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Juvenile Tyto alba furcata (Barn Owl) and Remains of Ascalapha odorata (Black Witch Moth) Found in Natural Limestone Cavity in Hellshire Hills, Jamaica
Seth E. Inman, C. Justin Proctor, and John M. Zeigers

Caribbean Naturalist, No. 37

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Caribbean Naturalist 1 S.E. Inman, C.J. Proctor, and J.M. Zeiger 22001177 CARIBBEAN NATURALIST No. 37N:1o–. 1317 Juvenile Tyto alba furcata (Barn Owl) and Remains of Ascalapha odorata (Black Witch Moth) Found in Natural Limestone Cavity in Hellshire Hills, Jamaica Seth E. Inman1,*, C. Justin Proctor2, and John M. Zeiger1 Abstract - While conducting avian surveys throughout the Caribbean island of Jamaica, the authors discovered a natural limestone cavity in the Hellshire Hills containing a juvenile Tyto alba furcata (Barn Owl) as well as over a hundred wing fragments of Ascalapha odorata (Black Witch Moth). To date, there are no known records of such abundant lepidopteran prey remains associated with Barn Owl in the scientific literature. Herein we discuss the significance of these novel findings in the greater context of Barn Owl life history, address the paucity of research on the Barn Owl in Jamaica, and the implications of studies that have drawn conclusions on the diet of the Barn Owl solely from the analysis of pellet remains. We also include a preliminary literature review of insectivory and pellet analyses among owls. Introduction The Barn Owl Tyto alba Scopoli (Barn Owl) is one of the most widespread avian species in the world, with a range that includes the Americas, Australia, Africa, Asia, and Europe. Twenty-nine subspecies are recognized in the Clements Checklist, 6 of them endemic to islands throughout the Caribbean (Clements et al. 2014). Barn Owls typically eat smaller rodents, and their pellet contents are often used as a reliable means of assessing small-mammal populations (Meek et al. 2012). Tyto alba furcata (Temminck) (American Barn Owl) is a subspecies endemic to the Greater Antillean islands of Cuba and Jamaica, and the Cayman Islands (Bruce et al. 2014). The taxon has been reported in the Hellshire Hills (Brown 1994), but surprisingly little information on the Jamaican population is available in the scientific literature, and most of what is known about the American Barn Owl is from Cuba. Despite some persecution by Jamaicans for folkloric cultural reasons, the Barn Owl remains a common breeding resident on the island (Sutton et al. 2009); this antagonistic behavior toward Barn Owls is not limited to Jamaica (see Viamontes et al. 2002 for similarities in Cuba). We observed a Barn Owl in the Hellshire Hills within a limestone cavity where pellets and other prey remains were abundant. Noteworthy amongst these remains were over a hundred wing fragments from Ascalapha odorata L. (Black Witch Moth) scattered on the dusty cavity floor. This large (10–15-cm wingspan) Noctuoid moth is a breeding resident in Florida, Texas, Mexico, the Caribbean, and South America, and some populations migrate yearly to the US and Canada (Ekrem et al. 2014). 1Cornell Laboratory of Ornithology, Ithaca, NY 14850, USA. 2Department of Natural Resources, Cornell University, Ithaca, NY 14853, USA. *Corresponding author - Manuscript Editor: Jason Townsend Caribbean Naturalist S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 2 Insectivory. Insectivory by Barn Owls around the world is well-documented, and some studies have even found that the species may consume larger quantities of arthropods seasonally or when confined to islands, but only Bunn et al. (1982) and Bruce et al. (2014) mention moths as potential prey. Barn Owls have been reported eating insects in North America (Johnston and Hill 1987, Marti 1988), South America (Begall 2005, Bellocq 1998, de Groot 1983, Sahores and Trejo 2004), Europe (Durant et al. 2013, Glue 1974), Africa (Bunn et al. 1982), and the Caribbean islands (Buden 1974, Debrot et al. 2001, Flikweert et al. 2007, Hernández-Muñoz and Mancina 2011). Most studies published in the scientific literature report that the main insect orders represented in the Barn Owl diet are Blattodea (roaches), Coleoptera (beetles), and Orthoptera (grasshoppers and crickets); these orders have highly chitinous exoskeletons that are able to survive digestion and can remain intact within pellets. Pellets are the sole evidence for prey items in most Barn Owl diet surveys, given that they conveniently accrue most bones and can reliably be found near a roost. The accuracy of ascertaining Barn Owl dietary preferences from pellets has been supported elsewhere (see Clark and Bunck 1991, Glue 1974, Platt et al. 2009); however, it must be explicitly recognized that this method is primarily useful to detect vertebrates. Insect remains are not as well preserved in pellets as bones. In addition, insects are also far more diverse than vertebrates and are represented by smaller-bodied organisms that break more easily into unrecognizable body parts and make calculating the number of insect prey per pellet difficult. Swengel and Swengel (1992), for example, counted each insect fragment found in Aegolius acadicus Gmelin (Northern Saw-whet Owl) pellets as an individual prey item to avoid underrepresentation, while Marti (1974) relied on mandible, leg, wing cover, and head parts to count insect individuals in Athene cunicularia Molina (Burrowing Owl) pellets. Sampling of insect remains It is possible that the biomass of insect prey has been underestimated in Barn Owl dietary preference studies because pellet analysis, the most common and widespread technique is deficient in assessing invertebrate remains. This deficiency has frequently led to interpretations of insect contributions to the Barn Owl diet as negligible (see Bellocq 1998, Glue 1974, Marti 1988). Indeed, Bunn et al. (1982:92), when mentioning invertebrates in their book on the Barn Owl, discuss them in an incidental manner, declaring that “in those instances where we have seen invertebrates taken, the [Barn] Owls have always been newly fledged juveniles in the process of learning the arts of hunting.” They describe moths and other invertebrate species as “ludicrously insignificant [prey] species”. However, these sorts of claims may not apply equally in all ecosystems. Discussing the diversity of prey taken by Barn Owls, Taylor (1994:30,40) writes that, “Insects in particular seem to feature significantly in hotter, drier environments,” and, “On many islands, especially the drier ones, Barn Owls often catch a greater proportion of non-mammal prey, usually birds and insects, than in mainland areas.” He continues to explain that, “This increased use of insects is probably the Caribbean Naturalist 3 S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 result of a shortage of other larger items and the smaller size of island subspecies of the Barn Owl … may be an evolutionary response to a dependence upon these small prey.” Bruce et al. (2014) support Taylor’s claims, tersely describing the Barn Owl’s diet: “Much non-mammalian prey is caught incidentally or opportunistically, but can constitute a significant part of the diet in some tropical and semi-arid regions, e.g., lizards and invertebrates in parts of Africa and on some islands.” Strangely, in what appears to be one of the very few studies of the Barn Owl diet in Jamaica—carried out in the dry limestone scrub forest of the island— McFarlane and Garrett (1989) made absolutely no mention of insects; bats, birds, and anoles comprised the entirety of non-rodent prey reported from hundreds of pellets. However, studies from other tropical and semi-arid islands report evidence of higher insectivory. For example, on Santa Cruz in the Galápagos Islands, de Groot (1983) found that insects accounted for 63% of prey items in 1156 pellets. Taylor (1994) cited an unpublished study in Sicily in which Barn Owls increased their rates of insectivory to the level of 70–80% of all prey items during the 2 most hot and dry months of the year, supporting his claim that Barn Owls will take advantage of alternative prey when it is temporarily abundant. At a single site in Bonaire, Flikweert et al. (2007) analyzed pellet contents of 125 samples from 13 different roosts around the limestone gorge of Roi Sangu and found that 87% of prey items were insects. Interestingly, over the course of a 7-year study in Cuba, Hernández-Muñoz and Mancina (2011) found that in disturbed, anthropogenic habitat Barn Owls caught more insects than in natural habitats—though rodents remained the dominant prey species by far in both habitat types. Antúnez and Flores (2002) conducted an analysis of Barn Owl pellet remains in Cuba and suggested that, given the fragility of insect bodies, the actual number of insect prey items consumed was likely higher than what they were able to detect. On the possible deficiency of pellet analyses. Biases against invertebrates in pellet data have been repeatedly determined in Burrowing Owls (Plumpton and Lutz 1993, York et al. 2002). Artuso (2010:128) conducted a study on Megascops asio L. (Eastern Screech-Owl) with multiple prey-recording methods, including observational (direct and video monitored), pellet analysis, and nest inspection; he found that 57% of invertebrates were detected only in the observational techniques, and that invertebrates were “by far the most common prey during breeding, despite their relatively small contribution to biomass consumed.” Simmons et al. (1991) found that pellet analyses were biased when comparing mammal and bird prey in Circus ranivorus Daudin (African Marsh Harrier), while Redpath et al. (2001) showed that pellet data has “poor predictive power” for smaller prey, especially in predictions of biomass percent, for Circus cyaneus L. (Hen Harrier); the latter 2 groups of authors recommended combining different diet-analysis methods. Although they did not specify a bias against invertebrates, Yom-Tov and Wool (1997) ascertained that the contents of Barn Owl pellets did not represent the proportions of available prey in rural Israel, and were biased toward larger prey. Although they do have prey prefernces, Barn Owls have a flexible hunting strategy, and pellet analyses may not accurately detect prey density (Tores et al. 2005). Caribbean Naturalist S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 4 Figure 1. Map depicting the Hellshire Hills, Jamaica, and surrounding geography. Black circle indicates the location of the 2 cavities that contained Black Witch Moth remains and a juvenile Barn Owl. Major cities and features are labeled for reference. Map shading depicts elevation, with lightest gray = lowest, and darkest gray = highest. Field-site Description The Hellshire Hills constitute an area of about 160 km2 located approximately 10 km south of Spanish Town in the central region of Jamaica’s southern coast (Fig. 1). Appropriately named, the Hellshire Hills region is hot and dry, receiving less than 1020 mm of rainfall a year; its karstic limestone hills and plateaus have low levels of both leaf litter and soil, and the dominant vegetation is cacti and dry scrub forest (Tole 2002). The Hellshire Hills are vulnerable to resource extraction in the form of limestone quarries and charcoal deforestation (we observed evidence of both of these practices). The rediscovery of Cyclura collei Gray (Jamaican Iguana), thought to be extinct between the 1940s and 1990 and now listed as critically endangered on the IUCN Red List (Grant et al. 2010), has helped raise awareness of the need for protection of the Hellshire Hills’ rapidly disappearing Caribbean dry forest (Wilson et al. 2004). The Hellshires are known as a biodiversity hotspot within the biologically diverse island of Jamaica, and are the only remaining natural habitat for the Jamaican Iguana. Anthropogenic disturbances common in the area such as logging have been minimized, and efforts are underway to remove populations of feral Felis catus L. (Domestic Cat) and Canis lupus familiaris L. (Domestic Dog), as well as the introduced Herpestes javanicus auropunctatus Hodgson (Small Indian Mongoose) in the deep interior of the hills as part of a concerted protection effort for the Jamaican Iguana (Lewis et al. 2011). The fact that a substantially large reptile went unnoticed for half a century prior to being rediscovered speaks to both the isolated nature of the Hellshire Hills and the lack of research carried out in the region. Caribbean Naturalist 5 S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 Although environmental interest is starting to increase, the future of the Hellshire Hills’ biodiversity would strongly benefit from greater attention from the scientific and conservation communities. Methods At ~1220 on 24 March 2015, while conducting point-count surveys for the probably extinct Tachycineta euchrysea euchrysea (Gosse) (Golden Swallow, Jamaican subspecies) in the Hellshire Hills region, we noticed a cavity in a limestone ridge just off a local access trail (17°55'33''N, 76°59'34''W; Proctor et al., in press). Closer inspection revealed that the cavity floor was covered with the wings of lepidopterans and orthopterans (Fig. 2). Curious as to their origin, we proceeded to a neighboring cavity, where we found owl pellets and urea stains outside the entrance. Both cavities were ~3–4 m in horizontal depth from entrance to back wall and 2–3 m in vertical height from ground to ceiling. The entrances were level with the ground outside, quite wide, and allowed a medium level of ambient light into the cavities. We describe the separate cavities as shelters because they did not appear to link anywhere else and were not deep enough to be considered caves. Upon closer observation into the recesses of the second cavity, we startled a juvenile Barn Owl that quickly hopped from where it had been roosting near the entrance to a low, narrow shaft at the back of the small cavity. After verifying that the shaft was longer than 1 m, and with no source of light to illuminate the tunnel, we withdrew from the cavity to allow the owlet time to return Figure 2. Profile view of the larger of the cavities found in the Hellshire Hills, Jamaica. A mix of Lepidopteran wings and dead leaves (lighter brown color) can be seen scattered across the rocky floor. Caribbean Naturalist S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 6 to the main cavity and so that we could acquire headlamps for better inspection of the environs. Approximately 20 minutes after the initial encounter, the owlet was quiet and motionless at the end of the same shaft, which measured roughly 2 m in length and 0.5 m in diameter. Despite the close proximity (~1.5 m) of one of the observers when approaching on hands and knees to take a photograph, the owlet did not emit any distress calls, nor did it attempt to defend itself by biting at the observer or rolling onto its back to feign death or strike with its talons, which are all commonly observed responses exhibited by juvenile Barn Owls when threatened (Bunn et al. 1982). We opportunistically collected both insect wings and owl pellets that were visible and in good condition. We made an effort to retrieve samples of wings that represented all moth species present; it was clear that fragments from one type of moth were far more numerous than others. We collected owl pellets and loose mammal skulls outside the cave entrance, along with a single adult Barn Owl feather. Our inability to obtain 100% of the prey remains eliminated the possibility of comparative quantitative analyses between categories of prey remains; thus, we have focused more closely on the general abundance and classification of prey remains where possible. Results Based on in-situ observations and photographs of the interior of both cavities, we estimated that between 100–150 moth wings, and ~50 orthopteran wing fragments were present on the floors. We identified all but 3 of the roughly 30 wings collected as those of Ascalapha odorata L. (Black Witch Moth); the species of the remaining 3 lepidopteran wings could not be determined. We identified the specimens (Fig. 3) using the Mississippi Entomological Museum’s digital guide to moth identification (Moth Photographers Group 2013) and Dr. Delano S. Lewis, systematic entomologist at Northern Caribbean University in Mandeville, Jamaica, confirmed these results. Both fore- and hindwings of Black Witch Moth were present in the cavities, although forewings appeared to have been more common, likely due to their larger size. We did not identify samples of orthopteran wings. We dissected 4 pellets and detected 7 Rattus rattus L. (Black Rat) skulls from the midden area outside the cavities among the pellets and excrement. We identified a total of 17 Mus musculus L. (House Mouse) and 4 Black Rat individuals based on the presence of House Mouse lower mandibles and Black Rat femurs; 1 passerine tarsus was found. Given the relatively nonsystematic sampling of pellets from outside the cavities, we provide these data primarily as anecdotal evidence and for context. It is important to note that we observed insect remains only as wings on the cavity floors and not within pellets; we are not trained entomologists who can identify body fragments other than obvious structures like wings. We identified the Barn Owl immediately based on the plumage color and facial disk of the juvenile; Jamaica is home to only the Barn Owl and Jamaican Owl, which look very different. Subsequent cross-referencing of our photograph (see cover photo) with Bunn et al. (1982) confirmed the bird as a juvenile Barn Owl (see below). Caribbean Naturalist 7 S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 Discussion Our goals are to bring attention to the unusual amount of lepidopteran prey remains in the Barn Owl roost found within the Hellshire Hills of Jamaica, and present a short literature review to suggest the importance of looking for evidence of insectivory when studying owls in the tropics. The results we gleaned from pellet dissections generally match the usual strong presence of rodent remains in Barn Owl pellets. However, the high numbers of moth wings scattered about the cavity floors indicate that invertebrates may play a significant role in juvenile and/ or adult Barn Owl diet in the Hellshire Hills. At the very least, foraging time, and thus, energy demands associated with the hunting of these invertebrates must be considerable. Using pellets alone to ascertain Barn Owl diet in the Hellshire Hills may not be an accurate representation of prey ratios, especially without an entomologist to review pellet contents. As Flikweert et al. (2007:75) correctly asserted, “Relatively few data are available on the dietary composition of Barn Owls from Figure 3. Black Witch Moth (Ascalapha odorata) forewings collected from the floor of a limestone cavity in the Hellshire Hills, Jamaica. Forewings from 2 different moths are shown in order to display both dorsal and ventral coloration patterns. Caribbean Naturalist S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 8 tropical regions, and in particular from (small) tropical islands.” We encourage further research on Barn Owl in Jamaica, as well as any ornithologically related research in the Hellshire Hills, given the area’s vulnerability to human development and ecological singularity on the island. On the juvenile Barn Owl Based on the sequential aging photographs in Bunn et al. (1982:99), the owlet in the Hellshires was likely very close to 42 days old (see Photo 9). Bunn et al. (1982) cite several authors in stating that the fledge-age range is 56–62 days, and especially given the abundant down on the individual we observed, the owlet had likely not yet fledged and was probably unable to hunt for itself, especially on the wing for dozens of moths. Thus, the hen likely provided the owlet with the moths. Artuso (2010:128) has noted from personal observations on the Eastern Screech-Owl that “invertebrates are important for the provisioning of broods, even though they contribute comparatively little biomass, as they can be captured close to the nest and delivered frequently, as often as every 45 seconds in some cases.” Conclusion We suggest that insectivory in Barn Owls remains understudied and unappreciated, primarily due to the relatively low percentage of total biomass that insects represent in pellet samples and the prevalence of pellet analysis as the sole source of data in dietary studies. Even in the unlikely case that this low biomass in pellets is an accurate reflection of actual diet, the fact remains that Barn Owls clearly catch many insects in addition to the better-known food sources—rodents, bats, and other vertebrates—that are more easily discernible as prey remains. Barn Owl insectivorous foraging, as was seen in case studies from the aforementioned islands of Cuba, the Galápagos, Bonaire, and Sicily, must require a significant amount of time and energy, and thus represents an important element of hunting and feeding behavior even apart from the subject of nutrition itself. Methods of ascertaining the types of insect prey on a more specific level than order or family might be created through observational data, stomach-content analyses, or sampling of pellets in a manner other than purely osteological. Acknowledgments Fieldwork was supported by the Smithsonian Institution’s James Bond Fund, the Cornell Lab of Ornithology, The Rufford Foundation, and Cornell University’s Betty Miller Francis Fund. Invaluable gear and materials were donated by IDEA WILD and Nature Balance Foods, and loaned by Cornell Outdoor Education. Noah Hamm of Cornell University helped provide access to library resources, and Delano S. Lewis verified the identification of Black Witch Moth. Carlos Arredondo Antúnez and Ana Trejo critiqued earlier drafts of the manuscript. We are grateful to the anonymous reviewers and the guest editor for taking the final steps in improving the strength of the paper. Special thanks go to Gary Graves from the Smithsonian Institution for creating the opportunity to pursue this research. Caribbean Naturalist 9 S.E. Inman, C.J. Proctor, and J.M. Zeiger 2017 No. 37 Literature Cited Antúnez, C.A,. and V.N.C. Flores. 2002. 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