DOI: 10.5061/dryad.c866t1gjh
title: Marketing birds: The traits birdwatching tourism companies highlight in Costa Rican tour itineraries

description: Birdwatching tourism has blossomed into a profitable industry, promoting
 peoples’ connections to nature and contributing billions of dollars to
 local economies annually. Understanding which species birdwatchers value
 and why could help distinguish flagship species for ecotourism and
 conservation marketing. Here, we draw from ecological, psychological, and
 economic perspectives to test hypotheses about which species’ attributes
 (i.e., traits) are most prevalent among birds marketed to birdwatchers. To
 do so, we implemented a multi-year course-based undergraduate research
 experience to quantify how often &gt;750 bird species appeared in
 &gt;150 itineraries describing tours traversing Costa Rica. We then
 associated itinerary mentions with data on species’ appearances,
 vocalizations, distributions, conservation statuses, and behaviors. We
 found that larger, more attractive species appeared frequently in trip
 itineraries, but species with pleasing vocalizations did not. We also
 found that range-restricted species appeared ten times more frequently in
 trip itineraries than cosmopolitan species. Species with tropical
 distributions that rarely associate with humans were also more frequently
 mentioned, suggesting that companies recognize the high value dedicated
 birdwatchers place on novel species. Surprisingly, IUCN status was not
 associated with itinerary scores, and commonly encountered species were
 more likely to be mentioned in trip itineraries than rarely encountered
 species. These findings suggest tour companies may balance species’
 perceived novelty to birdwatchers against the likelihood that they can
 ‘deliver’ them to their clients. Our findings broadly contribute to a
 deeper understanding of how human preferences shape interactions with
 biodiversity and may help bridge the gap between cultural ecosystem
 services and conservation strategies. Insights into which traits drive
 birdwatcher preferences could help conservation practitioners and tour
 operators identify species potentially at risk from unregulated
 birdwatching tourism as well as species that could be used as flagships to
 motivate ecotourism and conservation actions.
description: # Marketing birds: The traits birdwatching tourism companies highlight in
 Costa Rican tour itineraries Dataset DOI:
 [10.5061/dryad.c866t1gjh](10.5061/dryad.c866t1gjh) ## Description of the
 data and file structure We combined published functional trait databases
 with data collected from a multi-year course-based undergraduate research
 experience (CURE) to predict which of &gt;750 bird species were most
 likely to appear in &gt;150 tour itineraries marketed to birdwatchers
 visiting Costa Rica. See methods from associated paper.  ### Files and
 variables #### File: BirdwatchingTraitDatabase_Dryad.csv
 **Description:** Trait database.  ##### Variables * Order: Taxonomic order
 * Family: Taxonomic family * ScientifcName: Latin name (genus, species) *
 CommonName: English name  * included2020: Number of itineraries in which
 the species was present from the 2020 class * notincluded2020: Number of
 itineraries in which the species was not present from the 2020 class *
 fraction2020: Proportion of itineraries in which the species was present
 from the 2020 class * included2022: Number of itineraries in which the
 species was present from the 2022 class * notincluded2022: Number of
 itineraries in which the species was not present from the 2022 class *
 fraction2022: Proportion of itineraries in which the species was present
 from the 2022 class * included2020_More3Days: Number of itineraries in
 which the species was present from the 2020 class, only including
 itineraries for trips more than 3 days in length. *
 notincluded2020_More3Days: Number of itineraries in which the species was
 not present from the 2020 class, only including itineraries for trips more
 than 3 days in length. * fraction2020_More3Days: Proportion of itineraries
 in which the species was present from the 2020 class, only including
 itineraries for trips more than 3 days in length. *
 included2022_More3Days: Number of itineraries in which the species was
 present from the 2022 class, only including itineraries for trips more
 than 3 days in length. * notincluded2022_More3Days: Number of itineraries
 in which the species was not present from the 2022 class, only including
 itineraries for trips more than 3 days in length. *
 fraction2022_More3Days: Proportion of itineraries in which the species was
 present from the 2022 class, only including itineraries for trips more
 than 3 days in length. * included2020_More20Spp: Number of itineraries in
 which the species was present from the 2020 class, only including
 itineraries for trips with more than 20 species mentioned *
 notincluded2020_More20Spp: Number of itineraries in which the species was
 not present from the 2020 class, only including itineraries for trips with
 more than 20 species mentioned * fraction2020_More20Spp: Proportion of
 itineraries in which the species was present from the 2020 class, only
 including itineraries for trips with more than 20 species mentioned *
 included2022_More20Spp: Number of itineraries in which the species was
 present from the 2022 class, only including itineraries for trips with
 more than 20 species mentioned * notincluded2022_More20Spp: Number of
 itineraries in which the species was not present from the 2022 class, only
 including itineraries for trips with more than 20 species mentioned *
 fraction2022_More20Spp: Proportion of itineraries in which the species was
 present from the 2022 class, only including itineraries for trips with
 more than 20 species mentioned * AveTourLength_2020: The average tour
 length (in days) for all the trip itineraries a given species appeared on,
 in 2020 * AveTourLength_2022: The average tour length (in days) for all
 the trip itineraries a given species appeared on, in 2022 *
 AveTourSppRich_2020: The average number of species across all the trip
 itineraries a given species appeared on, in 2020 *
 AveTourSppRich_2022: The average number of species across all the trip
 itineraries a given species appeared on, in 2022 * ave_fraction: Average
 proportion of itineraries in which the species was present across both
 years * ave_fraction_More3Days: Average proportion of itineraries in which
 the species was present across both years, restricted to itineraries for
 trips that were greater than 3 days in length. *
 ave_fraction_More20Spp: Average proportion of itineraries in which the
 species was present across both years, restricted to itineraries for trips
 with more than 20 species reported. * ave_TourLength_BothYears: The
 average tour length (in days) for all the trip itineraries a given species
 appeared on, across both years * ave_TourRichness_BothYears: The average
 number of species across all the trip itineraries a given species appeared
 on, across both years * Mass_Avo: Body mass given as species average
 (incorporating both male and female body mass) from AVONET (grams) *
 BeakCulmen_Avo: Length from the tip of the beak to the base of the
 skull from AVONET (mm) * BeakNares_Avo: Length from the anterior edge of
 the nostrils to the tip of the beak from AVONET (mm) * BeakWidth_Avo:
 Width of the beak at the anterior edge of the nostrils from AVONET (mm) *
 BeakDepth_Avo: Depth of the beak at the anterior edge of the nostrils from
 AVONET (mm) * Tarsus_Avo: Length of the tarsus from the posterior notch
 between tibia and tarsus, to the end of the last scale of acrotarsium (at
 the bend of the foot) (mm) * Wing_Avo: Length from the carpal joint (bend
 of the wing) to the tip of the longest primary on the unflattened
 wing from AVONET (mm) * HandWing_Avo: 100*DK/Lw, where DK is Kipp’s
 distance and Lw is wing length (i.e., Kipp’s distance corrected for wing
 size) from AVONET. Species average HWI differ from estimates in Sheard et
 al. (2020) because of much higher sampling of individuals in some species,
 as well as taxonomic effects in the BirdLife list (mm) *
 Tail_Avo: Distance between the tip of the longest rectrix and the point at
 which the two central rectrices protrude from the skin, typically measured
 using a ruler inserted between the two central rectrices from AVONET (mm)
 * PrimaryHabitat_Avo: Habitat affiliation from AVONET. Desert (= drylands
 and other open arid habitats, often sandy with very sparse vegetation);
 Rock (= rocky substrate typically with no or very little vegetation,
 including rocky outcrops, rocky coastlines, arid stony steppes, rocky
 mountaintops and mountain slopes); Grassland (= open dry to moist
 grass-dominated landscapes, at all elevations); Shrubland (= low stature
 bushy habitats, included thornscrub, thorny or arid savanna, caatinga,
 xerophytic shrubland and coastal scrub); Woodland (= medium stature
 tree-dominated habitats, including Acacia woodland, riparian woodlands,
 mangrove forests, forest edges, also more open parkland with scattered
 taller trees); Forest (= tall tree-dominated vegetation with more or less
 closed canopy, including palm forest); Human modified (urban landscapes,
 intensive agriculture, gardens); Wetland (= wide range of freshwater
 aquatic habitats including lakes, marshes, swamps and reedbeds); Riverine
 (= associated with rivers and streams at all elevations); Coastal (=
 intertidal zones within immediate vicinity of beaches, estuaries, brackish
 to salty marshes, including mudflats, lagoons, alkaline wetlands, coastal
 dunes and harbours); Marine (= pelagic, on sea near coasts, including
 species in the intertidal zone on beaches, and those pelagic species
 nesting near the sea on cliffs, islets and islands). *
 HabitatDensity_Avo:  Habitat density from AVONET. 1= Dense habitats.
 Species primarily lives in the lower or middle storey of forest, or in
 dense thickets, dense shrubland etc. 2= Semi open habitats. Species
 primarily lives in open shrubland, scattered bushes, parkland, low dry or
 deciduous forest, thorn forest. 3= Open habitats. Species primarily lives
 in desert, grassland, open water, low shrubs, rocky habitats, seashores,
 cities. Also applies to species living mainly on top of forest canopy
 (i.e. mostly in the open) * HabitatStrata_Elton: Habitat strata score from
 Elton Traits, where higher numbers are higher up in the canopy. Formula
 is:
 (BelowWaterProportion)*0+(WaterSurfaceProportion)*1+(GroundProportion)*1+(UnderstoryProportion)*2+(MidStrataProportion)*3+(CanopyProportion)*4+(AerialProportion)*5 * Migration_Avo: Migration score from AVONET. 1 = Sedentary. 2 = Partially migratory, i.e. minority of population migrates long distances, or most of population undergoes short-distance migration, nomadic movements, distinct altitudinal migration, etc. 3 = Migratory, i.e. majority of population undertakes long-distance migration * TrophicLevel_Avo: Trophic level from AVONET. Herbivore = species obtaining at least 70% of food resources from plants; Carnivore = species obtaining at least 70% of food resources by consuming live invertebrate or vertebrate animals; Scavenger = species obtaining at least 70% of food resources from carrion or refuse; Omnivore = species obtaining resources from multiple trophic level in roughly equal proportion * TrophicNiche_Avo: Trophic niche from AVONET. Frugivore = species obtaining at least 60% of food resources from fruit; Granivore = species obtaining at least 60% of food resources from seeds or nuts; Nectarivore = species obtaining at least 60% of food resources from nectar; Herbivore = species obtaining at least 60% of food resources from other plant materials in non-aquatic systems, including leaves, buds, whole flowers etc.; Herbivore aquatic = species obtaining at least 60% of food resources from plant materials in aquatic systems, including algae and aquatic plant leaves; Invertivore = species obtaining at least 60% of food resources from invertebrates in terrestrial systems, including insects, worms, arachnids, etc.; Vertivore = species obtaining at least 60% of food resources from vertebrate animals in terrestrial systems, including mammals, birds, reptiles etc.; Aquatic Predator = species obtaining at least 60% of food resources from vertebrate and invertebrate animals in aquatic systems, including fish, crustacea, molluscs, etc; Scavenger = species obtaining at least 60% of food resources from carrion, offal or refuse; Omnivore = Species using multiple niches, within or across trophic levels, in relatively equal proportions * Lifestyle_Avo: Lifestyle class from AVONET. Aerial = species spends much of the time in flight, and hunts or forages predominantly on the wing; Terrestrial = species spends majority of its time on the ground, where it obtains food while either walking or hopping (note this includes species that also wade in water with their body raised above the water); Insessorial = species spends much of the time perching above the ground, either in branches of trees and other vegetation (i.e. arboreal), or on other raised substrates including rocks, buildings, posts, and wires; Aquatic = species spends much of the time sitting on water, and obtains food while afloat or when diving under the water's surface; Generalist = species has no primary lifestyle because it spends time in different lifestyle classes * SocialBond_Tob: Social bond score from Tobias et al 2016 (https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2016.00074/full). 1= solitary (does not form pairs or only just briefly during courtship); 2= short-term pair/group bonds (seasonal consorting of pair or group members AND unstable composition in the future); 3= long-term pair/group bonds (year-round consorting or those with seasonal pair/group bonds AND stable composition in subsequent breeding attempts) * Nocturnal_Elton: Whether (1) or not (0) the species is nocturnal from Elton traits.  * Territoriality_Tob:  Territoriality score from Tobias et al 2016 (https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2016.00074/full). 1= non-territorial (never defends territories); 2= seasonal or weak territoriality; 3= year-round territoriality * CommunalSignal_Tob:  Communal signaling score from Tobias et al 2016 (https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2016.00074/full). 0= non-communal signallers (species that do not sing communally); 1= communal signallers (species that sing in groups) * Duet_Tob:  Duet score from Tobias et al 2016 (https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2016.00074/full). 0= no duetting (species that do not duet); 1= duetting (species that sing in duets)  * Chorus_Tob:  Chorus score from Tobias et al 2016 (https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2016.00074/full).  0= no chorus (species that do not sing choruses); 1= chorus (species that sing as part of chorus) * MinLatitude_Avo: The minimum latitudinal extent of the species range (restricted to breeding and resident range) from AVONET (decimal degrees) * Max.Latitude_Avo: The maximum latitudinal extent of the species range (restricted to breeding and resident range) from AVONET (decimal degrees) * CentroidLatitude_Avo: The geometric centre of the species range (restricted to breeding and resident range) from AVONET (decimal degrees) * Range.Size_Avo: The total area of the mapped range of the species (not the Extent of Occurrence [EOO]) from AVONET (square km). We used maps shared by BirdLife International and restricted our analysis to areas of the range coded as Presence = 1 (Extant only), Origin = 1 & 2 (Native & Reintroduced), Seasonal = 1 & 2 (Resident and breeding). We calculated the total combined mapped area of these parts of the range using the areaPolygon function from the R package geosphere (Hijmans et al., 2011). This function accurately calculates the area in the World Geodetic System (WGS84) projection using spherical distances * FOO_ave: Average frequency of occurance across ebird checklists year round * Status_IUCN: IUCN Red List Status (Least concern, near threatened, endangered, vulnerable, critically endangered) * Trend_IUCN: Population trend (increasing, decreasing, stable, unknown) * ED_Jetz: Evolutionary Distinctness Score: the weighted sum of branch lengths from the root of the tree to the tip (i.e., a species). from Jetz et al 2014 Current Biology * AttractivenessMean_sant: Predicted appearance score frrm Santangeli et al 2023, where higher numbers indicate higher scoring species * AttractivenessSD_sant: Standard deviation in appearance scores from Santangeli et al 2023 * SexMeasured_sant: Which sex was assessed for appearance: average= unknown sex usually for species that can't be discerned; female= female bird only; male= male bird only; male_female_mean= both sexes measured so avereages were taken except for crest when the max value was taken * Colorfraction_black_sant: Fraction of the body in images from Santangeli et al 2023 that was black * Colorfraction_blueD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark blue * Colorfraction_blueL_sant: Fraction of the body in images from Santangeli et al 2023 that was light blue * Colorfraction_brownD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark brown * Colorfraction_brownL_sant: Fraction of the body in images from Santangeli et al 2023 that was light brown * Colorfraction_greenD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark green * Colorfraction_greenL_sant: Fraction of the body in images from Santangeli et al 2023 that was light green * Colorfraction_greyD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark grey * Colorfraction_greyL_sant: Fraction of the body in images from Santangeli et al 2023 that was light grey * Colorfraction_purpleD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark purple * Colorfraction_purpleL_sant: Fraction of the body in images from Santangeli et al 2023 that was light purple * Colorfraction_redD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark red * Colorfraction_redL_sant: Fraction of the body in images from Santangeli et al 2023 that was light red * Colorfraction_rufousD_sant: Fraction of the body in images from Santangeli et al 2023 that was dark rufous * Colorfraction_rufousL_sant: Fraction of the body in images from Santangeli et al 2023 that was light rufous * Colorfraction_white_sant: Fraction of the body in images from Santangeli et al 2023 that was white * Colorfraction_yellow_sant: Fraction of the body in images from Santangeli et al 2023 that was yellow * N.loci_sant: Number of color loci, a measure of colour diversity from Santangeli et al 2023 * Elaboration_sant: How much the overall plumage departs from the average bird (i.e. dull brown-grey coloration) from Santangeli et al 2023 * Sexdichtonomy_sant: Degree of dichromatism among sexes from Santangeli et al 2023 * Crest_sant: Crest presence from Santangeli et al 2023 0= = no crest, 1 = crest follows shape of head, 2 = short crest, and 3 = long crest * TipName: Label of tip in BirdTree.org * NumAudioRecordings: Average number of audio recordings across Xeno-canto and Macaulay libraries * AcousticsScore_Ave: Average acoustic aesthetic score * AcousticsScore_Modeled: Modeled acoustic aesthetic score accounting for variation among scorers (see Methods) ## Code/software All analyses were conducted in R vers 4.5.1 (see methods of associated paper).  ## Access information Data was derived from the following sources: * Online birdwatching tour itineraries (see methods) * Tobias, J. A., C. Sheard, A. L. Pigot, A. J. M. Devenish, J. Yang, M. H. C. Neate-Clegg, N. Alioravainen, T. L. Weeks, R. A. Barber, P. A. Walkden, H. E. A. MacGregor, et al. (2022). AVONET: morphological, ecological and geographical data for all birds. *Ecology Letters* 25:581–597. * Tobias, J. A., C. Sheard, N. Seddon, A. Meade, A. J. Cotton, and S. Nakagawa (2016). Territoriality, Social Bonds, and the Evolution of Communal Signaling in Birds. *Frontiers in Ecology and Evolution* 4:1–15. * Wilman, H., J. Belmaker, S. Jennifer, C. de la Rosa, M. M. Rivadeneira, and W. Jetz (2014). EltonTraits 1.0: Species-level foraging attributes of the world’ s birds and mammals. *Ecology* 95:2027. * Santangeli, A., A. Haukka, W. Morris, S. Arkkila, K. Delhey, B. Kempenaers, M. Valcu, J. Dale, A. Lehikoinen, and S. Mammola (2023). What drives our aesthetic attraction to birds? *npj Biodiversity* 2:1–7. * eBird (2021). eBird: An online database of bird distribution and abundance [web application]. [Online.] Available at http://www.ebird.org. * Jetz, W., G. H. Thomas, J. B. Joy, D. W. Redding, K. Hartmann, and A. O. Mooers (2014). Global Distribution and Conservation of Evolutionary Distinctness in Birds. *Current Biology* 24:919–930. * IUCN (2024). The IUCN Red List of Threatened Species.