Nesting behavior and nest site selection in monk parakeets (Myiopsitta monachus) in the Pantanal of Brazil

We examined nesting behavior in monk parakeets (Myiopsitta monachus) in their native habitat in the Brazilian Pantanal. Unique among parrots, monk parakeets build communal nest structures that contain many cavities, each belonging to an individual pair. We studied 41 parakeet colonies that had 104 nest structures. We hypothesized that nest structures would be located in trees providing the greatest support and protection from predators and inclement weather, and that nest sites and nest cavities would differ from random locations with respect to tree characteristics, location of houses, and presence of jabiru stork (Jabiru mycteria) nests, as suggested anecdotally by other authors. Fewer than half of the colonies were close (<350 m) to houses. There was a strong association with jabiru storks; 21 of 23 stork nests had monk parakeet nest cavities attached, accounting for 51% of parakeet colonies. Of the 21 jabiru-associated colonies, 6 had additional parakeet structures and 15 had only the jabiru-attached parakeet structure. Monk parakeet colonies associated with jabiru nests had significantly more nesting cavities than did monospecific monk parakeet colonies, due mainly to those attached directly to the jabiru nest. In jabiru-associated colonies, parakeet nest structures were located higher and in taller trees than in monospecific colonies. There was no difference in trunk diameters of parakeet nesting trees with or without jabirus. Although we tabulated 24 tree species as nest trees, nearest-neighbor trees, or matched-point trees, only 6 species were used for nesting. When compared to matched points, monk parakeet structures were preferentially located in piuva (Tabebuia spp.) and mandovi trees (Sterculia apetela). Parakeet structures were in taller trees with thicker trunk diameters than matched points. Most nest cavities (71%) faced in a northerly direction (northwest to northeast), away from cold southerly winds. Choice of a nest site and orientation appears to reflect structural, weather, and predator constraints. Thick, tall trees with stout branches provided stable sites for their large nest structures, which are known to collapse because of their own weight and strong winds. Nesting with jabirus confers structural advantages (they could attach many nests to the bottom of the jabiru nest, potentially gaining benefits from social facilitation), early warning, and predator defense.     

Rediscovery of the workerless inquiline antPogonomyrmex colei and additional notes on natural history (Hymenoptera: Formicidae)

Summary Pogonomyrmex colei is a workerless inquiline ant known only from nests ofP. rugosus, its closest relative. Ten of 776 (1.3%) host nests were parasitized at a site in central Arizona, while none of 1499 potential host colonies were parasitized at two other locales. Colonies ofP. colei are perennial, and host alate females in 9 of 10 colonies demonstrates that host queens survive parasitism. Three of 10 colonies died over 19 colony years of observation, while only 1 of 601 colonies became newly parasitized. Mating occurs in morning for up to 2–3 days following summer and fall rains and in afternoon during cool fall days. Mating is intranidal just outside the nest entrance, with males returning to the natal nest. MaleP. colei may be flightless because their wing area is reduced compared to host males. Females fly from the nest and locate potential host colonies by following trunk trails. Workers are the largest barrier to nest establishment, as they removed over 90% ofP. colei females placed in trunk trails or that entered host nests. Males and females ofP. colei andP. anergismus, the only other congeneric inquiline species, are diminutive compared to their hosts, with females 30% lighter than host workers. Fat content is lower and water content is higher inP. colei andP. anergismus females than in their hosts.     

Ontogenetic changes in forager polymorphism and foraging ecology in the leaf-cutting ant Atta cephalotes

In the leaf-cutting ant Atta cephalotes (L.) small colonies produce a relatively narrow size-range of small workers, whereas large colonies produce a much wider size-range of workers. In this study, I compared the foraging of four small A. cephalotes colonies (fewer than 5000 workers) with published data on foraging of large colonies to examine how colony size and worker size-range may be related to foraging ecology in leaf-cutting ants. I found that the foraging ecology of small A. cephalotes colonies is very different from that of large colonies. In small colonies, a relatively narrow size-range of foragers (1.4–6.7 mg, mean 3.3 mg) cut primarily herbs (ferns, grasses, and other small herbaceous plants) located within 7 m of the nest. In contrast, in large colonies, a broader size-range of workers (1.4–30 mg, mean 7.3 mg) participate in foraging, generally harvesting from trees 20–80 m from the nest, with larger workers cutting on trees with thicker and tougher leaves. Small colonies' dependence on small herbaceous plants near the nest may have a profound impact on distribution of A. cephalotes. A. cephalotes colonies are rarely found in primary forest, where the low occurrence of small herbaceous plants in the understory may preclude the establishment of young colonies.     

Old comb for nesting site recognition by <Emphasis Type="Italic">Apis dorsata?</Emphasis> Field experiments in China

The Asian giant honey bee, Apis dorsata, often conducts seasonal, long-distance migrations in southern China, between a preferred tree (having more than one nest) and alternate sites. Although worker bees cannot make a round-trip journey, colonies re-utilize preferred trees after an absence of several months. We performed comb experiments in which bases and all abandoned combs were entirely scraped off trees and their sites covered with plastic, or comb was moved to trees of the same species. Swarms of giant honey bees investigated trees where combs were removed and continued to nest on the same trees. In contrast, placing combs in nets on previously used trees, or on nearby trees of the same species, did not attract more swarms. The same number of colonies that left them returned to previously occupied trees. Our findings suggest that direct olfactory or sensory contact with old comb bases might regulate nest establishment, but individual trees, lacking normal visual or chemical cues of old nests, are relocated using behavioral devices that remain to be elucidated. Received 12 February 2007; revised 5 June 2007; accepted 13 September 2007.     

Trail following and stowaway behaviour of the myrmecophilous staphylinid beetle,Homoeusa acuminata,during foraging trips of its hostLasius fuliginosus (Hymenoptera: Formicidae)

Summary The behaviour of adultHomoeusa acuminata on trails of its hostLasius fuliginosus was investigated both in the field and in the laboratory. The beetles were active from May to September, accurately following the foraging trails of their hosts up to 20 metres from the nest. Most of the time, they were ignored by the ants, but if attacked they raised their abdomen as a possible appeasement or defensive behaviour. On trails the beetles most probably act as food robbers, feeding on prey collected by ants. The following method, called stowaway behaviour, was used by the beetles: when a beetle encountered an ant carrying a prey back to the nest it jumped on the prey, probably feeding on it while being transported.Laboratory experiments on circular artificial trails demonstrated thatH. acuminata follows a water extract of hindguts of the ants, the source of the trail pheromone. Both beetles and ants responded to an artificial trail of 0.03 hindgut equivalent per cm, but the mean distance followed by the beetles was about twelve times higher than that covered by the ants themselves. In contrast, experiments with solutions of the six fatty acids reported as the active components of the trail pheromone showed that the beetles did not respond at all, and that the ants only respond to the fatty acids at a very high concentration.     

Seasonal spatial dynamics and causes of nest movement in colonies of the invasive Argentine ant (Linepithema humile)

Abstract.  1. Colony organisation and movement behaviour of the Argentine ant ( Linepithema humile ) was studied over 3 years in field populations in California and in captive colonies in the laboratory. This invasive species is highly polydomous and unicolonial; colonies consist of expansive and fluid networks of nests and trails. The spatial and temporal organisation of colonies may contribute to ecological dominance.
2. Argentine ant nests and inter-nest trails shift in size, abundance, and location, so that colony networks are spatially contracted in the winter and expanded spring to autumn. Colonies occupy permanent sites; ants migrated to and from the same winter nest locations year after year, and occupied 30% of the same nests repeatedly during seasonal migrations.
3. Nests were moved on average 2–3 m. Forty-two per cent were occupied less than 1 month, 4% the entire study, and the other 54% lasted 3.9 ± 2.3 months (mean ± SD).
4. Nests were located within 2–4 m of woody plants, in warm sites in the winter and cool sites in the summer. Both humidity and food availability influenced nest-site choice in laboratory colonies. However, when faced with a trade-off between factors, the ants chose humid nest boxes over nest boxes near food, and ants moved nests only in response to changes in humidity and not distance to food.
5. The results indicate that L. humile colonies are seasonally polydomous, and that nest movements are driven by changes in microclimate. Colony organisation maintains high local density and increases food supply, which may improve the competitive ability of L. humile colonies and reduce opportunities for species coexistence.     

The self-organizing exploratory pattern of the argentine ant

Workers of the Argentine ant, Iridomyrmex humilis,start to explore a chemically unmarked territory randomly. As the exploratory front advances, other explorers are recruited and a trail extends from it to the nest. Whereas recruitment trails are generally constructed between two points, these exploratory trails have no fixed destination, and strongly resemble the foraging patterns of army ants. A minimal model shows how the exploratory pattern may be generated by the individual workers' simple trail-laying and -following behavior, illustrating how complex collective structures in insect colonies may be based on self-organization.     

Trail system in red wood ants (<Emphasis Type="Italic">Formica rufa</Emphasis> Group) under recreation press

The influence of recreation press on the forager trail system in red wood ants was examined. The model settlement of these ants was situated in green moss-pine forest, comprising areas with a different degree of the recreation press. In 2007, trail systems of 36 anthills were mapped (a total of 136 trails). In 2009, two nests from each group at each digression stage were selected as model objects. These nests possessed close to average values of the cupola diameter (d) and the number of trophic trails (n _c ). In these 6 model nests, trail systems (a total of 33 trails) were re-mapped and the strength of columns was estimated. It had been found that the recreation press decreases the population number in families of red wood ants. Trophic trails become shorter, their structure is simplified, and the number of trees with aphid colonies decreases. When the size of the family decreases owing to irregular decrease in the population number of separate columns, columns possessing non-ramified (less “efficient”) trails disappear first. Weakening of the family decreases the mobility of the trail network; trees with aphid colonies become more important.     

Nest demographics and foraging behavior of <Emphasis Type="Italic">Apterostigma collare</Emphasis> Emery (Hymenoptera,Formicidae) provide evidence of colony independence

Apterostigma collare Emery is a highly derived fungus-growing ant within the Tribe Attini whose small, fungal nests are found in tropical rain forests. This study focuses on determining the colony structure of A. collare, specifically searching for evidence of polydomy or independence. We surveyed and observed nests in the field, and performed foraging bioassays and dissected nests in the laboratory. We determined the size and contents of nests in field populations. Nests found near other nests were not statistically different in size compared to nests found alone. There was also no statistical difference between near and lone nests regarding the presence of a queen in the nest. Most nests contained one queen with brood and workers, regardless of their proximity to other nests. Observations also were made of foraging and trail-marking behaviors. Foraging activity observed in the field revealed that workers left the nest area and followed trails upwards into the canopy, but they did not interact with foragers from other nearby nests. In a laboratory foraging arena, foragers marked a trail to a food source by dragging the gaster. Bioassays showed that A. collare workers preferred their own foraging trails, but not those of other conspecific colonies. All results suggest that each nest represents an independent colony, supporting a previous report that nests found in close proximity do not constitute a polydomous colony. Received 19 July 2006; revised 23 March 2007; accepted 6 June 2007.     

The size–distance relationship in the wood ant Formica rufa

1. The size–distance relationship among honeydew‐collecting foragers of the red wood ant Formica rufa was investigated. Within the colony territory, the size (as measured by head width) and fresh weight of samples of foragers were determined for ants ascending and descending trees near, and farther from, the central nest mound. 2. The mean size of the ants was significantly higher at far trees than at near trees in six out of the seven colonies investigated, confirming the general presence of the size–distance relationship. 3. In three colonies, a load–distance relationship was also found. For a given head width, honeydew‐carrying ants descending far trees were significantly heavier than those descending near trees (i.e. they were carrying heavier loads from trees farther away from the central nest mound). 4. This is the first time that both load–distance and size–distance relationships have been reported in foraging workers from the same ant colony. 5. The combined effects of these characteristics suggest that colony foraging efficiency is enhanced by far trees being visited by the larger workers that then return with heavier loads of honeydew.     

Intercolonial differences in raiding activities in the Japanese slave-making antPolyergus samurai

Summary Intercolonial differences in raiding activity were investigated in the field on 9 colonies of the slavemaking antPolyergus samurai. Duration of raiding season, the mean start time of the first raiding trip in a day, the mean speed of outbound and inbound trips, and the mean distance to target nests varied significantly among colonies. The variance in start time of raiding was correlated with that in soil temperature and diurnal change pattern in soil temperature at the nest sites. The speed of trips and the first day with trips in the season also correlated with soil temperature at the nest site. Simple environmental factors well explained the observed variances in raiding activity among colonies. Therefore, raiding behaviour ofP. samurai seemed to be a stereotyped behaviour that is regulated through simple environmental factors.     

Colony structure and spatial distribution of food resources in the polydomous meat ant <Emphasis Type="Italic">Iridomyrmex purpureus</Emphasis>

Polydomous social insects may reduce the costs of foraging by the strategic distribution of nests throughout their territory or home-range. This efficiency may most likely be achieved if the resources are relatively stable in place and time, and the colonies and nests are distributed in response to the location of the resources. However, no study has investigated how the distribution of food sources influences the spatial patterns of nests within polydomous colonies under natural conditions. Our two year study of 140 colonies of the Australian ant Iridomyrmex purpureus revealed that the decentralization of nests within colonies is associated with the distribution of trees containing honey-dew producing hemiptera. We show there is a positive correlation between the maximum distance between trees containing hemiptera and the maximum distance between nests within a colony. In addition, we demonstrate the mechanism by which this pattern may arise; new nests are built nearer to trees containing hemiptera than existing nests. Further, the distance between trees containing hemiptera and the nearest nests was negatively correlated with the length of exploitation of that tree. Finally, we show that most food is delivered to the nearest nest after which other ants redistribute it between the nests. Combined, these data suggest that foraging efficiency may be an important selection pressure favouring polydomy in I. purpureus. Received 6 April 2006; revised 29 September; accepted 4 October 2006.     

A shift in colony founding behaviour in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2

Summary. A shift in colony founding behaviour from single queen (haplometrosis) to multiple queens (pleometrosis) was observed locally in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2, which is associated with Macaranga trees in Borneo. In addition, about a quarter of all mature colonies (27 of 95 trees examined) were found to be multiple queen colonies. They arose either directly from pleometrotic founding colonies or secondarily by adoption of additional queens. Using microsatellite markers, we showed that queens in colonies founded through pleometrosis are unrelated and each queen participates in producing worker offspring, albeit with significant skew in a third of the colonies. In mature polygynous colonies, all resident queens contributed to the production of workers and sexual offspring. Relatedness of queens in mature polygynous colonies was not significantly higher than in foundress associations. We hypothesize that increased nest site limitation in this specific interaction trigger the observed shift in colony founding behaviour. Crematogaster msp. 2 inhabits the light demanding pioneer plant species Macaranga pearsonii that is typical for early successional stages of secondary forests. Thus suitable host-plants for colonisation are abundant for only a short time in highly disturbed sites and become increasingly sparse when secondary forest matures.Received 6 September 2004; revised 29 November; accepted 10 December 2004.     

Food and reproductive strategies of the myrmecophilous staphylinid beetleHomoeusa acuminata (aleocharinae) on the foraging trails of its host antLasius fuliginosus (Hymenoptera: Formicidae)

Summary The trail following behaviour of the adult myrmecophilous beetleHomoeusa acuminata on the foraging trails on the host antLasius fuliginosus was examined in June and all the observed behaviours were quantified. The beetle appears as a food robber, using the stowaway behaviour which consists of attaching itself to prey transported by ants to the nest. Foraging trails are also used by the beetles as a meeting place for mating. However, the beetle does not enter into the nest and appears as a symbiont poorly integrated in the social life of its host.     

Dispersal of Eucalyptus torelliana seeds by the resin-collecting stingless bee,Trigona carbonaria

Ants are the only group of invertebrates currently identified as significant dispersers of seeds, but we report here the dispersal of Eucalyptus torelliana seeds by bees. Fruits of E. torelliana produce resin which is collected by workers of the stingless bee Trigona carbonaria. Seeds adhere to resin in the workers' corbiculate and are transported to the nest. Workers transported seeds distances of more than 300 m from the parent tree and seeds at the nest were viable and capable of germination. Seeds were removed from the nests by workers and discarded away from the nest, and E. torelliana trees became established in the vicinity of colonies of T. carbonaria. Mellitochory may be a novel method of seed dispersal where bees are dispersers, and associated with fruits that produce resin as an attractant for bees.     

Intraspecific interactions in a community of arboreal nesting termites (Isoptera: Termitidae)

In coconut plantations of northern New Guinea, the arboreal nesting termite community comprises three species:Nasutitermes princeps, N. novarumhebridarum, andMicrocerotermes biroi. In orde to assess the importance of intraspecific interactions in this community, we conducted pairwise encounters between batches of individuals in the laboratory and between entire nest populations in seminantural conditions. Three levels of agonism were defined in laboratory bioassays: anagonism, moderate agonism, and strong agonism. Anagonism was observed during all control tests with homocolonical groups and in some tests with allocolonial groups of all species. Moderate agonism included initial aggressiveness that subsequently faded out, and initially passive encounters where aggression progressively built up and led to fighting. Strong agonism corresponded to initial aggressiveness and fighting. Results obtained in alboratory bioassays were consistent with bioassays in seminatural conditions. WhenNasutitermes colonies were anagonists in laboratory bioassays, their foraging trails merged without aggression in field tests.N. princeps nests that were moderately agonistic in laboratory tests fought and either continued to avoid each other or finally joined after elimination of the most aggressive individuals. The most aggressiveM. biroi andN. princeps colonies fought and their foraging trails diverged afterward. Direct attacks on alien nests were winnessed inM. biroi. In all species, anagonism occurred in 21–34% of the combinations tested, between either geographically close or distantcolonies. An exeption was a group of 112 anagonist nests ofN. princeps, which most probably constituted a supercolony. The level of agonism betweenNasutitermes colonies was constant during the wet and dry season. Termite colonies excluded each other, both intra- and interspecifically, from the coconut trees, and their territories seem distributed in a mosaic pattern. Agonism between colonies may result in the elimination of the weakest colonies or in trail divergence, maintaining this mosaic. In cotrast, lack of agonism between some colonies suggests the possibility of colony fusion and gene exchanges without nuptial flights.     

Suburban sprawl: environmental features affect colony social and spatial structure in the black carpenter ant,Camponotus pennsylvanicus

1. In social insects, the number of nests that a colony inhabits may have important consequences for colony genetic structure, the number of queens, sex allocation, foraging efficiency, and nestmate recognition. Within the ants, colonies may either occupy a single nest (monodomy) or may be organised into a complex network of nests and trails, a condition known as polydomy. 2. The current study is a large‐scale, long‐term, comprehensive field examination of various features of colony social and spatial structure in the facultatively polydomous black carpenter ant, Camponotus pennsylvanicus (DeGeer). The study examined the density, persistence, and the spatiotemporal distribution of colonies across a gradient of land disturbance associated with urban development. The temporal and spatial pattern of nest use was compared between fragmented landscapes where nesting sites were interspersed among human‐built structures (urban plots) and less disturbed landscapes with higher tree density (suburban plots). In addition, nesting site fidelity and changes in colony spatial structure were monitored over 7 years. 3. Long‐term monitoring and extensive sampling over a large spatial area allowed the first comprehensive insight into the spatiotemporal dynamics of colony and population structure in C. pennsylvanicus. A total of 1113 trees were inspected over 233 ha. Camponotus pennsylvanicus were active on 348 of the 1113 trees (31%) and these represented 182 distinct colonies. The colonisation rate remained relatively stable over 7 years suggesting that an equilibrium point had been reached. Relative to the suburban plots, tree density was 65% lower in the urban plots. The proportion of trees colonised by C. pennsylvanicus was significantly higher in the urban plots suggesting that intraspecific competition for nesting sites may be especially high in areas with lower tree density. Colony spatial structure also differed significantly between habitats and a higher incidence of monodomy was observed in the urban environment. The average number of trees per colony across all subplots was 1.95 (range 1–4) indicating that C. pennsylvanicus are weakly polydomous. 4. The composite picture that emerges for C. pennsylvanicus colonies in the urban habitat is a chain reaction of events: (i) the urban habitat has a lower tree density, (ii) lower tree density results in higher tree colonisation rate, (iii) higher tree colonisation rate results in simpler colony spatial structure (i.e. higher incidence of monodomy), and (iv) simpler colony spatial structure results in numerically smaller colonies. Long‐term monitoring of the spatiotemporal pattern of nest site use in selected colonies revealed a unique trend. While worker counts in selected colonies remained relatively stable throughout the course of the study, colony spatial structure changed considerably with 28% of colonies experiencing a change. Furthermore, the likelihood of detecting a change in colony spatial structure increased with the amount of time passing from the initial inspection. 5. In conclusion, tree density has a significant effect on a number of important colony features in C. pennsylvanicus. Besides tree density, other environmental features such as human‐built structures cause habitat fragmentation and may act as natural barriers to worker dispersal and/or foraging. Such barriers may ultimately affect the social and/or spatial structure at both the colony and the population level.     

Acanthocephalan (Oligacanthorhynchidae) parasitism of the Caribbean termite <Emphasis Type="Italic">Nasutitermes acajutlae</Emphasis>: implications for reproductive success

Summary Few studies have examined the impact of parasitism on free-living social insects. We documented the association between an acanthocephalan parasite and survival, reproduction and growth of a Caribbean termite (Nasutitermes acajutlae). We measured these parameters yearly in 100 termite colonies for 4 years. In 2001 and 2002 we also compared the rate at which parasitized and unparasitized colonies rebuilt damaged foraging trails because trails protect foragers from predation as well as fluctuations in temperature and humidity. Although there were no significant differences in growth or survival between parasitized and unparasitized colonies, parasitized colonies reproduced significantly less often and rebuilt foraging trails significantly more slowly than unparasitized colonies. The average parasitized colony may sustain a loss of alate production =30% in the year in which it is parasitized, and a loss of up to 19% if it is parasitized once in a 5-year period. We conclude that the acanthocephalan parasite is associated with significant alteration of fitness parameters of individual N. acajutlae colonies. However, prevalence of parasitism is low (6.8%) and alate production by non-parasitized colonies is likely to compensate for losses due to parasitism. Therefore, it seems unlikely that this parasite affects N. acajutlae at the population level.Received 5 October 2003; revised 19 November 2003; accepted 3 December 2003.     

Chemical trail systems,orientation, and territorial interactions in the antLasius neoniger

Foraging and territoriality in the ant Lasius neonigerinvolves a series of trails which channel foragers away from adjacent colonies. Experimental studies suggest that the trails are composed of colony-specific, persistent orientation components of hindgut material that accumulate on trails during foraging. A less durable component of the hindgut trail pheromone regulates recruitment. Foraging directionality and the use of a trail could be modified by experimentally arranging confrontations with conspecifics. The orientation of foragers is mediated by visual as well as chemical cues. Components of the foraging and territorial system of L. neonigerappear to include (1) a network of subnests which change in position seasonally within each polydomous nest; (2) a series of trails emanating from each subnest that adjusts search toward resource patches and away from aggressive, neighboring conspecifics; and (3) trail communication involving an ephemeral component of the hindgut trail pheromone that regulates the organization of cooperative prey retrieval and a more persistent component that serves as an orientation guide.     

Nesting habitat characteristics of Marbled Murrelets occurring in near‐shore waters of the Olympic Peninsula,Washington

Marbled Murrelets (Brachyramphus marmoratus) are listed as threatened in the portion of their range extending from British Columbia to California due to loss of nesting habitat. Recovery of Marbled Murrelet populations requires a better understanding of the characteristics of their nesting habitat in this part of their range. Our objective, therefore, was to describe their nesting habitat in Washington State and Vancouver Island, British Columbia. We captured Marbled Murrelets from 2004 to 2008, fitted them with radio transmitters, and followed them to nests (N = 20). We used Cohen's unbiased d effect size to assess differences between forest plots surrounding nest sites and nearby control sites (N = 18). Nest sites had less canopy cover of the dominant conifers and fewer, but larger, trees than control sites. Nest sites also had greater percentages of trees with platforms >10 cm diameter and >15 cm diameter, and more platforms of these sizes than control sites. The mean diameter at breast height of nest trees was 136.5 cm (range = 84–248 cm) and all but one nest was in dominant or co‐dominant tree species. At the landscape scale, we used vegetation maps derived from remotely sensed data and found greater canopy cover, higher density of mature trees, more platforms >10 cm/ha, and more old‐growth habitat at nest sites than at random sites. Our findings suggest that, at the site scale, nesting Marbled Murrelets selected the most suitable features of forest structure across expansive potentially suitable habitat. Our landscape‐scale analysis showed that habitat features in nesting stands differed from those features in available stands in the murrelet's range in Washington. We also found that stands with nests were less fragmented than available forest across murrelet range. All nest sites of radio‐tagged birds in Washington were in protected areas in mostly undisturbed forest habitat. Conservation of these areas of inland nesting habitat will be critical to the recovery of Marbled Murrelet populations.