐site aggregations did not explain a significant portion of genetic variation within nesting sites. However, a small but positive correlation between geographic and genetic distance was detected using a Mantel's test. Hence, the relationship between geographic distance and genetic distance among hatchlings within nest sites, while detectable, was not strong enough to have a marked effect on fine‐scale genetic structure. Spatial and genetic data combined determined that the nesting sites included nesting females from multiple locations, and the hypothesis of “natal philopatry” was not supported because females nesting in the same cluster were no more closely related to each other than to other females from the same nesting site. These findings imply that nesting aggregations are more likely associated with cryptic habitat variables contributing to optimal nesting conditions.     

Competition, predation and nest niche shifts among tropical cavity nesters: phylogeny and natural history evolution of parrots (Psittaciformes) and trogons (Trogoniformes)

Nest site selection by birds is a critically important life history trait as competition for suitable sites can be intense, and because birds are at their most vulnerable to predators during nesting. Previous studies show that the clutch size and nestling period evolve in response to competition for nest sites and nest predation, respectively. This provides the opportunity to study the relative contribution of competition and predation to the evolution of nesting niche. Using previously published phylogenies for parrots and trogons, I found evidence for at least 13 independent evolutionary transitions from tree cavities to alternative nesting niches (including termitaria, cliffs, and burrows). I analyzed variations in clutch size, incubation period and nestling period for 16 phylogenetically controlled pairs of species to test the relative roles of competition for tree cavities and nest predation, in favoring evolutionary switches to alternative nest sites. Tree cavity nesting species did not have larger clutch sizes as predicted if competition for tree cavities leads birds to invest heavily in nesting once they obtain a nest site (the limited breeding opportunities hypothesis). Instead I found that shifts to alternative nesting niches were accompanied by an increase in nestling period. As nestling period is a surrogate measure for long-term nest predation rates, this finding suggests that nest predation has been more important than competition in niche diversification among cavity nesting parrots and trogons. The timing of events in South America suggests that the explosive radiation of mammalian nest predators during the Upper-Oligocene, Lower-Miocene (20–30 million years ago) corresponded with the radiation of parrot and trogon taxa that exploit novel nesting niches.     

Relationship between tropical cyclones and the distribution of sea turtle nesting grounds

Aim This study examines the relationship between the distribution of existing sea turtle nesting sites and historical patterns of tropical cyclone events to investigate whether cyclones influence the current distribution of sea turtle nesting sites. The results, together with information on predicted cyclone activity and other key environmental variables, will help in the identification and prediction of future nesting sites for sea turtles as changes to the coastal environment continue. Location Queensland, Australia. Methods We used data on the nesting distribution of seven populations of four species of sea turtles [green (Chelonia mydas), flatback (Natator depressus), hawksbill (Eretmochelys imbricata) and loggerhead (Caretta caretta)] from the eastern Queensland coast, and tropical cyclone track data from 1969 to 2007 to explore the relationship between (1) sea turtle nesting phenology and cyclone season, and (2) sea turtle nesting sites and cyclone distribution. Furthermore, using two green turtle populations as a case study, we investigated the relationship between cyclone disturbance and sea turtle reproductive output, nesting site and season. Bootstrapping was used to explore if current sea turtle nesting sites are located in areas with lower or higher cyclone frequency than areas where turtles are currently not nesting. Results All populations of sea turtles studied here were disturbed by cyclone activity during the study period. The exposure (frequency) of tropical cyclones that crossed each nesting site varied greatly among and within the various sea turtle populations. This was mainly a result of the spatial distribution of each population’s nesting sites. Bootstrapping indicated that nesting sites generally have experienced lower cyclone activity than other areas that are available for nesting. Main conclusions Tropical cyclones might have been sufficiently detrimental to sea turtle hatching success on the eastern Queensland coast that through a natural selection process turtles in this region are now nesting in areas with lower cyclone activity. Therefore, it is important that future studies that predict climate or range shifts for sea turtle nesting distributions consider future cyclone activity as one of the variables in their model.     

Conspecific attraction and nest site selection in a nomadic species,the zebra finch

Conspecific nesting density affects many aspects of breeding biology, as well as habitat selection decisions. However, the large variations in breeding density observed in many species are yet to be fully explained. Here, we investigated the settlement patterns in a colonial species with variable breeding density and where resource distribution could be manipulated. The zebra finch, Taeniopygia guttata, is a classic avian model in evolutionary biology but we know surprisingly very little about nest site selection strategies and nesting densities in this species, and in fact, in nomadic species in general. Yet, important determinants of habitat selection strategies, including temporal predictability and breeding synchrony, are likely to be different in nomadic species than in the non‐nomadic species studied to date. Here, we manipulated the distribution of nesting sites (by providing nest boxes) and food patches (feeders) to test four non‐exclusive habitat selection hypotheses that could lead to nest aggregation: 1) attraction to resources, 2) attraction to breeding conspecifics, and 3) attraction to successful conspecifics and 4) use of private information (i.e. own reproductive success on a site). We found that wild zebra finches used conspecific presence and possibly reproductive success, to make decisions over where to locate their nests, but did not aggregate around water or food within the study areas. Moreover, there was a high degree of inter‐individual variation in nesting density preference. We discuss the significance of our results for habitat selection strategy in nomadic species and with respect to the differential selection pressures that individuals breeding at different densities may experience.     

Nesting biology of Black-shouldered Kite (Elanus caeruleus) in oil palm landscape in Carey Island,Peninsular Malaysia

Black-shouldered Kite (Elanus caeruleus) is a well-known raptor that inhabits open areas such as oil palm plantation or paddy field. To determine preferable habitat and nesting site of Black-shouldered Kite in oil palm landscape, we conducted a study on Black-shouldered Kite’s nesting biology in Carey Island, Selangor, Malaysia. We divided the island into six types of habitat and conducted road-side count of Black-shouldered Kite from April 2009 to February 2011. Whenever the Black-shouldered Kite was detected, we thoroughly searched the surrounding area for their nest. In total, we have recorded forty nests. The nests were built on 15 species of trees but most of the trees shared common physical characteristics. Some novice breeders also used oil palm tree as their nesting site. Structure and building materials of nests constructed on oil palm trees were different from nests built on other trees. Of all breeding attempts, only four nests which were located in residential area adjacent to young oil palm habitat were successful. Among important characteristics of successful nesting site include taller trees with strong branches and good leaf coverage. These trees not only protect nests from predator detection (except from other predatory birds) and physical environment but also facilitate Black-shouldered Kite’s foraging activities by providing good vantage point.     

Honey bee (<Emphasis Type="Italic">Apis mellifera capensis/A. m. scutellata</Emphasis> hybrid) nesting behavior in the Eastern Cape,South Africa

Little is known about the natural history of wild honey bee (Apis mellifera) colonies in the Eastern Cape Province of South Africa. The goal of this research was to examine nest site characteristics of honey bee (A. m. capensis/A. m. scutellata hybrid) colonies sampled from a variety of habitats (nature reserves, livestock farms, and an urban setting) in the Eastern Cape. We also determined how nest site location related to various colony strength parameters. In general, colonies not nesting in ground cavities tended to nest in locations >6 m high when nesting in cliffs and buildings and >2 m high when nesting in trees. Colonies typically nested in cavities whose entrances faced a southeasterly direction and were ~40 L in volume. We sampled a subset of colonies to determine the relationship between nest type and the following colony strength parameters: total area of comb in the colony, the volume of stored honey, pollen, and brood, adult bee population, the weight per adult bee, and the bee/nest cavity volume ratio. In general, colonies nesting in cliffs tended to be stronger than those nesting in the ground or trees. Our findings provide new insights into the nesting biology of honey bees in the Eastern Cape, South Africa, perhaps leading to the formation of conservation recommendations for honey bees in this region.     

Effects of management on double-crested cormorant nesting colony fidelity

The increase of double-crested cormorant (Phalacrocorax auritus; hereafter, cormorant) populations during the last 2 decades has impacted many stakeholder groups. The negative effects of nesting cormorants on trees and other vegetation have motivated private organizations and government agencies to manage nesting colonies and reduce their impacts to private property and public resources. Management-induced reproductive failure has been shown to influence cormorant inter-annual nesting colony fidelity, but not complete abandonment from a nesting colony site. We attached very high frequency (VHF) transmitters and Global Positioning System (GPS) transmitters to nesting cormorants to monitor their movement response on a managed site (Young Island, VT [YI]) and an unmanaged site (Four Brothers Islands, NY [FB]). Additionally, we monitored these sites to determine the influence of management activities on subsequent-year colonization. On YI, management consisted of egg-oiling all cormorant nests (some nests had been oiled in previous years) and culling approximately 20% of adults. Annual dispersal rates did not differ between managed and unmanaged sites, but a nesting period interaction occurred with greater dispersal on the managed site following the incubation period. After 4 years of both egg oiling and culling, cormorant nesting on YI declined to zero. Simultaneously, cormorant numbers increased on the nearby unmanaged FB. We propose either the cumulative effect of partial or complete reproductive failure (8 yr) or simply the inclusion of adult culling (4 yr) caused the abandonment. From a colony-specific management perspective, the rapid decline was beneficial to the goal of restoring the vegetative community on YI. The effects of adult culling at nesting colonies, prior-year reproductive failure caused by egg oiling, or the combination of these factors may be required for complete and rapid nesting site abandonment. The use of culling adult breeders reduced nesting and likely limits the cost and logistics of control and allows more rapid initiation of mitigation measures and island habitat restoration. © 2011 The Wildlife Society.     

Use of Restored Riparian Habitat by the Endangered Least Bell's Vireo (Vireo bellii pusillus)

A primary objective of riparian restoration in California is the creation of habitat for endangered species. Four restoration sites in San Diego County were monitored between 1989 and 1993 and evaluated for their suitability as nesting habitat for Vireo bellii pusillus (Least Bell's Vireo), a state and federally endangered obligate riparian breeder. Vegetation structure at each site was quantified annually and compared to a model of canopy architecture derived from Least Bell's Vireo territories in natural habitat. Vireo use of restored habitat was documented through systematic surveys and nest monitoring. By 1993, only one site in its entirety met the habitat suitability criteria of the model, but portions of each site during all years did so. Differences between sites in the time required to develop suitable habitat—well-developed layered vegetation from the ground to under 8m in height)—were attributable largely to variation in annual rainfall. Vireos visited restoration sites to forage as early as the first growing season, but they did not establish territories or nest there until at least part of the site supported suitable habitat as determined from the model. Placement of territories and nests coincided with patches of dense vegetation characteristic of natural nesting areas. Occupation of restored sites was accelerated by the presence of adjacent mature riparian habitat, which afforded birds nest sites and/or foraging habitat lacking in the planted vegetation. Vireos nesting in restored habitat achieved success comparable to that of vireos nesting in surrounding natural habitat, and there was no evidence that productivity was reduced in created areas. These findings indicate that creating nesting habitat for this target species is feasible and suggest that the critical components of vireo nesting habitat have been captured in both the design and quantitative assessment of restoration sites.     

Nesting Behavior of <Emphasis Type="Italic">Gorilla gorilla diehli</Emphasis> at Kagwene Mountain,Cameroon: Implications for Assessing Group Size and Density

We recorded nesting data at 569 fresh night nest sites, comprising 7032 individual nests, of Cross River gorillas inhabiting the Kagwene Mountain in western Cameroon. The mean night nest group size was 12.4. Overall, 55% of night nests were constructed on the ground and 45% in trees. Significantly more arboreal nests were constructed in the wet season (69%), vs. the dry season (19%). Day nest construction was common at Kagwene (n = 260 nest sites, mean nest group size = 5.98) and we encountered significantly more day nest sites in the wet season. Nest site reuse was also common (35%), though not related to season. Our results of nesting habits concur with those from other western gorilla studies, in which rainfall influences arboreal nesting. However, we encountered wet season arboreal nesting, day nest construction, and overall nest site reuse more frequently than reported for other sites. Our observations have considerable implications when estimating group size and density using traditional nest count data. The gorillas at Kagwene inhabit the highest altitudinal range of all Cross River gorilla subpopulations and rainfall is also high; therefore other subpopulations may demonstrate different nesting characteristics. However, one should consider our findings when attempting to estimate Cross River gorilla density at other localities through nest site data.     

Soil temperature, digging behaviour, and the adaptive value of nest depth in South American species of Acromyrmex leaf-cutting ants

In leaf-cutting ants, workers are expected to excavate the nest at a soil depth that provides suitable temperatures, since the symbiotic fungus cultivated inside nest chambers is highly dependent on temperature for proper growth. We hypothesize that the different nesting habits observed in Acromyrmex leaf-cutting ants in the South American continent, i.e. superficial and subterranean nests, depend on the occurrence, across the soil profile, of the temperature range preferred by workers for digging. To test this hypothesis, we first explored whether the nesting habits in the genus Acromyrmex are correlated with the prevailing soil temperature regimes at the reported nest locations. Second, we experimentally investigated whether Acromyrmex workers engaged in digging use soil temperature as a cue to decide where to excavate the nest. A bibliographic survey of nesting habits of 21 South American Acromyrmex species indicated that nesting habits are correlated with the soil temperature regimes: the warmer the soil at the nesting site, the higher the number of species inhabiting subterranean nests, as compared to superficial nests. For those species showing nesting plasticity, subterranean nests occurred in hot soils, and superficial nests in cold ones. Experimental results indicated that Acromyrmex lundi workers use soil temperature as an orientation cue to decide where to start digging, and respond to rising and falling soil temperatures by moving to alternative digging places, or by stopping digging, respectively. The soil temperature range preferred for digging, between 20°C and maximally 30.6°C, matched the range at which colony growth would be maximized. It is suggested that temperature-sensitive digging guides digging workers towards their preferred range of soil temperature. Workers’ thermopreferences lead to a concentration of digging activity at the soil layers where the preferred range occurs, and therefore, to the construction of superficial nests in cold soils, and subterranean ones in hot soils. The adaptive value of the temperature-related nesting habits, and the temperature-sensitive digging, is further discussed.     

Nesting Behaviour of the Giant Honeybees <Emphasis Type="Italic">Apis</Emphasis><Emphasis Type="Italic">dorsata</Emphasis> Occurring in Jhargram,West Bengal,India

The giant honeybees Apis dorsata are habituated to construct combs in trees, houses, caves as well as in overhead water reservoir occurring in their nesting localities. To verify their preference for nesting sites if any, surveys were conducted in recent years (2013–16), during nesting seasons of these bees in Jhargram area of West Bengal, India. It is revealed that A. dorsata construct their combs in big, tall, aged simul (Bombax ceiba), bot (Ficus benghalensis) trees mostly, irrespective of localities. Also they were seen to construct nest at the underside of the overhead water reservoir ignoring nesting potential trees occurring nearby. Of course, nesting in the houses, and on the walls of culvert is not uncommon. As the bees constructed more than 100–200 nests at the same nesting site e.g., a tree or/and an overhead water reservoir, depending upon the availability of space for construction of nest it is concluded that these insects prefer colonial nesting.