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Factors associated with arrival timing and condition of migrant landbird species in northeastern Pennsylvania

Factors associated with arrival timing and condition of migrant landbird species in northeastern... Anim. Migr. 2022; 9: 37–47 Research Article Robert J. Smith*, Margret I. Hatch, Jason M. Graham Factors associated with arrival timing and condition of migrant landbird species in northeastern Pennsylvania https://doi.org/10.1515/ami-2022-0119 1 Introduction received March 29, 2022; accepted July 25, 2022 Timing of spring migration in passerines, and conse- Abstract: Extrinsic and intrinsic factors operating during quently arrival at the breeding grounds has reproductive and prior to the passerine spring migratory period have consequences [1-3] and may be influenced by intrinsic and been associated with both migratory timing and condi- extrinsic factors operating both during passage as well as tion. Here we take advantage of a long-term data set to prior to the migratory event. For example, genetic control answer questions about how extrinsic factors encoun- mechanisms influence onset of migratory restlessness, tered on the wintering grounds (El Niño Southern Oscil- temporal pattern and direction of migratory activity as well lation, ENSO) and en route (temperatures south of our as annual patterns of body mass [4]. Further, within-spe- study site) along with intrinsic factors (age, sex if possi- cies migratory strategies may differ, leading to intra-spe- ble) influenced both arrival timing and condition at our cific differences in timing of passage [5] and arrival at the site in northeastern Pennsylvania. Older birds preceded migratory destination [6]. The classic example of differ- younger, male Common Yellowthroats (Geothlypis trichas) ential passage in North American landbirds is the early preceded females and within a year later arriving Gray Cat- migration and arrival of males to breeding areas [e.g., birds (Dumetella carolinensis) and Common Yellowthroats 7, 8]. Perhaps due to advantages in gainingaccess to the were in better condition. We found that Gray Catbirds and highest quality territories [9, 10 - rank advantage hypoth- Common Yellowthroats migrating during warmer years esis], more mating opportunities [11 - mate opportunity arrived in better condition. Finally, we found evidence that hypothesis], differences in the sex-specific fitness costs ENSO, likely via influencing weather and food availability and benefits of arrival timing [12 - sexual conflict hypoth- during the winter, was associated with arrival timing in esis] or some combination [13]. Finally age and sex-related Veery (Catharus fuscescens), Common Yellowthroats and dominance asymmetries, via influencing en route [14] or possibly Gray Catbirds. Our results support the hypothesis winter habitat occupancy [15, 16], have also been related that events experienced earlier, either between (wintering to timing of arrival in breeding areas as well as reproduc- to migratory periods) or within (earlier vs. later in migra- tive success for that season [3]. tion) phases of the avian annual cycle may carry over, Extrinsic factors have also been linked to timing of influencing fitness later in time or in subsequent phases departure from the wintering grounds [17-20], en route of the annual cycle. passage timing [20-24] and condition (25) as well as timing Keywords: arrival timing; carry-over effects; landbird; of arrival at northerly breeding grounds [6, 21, 26, 27]. spring migration For example, Studds and Marra [17] demonstrated rela- tionships between rainfall, food resources and departure dates of American Redstarts (Setophaga ruticilla) from wintering grounds in Jamaica with birds departing earlier in years with more rainfall and food resources. *Corresponding author: Robert J. Smith, Department of Biology, More recently, wintering ground environmental con- The University of Scranton, Scranton, PA 18510, USA, Email: Robert. ditions, as indicated by the El Niño Southern Oscillation Smith@scranton.edu Margret I. Hatch, Penn State Scranton, 120 Ridge View Drive, Dun- (ENSO), have been related to passage timing [27-29] as more, PA 18512, USA well as energetic condition in spring landbird migrants Jason M. Graham, Department of Mathematics, The University of passing through both southerly [25] and northerly [28, 30] Scranton, Scranton, PA 18510, USA Open Access. © 2022 Smith et al., published by De Gruyter. This work is licensed under the Creative Commons Attribution 4.0 License. 38   Smith et al. stopover sites in North America. ENSO events cause vari- habitat. We captured birds in shrub-dominated habitats ation in precipitation and temperature in parts of South through both the spring migratory period and into the rd nd America and the Caribbean-Central America region [31, breeding season (from the 3 week of April through the 2 32] both of which influence habitat quality for wintering week of June). We delineated the endpoint of spring migra- migrant songbirds [33, 34]. Further, both Hüppop and tion for each species by examining histograms of first cap- Hüppop [35] and Tøttrup et al. [36] found en route temper- tures, determining the date when number of first captures ature to be associated with mean spring passage timing dropped to zero. We operated between 10 (2004, 2011 – in a majority of short and long distance migrants moving 2019, 2021) and 16 (2005 – 2010) permanently positioned through Europe while in North America Marra et al. [22] mist-nets, checking nets at 30 minute intervals. Nets were found median passage dates to be inversely related to en opened by sunrise, remaining open until late-morning. route temperatures in the majority of species examined. We did not capture birds in the event of excessively low Finally, Balbontín et al. [21] found an advance in timing temperatures or rain. For each individual captured we of arrival at breeding grounds for barn swallows (Hirundo recorded species, age and when possible to determine rustica) when ecological conditions en route improved by plumage, sex [39], mass and tarsus length along with while Smith et al. [6] demonstrated relationships between capture date and time. Birds were banded with a U.S. annual variation in environmental conditions and timing Geological Service aluminum leg band and all recaptures of arrival at northerly breeding grounds in American Red- measured without reference to previous records. In all starts. subsequent analyses we only use data collected the first Even as a large body of literature exists linking extrin- time within a year an individual was captured and include sic factors to en route passage timing [6, 21-23, 37] less is only those species for which we captured 10 or more indi- known about how these factors influence en route condi- viduals within a year for at least 80% of the years encom- tion and especially if and how ENSO events taking place passed by this study (veery – 82%, Gray Catbird – 100%, on the wintering grounds influence arrival timing and Common Yellowthroat – 100%). For all individuals within condition of birds at northerly stopover and breeding each species we estimated body condition using the areas [28]. Our objective was to document passage timing Scaled Mass Index of Peig and Green [40], using tarsus and size-corrected body mass at first capture using a long- length to adjust mass as if all individuals within a species term (17 yr) dataset collected during spring migration in were the same size. northeastern Pennsylvania. We tested the hypotheses that We estimated environmental conditions on the win- 1) extrinsic factors encountered both on the wintering tering grounds following the methodology of Paxton et al. grounds (as indicated by ENSO) as well as during migra- [41]. We identified years between 2004 and 2021 as El Niño, tion (en route temperatures), and 2) intrinsic factors (age La Niña or non-ENSO during winter using the National and when possible sex) influenced condition and timing Weather Service Climate Prediction Center (ONI; https:// of arrival at our site in northeastern Pennsylvania, USA. www.cpc.ncep.noaa.gov/products/precip/CWlink/MJO/ enso.shtml mean monthly values of the standardized ENSO conditions (January to March). This index is a measure of departure from long-term average sea surface 2 Methods temperature (SST) in the east-central Pacific Ocean within o o 0 o the Nino 3.4 region (5 N-5 S, 120 -170 W). The index clas- We collected data describing arrival phenology and sifies El Niño and La Niña events based on a threshold size-corrected body mass of the above species in Lacka- of 3 consecutive months above the 0.5 C SST anomaly (El wanna County, Benton Township, northeastern Pennsyl- Niño) or below the -0.5 C SST anomaly (La Niña). vania at both Lackawanna State Park and private lands We investigated the effect of en route conditions on immediately adjacent to the Park from 2004 – 2019, 2021. arrival timing and condition by downloading temperature All netting locations were within 1.8 km of each other. data collected from The National Centers for Environmen- Shrub habitat was approximately 25-35 years post agricul- tal Prediction (NCEP)/National Center for Atmospheric ture and was a mix of exotic (primarily honeysuckle [Lon- Research (NCAR) Reanalysis data set [42] using R package icera spp.]) and native shrubs (primarily dogwood [Cornus RNCEP, version 1.0.7 [43]. This data set is high-quality, well spp.] and prior to infestation by exotic Viburnum Leaf documented, freely available and is increasingly being Beetle (Pyrrhalta viburni) in 2008, arrowwood viburnum used in ecological research [43], including bird migration (Viburnum dentatum) (see 38) as well as a small number [44, 45]. To estimate annual variation in spring tempera- of saplings of most tree species found in nearby forested tures encountered by migrating birds we selected temper- Landbird arrival in northeastern Pennsylvania    39 ature data from areas south of our study site bounded by sex on passage timing. We treated capture day, April and 30.0 to 39.9 N and -85.0 to -78.0 W for April and May, 2004- May temperatures as continuous variables and age, sex 2021 (Figure 1, En route Stations). We selected this area to and ENSO as factors (Table 1). To evaluate size-corrected maximize the likelihood that temperature data reflected body mass at arrival we used a similar modeling approach what the individuals captured at our site experienced though because mass may change with date we included en route as most populations of migrant landbirds tend ordinal date as a numerical variable in the model (Table to follow a north-south axis [46, 47]. Further, modeling 1). For Veery we omitted data from 2004, 2012 and 2018 (Gray Catbird, Common Yellowthroat; 48], geolocator data because we captured less than 10 birds in these years. (Veery; 49) geolocator and recapture data (Gray Catbird; We evaluated assumptions for each model by following 49) and genetic data (Common Yellowthroat; 48, 50, 51) the methodology of Zuur et al. [57]. Using a lme approach support our assumption that our temperature data reflect allowed us to account for heterogeneity and temporal what birds experienced prior to arrival at our site. autocorrelation as necessary [57]. We screened data for Because ENSO events may influence temperature and multicollinearity using function vif from package ‘usdm’ precipitation in the southeastern United States [52, 53] we [58] and used the package ‘emmeans’ [59] to calculate evaluated another NCEP data set, using temperature data parameter estimates and where necessary Tukey post hoc from stations within an area bounded by 25.0 to 32.5 N and tests. While we performed post hoc tests using all three -93.7 to -78 W (Figure 1, Gulf Coast stations). We looked for an influence of winter ENSO on environmental conditions during migration by comparing yearly April and May tem- peratures between El Niño, La Niña and non-ENSO years using climate data obtained from en route weather stations (En route stations, Figure 1) using Kruskal-Wallis tests. Similarly, we compared El Niño, La Niña and non-ENSO years within the southeastern United States (Gulf Coast stations, Figure 1) to look for evidence of ENSO effects on the Gulf Coast in April and May. We also used Spearmans’ correlations to look for evidence of relationships between yearly April and May temperatures and ENSO index using both temperature data sets (en route and Gulf Coast). We performed all statistical analyses in R version 4.1.2 [54]. We used a linear mixed modeling approach similar to Rockwell et al. [55] where we controlled for individual by entering band number as a random factor because we captured some individuals across multiple years. We used Figure 1. Areas for which we downloaded NCEP temperature data to evaluate en route temperature (En route Stations) effects on timing lme in R package ‘nlme’ [56] to assess the influence of age, and condition of birds arriving at our site in northeastern Pennsylva- April and separately, May temperatures south of the study nia (Study Site) and to evaluate ENSO effects on temperatures along area, ENSO, and in the case of Common Yellowthroat, the Gulf Coast (Gulf Coast Stations). Table 1. Mixed models used to evaluate winter (El Niño Southern Oscillation, ENSO) and en route conditions (April and May temperature) on arrival day and size-corrected body mass. Band number was included as a random effect to control for repeated captures across years. Species Response Variable Model Veery Arrival day Arrival day ~ age + April temperature + May temperature + ENSO Gray Catbird Arrival day Arrival day ~ age + April temperature + May temperature + ENSO Common Yellowthroat Arrival day Arrival day ~ age + sex + April temperature + May temperature + ENSO Veery Size-corrected mass Size-corrected mass ~ age + April temperature + May temperature + ENSO Gray Catbird Size-corrected mass Size-corrected mass ~ age + April temperature + May temperature + ENSO Common Yellowthroat Size-corrected mass Size-corrected mass ~ age + sex + April temperature + May temperature + ENSO 40   Smith et al. age categories we only report parameter estimates for P = 0.48) temperatures using data collected from en route age categories second year (SY – entering first breeding stations. Further, examination of temperature data from season) and after second year (ASY – entering at least the only stations in the southeastern United States (Gulf Coast second breeding season hence older than SY) as birds cat- Stations) revealed no difference in average April (χ = 0.71, egorized as after hatch year (AHY) could have been either df = 2, P = 0.70) or May (χ = 0.43, df = 2, P = 0.81) tem- SY or ASY individuals. peratures between El Niño, La Niña or non-ENSO years. Finally, we found no relationship between ENSO index and April (Spearman’s r = 0.02, n = 18, P = 0.95) nor May (Spearman’s r = 0.18, n = 18, P = 0.48) temperatures. 3 Results 3.1 ENSO 3.2 Extrinsic factors We identified 11 years as either El Niño (2005, 2010, 2016, We found no evidence that temperatures experienced 2019) or La Niña (2006, 2008, 2009, 2011, 2012, 2018, 2021) south of our site during April influenced arrival timing in over the 17 years of this study (2004 – 2019, 2021). We found any species nor did May temperatures south of our study no evidence that ENSO events influenced average temper- area appear to influence arrival timing in Veery (Table 2). atures birds experienced during migration. For example, We did find that Gray Catbird and Common Yellowthroat we found no difference in average April (χ = 0.53, df = 2, migrating in warmer Mays arrived earlier at our site (Table P = 0.772) or May (χ = 0.05, df = 2, P = 0.98) temperatures 2). Within a year later arriving individuals had higher between El Niño, La Niña or non-ENSO years nor did we size-corrected body mass in Gray Catbird and Common find relationships between ENSO and April (Spearman’s r Yellowthroat though arrival day was not associated with = 0.02, n = 18, P = 0.95) or May (Spearman’s r = 0.18, n = 18, size-corrected body mass in Veery (Table 2). Finally, both Table 2. Mixed-model results of the influence of extrinsic factors on arrival day (Ordinal) and size-corrected body mass in landbird species, using habitat in and around Lackawanna State Park, Lackawanna County, northeastern Pennsylvania, during spring migration, 2004 – 2019, 2021 (years 2004, 2012 and 2018 omitted for Veery due to within-year sample sizes - see Methods) Species Response Variable N Β SE F df P Veery April temperature Arrival day 260 -0.268 0.788 0.12 1,43 0.74 May temperature Arrival day 260 -0.700 0.838 0.699 1,43 0.41 Arrival day Size-corrected mass 255 0.018 0.016 1.28 1,42 0.27 April temperature Size-corrected mass 255 -0.169 0.172 0.97 1,42 0.33 May temperature Size-corrected mass 255 0.170 0.180 0.90 1,42 0.35 Gray Catbird April temperature Arrival day 1,158 -0.518 0.316 2.69 1,111 0.104 May temperature Arrival day 1,158 -2.268 0.297 58.17 1,111 < 0.001 Arrival day Size-corrected mass 1,111 0.026 0.001 10.90 1,93 0.001 April temperature Size-corrected mass 1,111 0.109 0.083 1.73 1,93 0.19 May temperature Size-corrected mass 1,111 0.347 0.078 19.72 1,93 < 0.001 Common Yellowthroat April temperature Arrival day 689 -0.144 0.332 0.19 1,68 0.67 May temperature Arrival day 689 -1.727 0.285 36.75 1,68 < 0.001 Arrival day Size-corrected mass 674 0.012 0.004 7.72 1,64 0.007 April temperature Size-corrected mass 674 0.102 0.038 7.05 1,64 0.01 May temperature Size-corrected mass 674 -0.020 0.038 0.28 1,64 0.60 Landbird arrival in northeastern Pennsylvania    41 Gray Catbird (May) and Common Yellowthroat (April) 3.3 Intrinsic factors migrating in warmer temperatures arrived with higher size-corrected body mass (Table 2) though en route tem- Older birds arrived earlier than younger in Veery, Gray peratures were not associated with size-corrected mass in Catbird and Common Yellowthroat (Table 3). We also Veery (Table 2). found older individuals arrived with higher size-cor- We found evidence that Veery arrived later follow- rected mass in Gray Catbird though found no age effect ing La Niña years (F = 6.30, P = 0.004, Figure 2) while on size-corrected mass in Veery nor Common Yellowthroat 2,43 Common Yellowthroat arrived earlier following El Niño (Table 3). Finally, we found evidence that males arrived events relative to non-ENSO years (F = 5.20, P = 0.008, before females in Common Yellowthroat (Table 3) though 2,68 Figure 3). We also found a possible effect of ENSO on Gray found no effect of sex on size-corrected mass at first Catbird (F = 2.98, P = 0.055) with birds arriving fol- capture in Common Yellowthroat (Table 3). 2,111 lowing El Niño winters on ordinal day 137 ± s.e. 0.59, La Niña winters on ordinal day 139 ± se 0.482 and following non-ENSO years on ordinal day 139 ± s.e. 0.55. Finally, we 4 Discussion found no evidence of ENSO effects on size-corrected mass in any species: Veery (F = 2.05, P = 0.14); Gray Catbird 2,42 Timing and condition upon arrival at northerly breed- (F = 0.99, P = 0.37); Common Yellowthroat (F , = 1.52, 2, 93 2 64 ing grounds, both of which may be influenced by events P = 0.23). encountered prior to arrival at the migratory destination, may have reproductive consequences [1-3] such that indi- Table 3. Estimated arrival day and size-corrected body mass by status (age ASY - after second year, SY – second year and in Common Yel- lowthroat sex), for landbird species using habitat in and around Lackawanna State Park, Lackawanna County, northeastern Pennsylvania, during spring migration 2004 – 2019, 2021 (years 2004, 2012 and 2018 omitted for Veery due to within-year sample sizes - see Methods). Parameter estimates are arrival day (Ordinal) or size-corrected body mass (g), post hoc P represents results of Tukey post hoc tests compar- ing three age groups (see Methods). Species Response Variable F df P Status N Parameter SE Post hoc Estimate P Veery Arrival day 15.70 2,43 < 0.001 ASY 124 135 0.983 <0.001 SY 97 143 1.025 Size-corrected mass 3.38 2,42 0.04 ASY 121 30.5 g 0.265 0.129 SY 95 29.7 g 0.267 Gray Catbird Arrival day 28.31 2,111 < 0.001 ASY 341 137 0.612 < 0.001 SY 596 141 0.388 Size-corrected mass 16.81 2,93 < 0.001 ASY 315 35.9 g 0.192 < 0.001 SY 583 34.9 g 0.099 Common Arrival day 22.94 2,68 < 0.001 ASY 282 135 0.483 < 0.001 Yellowthroat SY 185 140 0.513 Size-corrected mass 1.29 2,64 0.28 ASY 274 9.9 g 0.063 0.364 SY 180 9.8 g 0.074 Arrival day 15.49 1,613 < 0.001 Female 238 139 0.449 < 0.001 Male 451 136 0.416 Size-corrected mass 3.11 1,601 0.08 Female 232 9.8 g 0.065 0.079 Male 442 9.9 g 0.049 The overall effect of age was significant though post-hoc comparisons revealed no difference between ASY and SY birds. 42   Smith et al. Figure 2. Arrival day (estimated marginal means) by ENSO effect Figure 3. Arrival day (estimated marginal means) by ENSO effect occurring the prior January – March of the wintering period for Veery occurring the prior January – March of the wintering period for using habitat in and around Lackawanna State Park, Lackawanna Common Yellowthroat using habitat in and around Lackawanna County, northeastern Pennsylvania, during spring migration 2005 – State Park, Lackawanna County, northeastern Pennsylvania, during 2011, 2013 - 2017, 2019, 2021. Whiskers represent ± 1 s.e. Bars with spring migration 2004 – 2019, 2021. Whiskers represent ± 1 s.e. different letters are significantly different at P < 0.02. Bars with different letters are significantly different at P < 0.01. viduals arriving early and/or in superior condition realize cause of these carry over effects [55] because differences higher seasonal reproductive performance. For Veery, in food resource abundance between wintering habitats Common Yellowthroat and possibly Gray Catbird, we are associated with multiple measures of overwintering found that arrival timing at our site was associated with fitness correlates [e.g., 33, 55, 64, 65, 66]. events likely encountered on the wintering grounds. We Precipitation is one ecological factor which plays a key also found that environmental conditions experienced role in determining phytophagous insect abundance [67, en route, as measured by average monthly temperatures 68]. These insects are important food resources for winter- south of our study area, were related to arrival timing ing landbirds [17, 33, 34] and have been identified as one of (Gray Catbird, Common Yellowthroat) and condition (Gray the most important factors determining habitat quality for Catbird, Common Yellowthroat) of birds captured at our wintering migrant songbirds [33, 34]. Relative to xeric hab- northerly site. Finally, we found intrinsic factors such as itats, mesic areas have higher food abundance, permitting sex (Common Yellowthroat) and age to be associated with birds using these areas to more quickly acquire and store arrival time (Veery, Gray Catbird, Common Yellowthroat) energy and nutrients for migration as well as departing and condition (Gray Catbird). earlier from their wintering grounds than individuals from Our results suggesting ENSO effects on arrival timing more xeric habitats [17, 69]. at our study site add to the increasing evidence that eco- ENSO events cause regional variation in precipitation logical factors operating during the non-breeding season and temperature during the winter throughout much of may carry over, affecting individuals and population pro- northern South America, areas within the Caribbean as cesses in subsequent seasons. For example, measures of well as Florida and coastal areas of several Gulf Coast states winter precipitation and primary productivity in Africa [32, 53, 70]. This variation, in turn, may influence habitat have been related to both timing of spring arrival and quality, differentially affecting birds depending upon onset of nesting in European migrants [26, 60]. Further, where they overwinter and what ENSO phase they expe- poor winter habitat quality has been associated with rience. Further, evidence suggests that these differential poor body condition in en route migrants [19, 61] as well effects may influence birds during migration and as they as delayed arrival and poor arrival condition at northerly arrive at the breeding grounds. For example, Paxton et al. stopover and breeding areas [15, 30, 55, 62], later breed- [41] provided evidence that condition of South American ing dates [3, 26], reduced reproductive success [3, 55, 63] wintering species migrating through coastal Alabama was and lower return rates [17]. Evidence suggests variation in influenced by ENSO events encountered on the wintering resource abundance on the wintering grounds as a major grounds while González-Prieto and Hobson [30] found Landbird arrival in northeastern Pennsylvania    43 that condition of both American Redstarts and Palm War- the eastern seaboard (Common Yellowthroat - North and blers (Setophaga palmarum) was enhanced upon arrival South Carolina; Gray Catbird – South Carolina through in Manitoba, Canada during springs preceded by La Niña Connecticut) evidence suggests that Gray Catbirds and events. Further they found that arrival day in Yellow War- Common Yellowthroats breeding in the mid-Atlantic likely blers (Setophaga petechia), Wilson’s Warblers (Cardellina winter in southern Florida, islands in the Caribbean [48, pusilla) and Alder Flycatchers (Empidonax alnorum) was 73] and for Common Yellowthroat, possibly the eastern advanced in years preceded by La Niña events. Horn et al., coast of Central America [74]. El Niño effects in the Gulf [28] found limited evidence that winter ENSO events influ- Coast region [52], Florida and islands in the Caribbean enced body condition (1 of 7 species) and more evidence [70] include above normal precipitation and below normal that ENSO events influenced arrival timing (5 of 7 species) temperatures [53, 75] though the effects seem to end by of landbirds passing through Long Point Bird Observatory March [53, 70]. We unfortunately do not know specifically in southern Ontario. where birds using our site overwinter. However, if they While we found no evidence ENSO influenced body did overwinter in the Gulf region, Florida or in localized condition we did find that Veery delayed arrival in north- areas within the Caribbean-Central American region more eastern Pennsylvania after experiencing La Niña events strongly affected by ENSO, we hypothesize that during the prior winter. Evidence suggests Veery breeding in El Niño years birds experienced higher food abundance, the mid-Atlantic (which includes Pennsylvania) winter enhancing body condition which permitted early winter in central South American [49] which, combined with departure, in turn advancing arrival timing at our north- our results suggest birds captured at our site experienced erly site. Overall, our results are consistent with prior areas in South America where La Niña reduced habitat studies suggesting that local winter environmental condi- quality. We also found evidence that ENSO influenced tions influence condition and wintering ground departure arrival timing in Common Yellowthroat and possibly Gray in migratory birds [17, 30, 69] which in turn carry over to Catbird. Interestingly, even with considerable overlap other phases of the avian annual cycle. of the wintering ranges of the species influenced by La Timing of arrival at our site by Gray Catbird and Niña in the González-Prieto and Hobson [30] study and Common Yellowthroat, as well as arrival condition in Gray Common Yellowthroat and Gray Catbird, our findings Catbird and Common Yellowthroat appeared influenced suggest that birds arriving at our site wintered in areas by factors encountered en route. Our results suggesting where ENSO effects were different than birds captured in that Gray Catbird and Common Yellowthroat migrating in the Gonzalez-Prieto and Hobson [30] study. For example, warmer Mays arrived at our site earlier are similar to the we found that Common Yellowthroat arrived earlier fol- findings of Marra et al. [22] who found, in these species, lowing El Niño winters compared to non-ENSO years. that birds experiencing warmer temperatures en route Regardless, our results, combined with those of Paxton arrived earlier at Long Point Bird Observatory, Ontario et al. [41], Gonzalez-Prieto and Hobson [30] and Horn et Canada. Moreover, our arrival timing results support the al., [28] suggest that ENSO effects encountered during the hypothesis that some species adjust their migration rate to winter influence landbird migrants both early in migra- match annual variation in en route temperatures or asso- tion, when birds are closer to the wintering grounds and ciated factors such as leaf emergence and resource avail- later in migration, when birds are closer to and arriving at ability [6, 21, 22]. Low temperatures restrict emergence northerly breeding grounds and further, that ENSO effects of arthropods at northerly latitudes [6, 30, 76], a primary differentially influence landbirds dependent upon over- food resource during spring migration and low food avail- wintering location. Further, our results (no effect of ENSO ability during colder springs increases thermoregulatory on body condition) support the hypothesis that negative costs during migration [77]. ENSO effects on condition are strongest during the earlier Timing and rate of migration may not be directly stages of spring migration and that birds may be able to regulated by temperature but by factors, such as leaf compensate for poor departure condition while en route emergence and associated resource availability, as well [28]. as other climatic factors resulting from the northward While both Gray Catbird and Common Yellowthroat advancing vernal front [22]. The northward progression have wide wintering ranges [71, 72] encompassing much of spring leaf-out is strongly associated with temperature and may provide important information about habitat of Mexico (Common Yellowthroat) or Caribbean coastal for spring migrating landbirds [22]. Leaf-out is correlated Mexico (Gray Catbird) and the Caribbean-Central America with increases in herbivorous arthropods [6, 78], impor- region as well as throughout the Gulf Coast region of the tant prey for insectivorous landbird migrants and foliage United States, Florida as well as a number of states along 44   Smith et al. may offer camouflage to migrating landbirds from pred- intrinsic factors influenced arrival timing and condition ators [22]. Given the energetic requirements of migration in northeastern Pennsylvania, there are several possi- [79] presence of adequate food resources is critical to max- ble reasons for differential arrival by sex and age. For imizing fat deposition rate [80, 81]. Further, temperature example, evidence suggests males generally precede is associated with an assortment of climatic conditions females to breeding areas (protandry) in an attempt to important to the timing and rate of migration, including obtain the highest quality territories and/or to maximize winds, cold fronts and late season snow storms [37], all of mating opportunities [11, 83, 84] hence males suffer a which may influence migration speed [22, 82]. greater cost than females from arriving late [11]. Further, Our results indicating that size-adjusted body mass younger birds have more time left for future reproduction at first capture of both Gray Catbird and Common Yel- relative to older individuals hence are expected to be more lowthroat was higher for those individuals migrating in risk-averse, delaying arrival [83]. These ultimate causes years with warmer April (Common Yellowthroat) or May for differential arrival may manifest via classes of indi- (Gray Catbird) temperatures are sensible as one would viduals departing wintering areas at different times and/ expect migrants experiencing better conditions en route or migrating different distances [46, 85-87] or at different to be in better condition as warmer temperatures reduce speeds [46, 88, 89]. Further, there is an increasing body of thermoregulatory costs while also providing more arthro- evidence suggesting that age- and sex-related behavioral pods. asymmetries, via influencing winter habitat occupancy, The El Niño-Southern Oscillation may also influence influence departure time [15] and condition from the win- precipitation and temperature patterns within North tering grounds [17] as well as arrival time and condition of America, notably in the southeastern United States [52, arrival to northerly breeding grounds [3, 15, 16]. Another 53]. El Niño events tend to induce above normal precip- reason might be due to age-related experience or domi- itation and below normal temperatures [53, 75] while La nance asymmetries experienced en route. For example, Niña events tend to induce above average temperatures younger, inexperienced individuals (SY birds on their first [52] in the southeastern U.S. Hence, our findings suggest- spring migration) may be less adept at dealing with en ing that ENSO events influenced arrival timing might be route exigencies [2, 14, 55]. Spending more time deposit- a consequence of ENSO events influencing precipitation ing fat may come at a cost with respect to migratory timing and temperature patterns within habitats along the north- [2], in turn delaying arrival at our northerly site. Further, ern Gulf Coast. younger individuals forced into suboptimal habitats [14, We feel our results generally support the effect of 90] may experience conditions, e.g., fewer food resources, ENSO encountered on the wintering grounds rather than that suppress individual condition, both delaying arrival en route for a number of reasons. First, we found no differ- and negatively influencing condition at northerly stopo- ence between La Niña, El Niño and non-ENSO years with ver and breeding locations. Taken together, our results respect to mean temperature, in either April or May (when suggest species-specific responses to how birds deal with most individuals of both species migrate), from weather temperatures en route and that more work examining stations throughout the entire range from which we col- species differences in condition upon arrival at the breed- lected temperature data nor from those stations limited ing grounds is warranted. to the southeastern United States. Further, we found no association between ENSO index and April or May tem- peratures throughout the entire range from which we Acknowledgements collected temperature data nor from those stations in the southeastern United States. Finally, Veery pass through Funding was provided by the Pennsylvania Department the Gulf region in late April to early May [49] while Gray of Conservation and Natural Resource Conservation Catbird and Common Yellowthroat begin migration or Program, Penn State University Scranton and the Univer- pass through the Gulf region in late March to mid-April [72] sity of Scranton. We thank Ms. A. Bushko and Lackawanna and prior work suggests ENSO effects in the Gulf region State Park for permission to capture birds on their proper- typically end in March [but see 53, 75]. ties and are grateful to M. Carey for providing insight into We found that male Common Yellowthroats arrived the ecology of this system. We thank Dr. Andrew Davis before females and that older birds arrived earlier (Veery, and 2 anonymous reviewers whose comments signifi- Gray Catbird, Common Yellowthroat) and with higher cantly improved the manuscript. We thank The University size-corrected body mass (Gray Catbird) than younger of Scranton and Penn State Scranton for supporting our birds. While our results are insufficient to evaluate how work. Finally, B. Smith and numerous University of Scran- Landbird arrival in northeastern Pennsylvania    45 [15] Marra PP, Hobson KA, Holmes RT. Stable-carbon isotopes ton undergraduates contributed to this project. All proce- link winter and summer events in a migratory bird. Science. dures performed in this study complied with the laws and 1998;282:1884-6 regulations within the United States of America, and the [16] Norris DR, Marra PP. Seasonal interactions, habitat quality, and population dynamics in migratory birds. Condor. 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Factors associated with arrival timing and condition of migrant landbird species in northeastern Pennsylvania

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Anim. Migr. 2022; 9: 37–47 Research Article Robert J. Smith*, Margret I. Hatch, Jason M. Graham Factors associated with arrival timing and condition of migrant landbird species in northeastern Pennsylvania https://doi.org/10.1515/ami-2022-0119 1 Introduction received March 29, 2022; accepted July 25, 2022 Timing of spring migration in passerines, and conse- Abstract: Extrinsic and intrinsic factors operating during quently arrival at the breeding grounds has reproductive and prior to the passerine spring migratory period have consequences [1-3] and may be influenced by intrinsic and been associated with both migratory timing and condi- extrinsic factors operating both during passage as well as tion. Here we take advantage of a long-term data set to prior to the migratory event. For example, genetic control answer questions about how extrinsic factors encoun- mechanisms influence onset of migratory restlessness, tered on the wintering grounds (El Niño Southern Oscil- temporal pattern and direction of migratory activity as well lation, ENSO) and en route (temperatures south of our as annual patterns of body mass [4]. Further, within-spe- study site) along with intrinsic factors (age, sex if possi- cies migratory strategies may differ, leading to intra-spe- ble) influenced both arrival timing and condition at our cific differences in timing of passage [5] and arrival at the site in northeastern Pennsylvania. Older birds preceded migratory destination [6]. The classic example of differ- younger, male Common Yellowthroats (Geothlypis trichas) ential passage in North American landbirds is the early preceded females and within a year later arriving Gray Cat- migration and arrival of males to breeding areas [e.g., birds (Dumetella carolinensis) and Common Yellowthroats 7, 8]. Perhaps due to advantages in gainingaccess to the were in better condition. We found that Gray Catbirds and highest quality territories [9, 10 - rank advantage hypoth- Common Yellowthroats migrating during warmer years esis], more mating opportunities [11 - mate opportunity arrived in better condition. Finally, we found evidence that hypothesis], differences in the sex-specific fitness costs ENSO, likely via influencing weather and food availability and benefits of arrival timing [12 - sexual conflict hypoth- during the winter, was associated with arrival timing in esis] or some combination [13]. Finally age and sex-related Veery (Catharus fuscescens), Common Yellowthroats and dominance asymmetries, via influencing en route [14] or possibly Gray Catbirds. Our results support the hypothesis winter habitat occupancy [15, 16], have also been related that events experienced earlier, either between (wintering to timing of arrival in breeding areas as well as reproduc- to migratory periods) or within (earlier vs. later in migra- tive success for that season [3]. tion) phases of the avian annual cycle may carry over, Extrinsic factors have also been linked to timing of influencing fitness later in time or in subsequent phases departure from the wintering grounds [17-20], en route of the annual cycle. passage timing [20-24] and condition (25) as well as timing Keywords: arrival timing; carry-over effects; landbird; of arrival at northerly breeding grounds [6, 21, 26, 27]. spring migration For example, Studds and Marra [17] demonstrated rela- tionships between rainfall, food resources and departure dates of American Redstarts (Setophaga ruticilla) from wintering grounds in Jamaica with birds departing earlier in years with more rainfall and food resources. *Corresponding author: Robert J. Smith, Department of Biology, More recently, wintering ground environmental con- The University of Scranton, Scranton, PA 18510, USA, Email: Robert. ditions, as indicated by the El Niño Southern Oscillation Smith@scranton.edu Margret I. Hatch, Penn State Scranton, 120 Ridge View Drive, Dun- (ENSO), have been related to passage timing [27-29] as more, PA 18512, USA well as energetic condition in spring landbird migrants Jason M. Graham, Department of Mathematics, The University of passing through both southerly [25] and northerly [28, 30] Scranton, Scranton, PA 18510, USA Open Access. © 2022 Smith et al., published by De Gruyter. This work is licensed under the Creative Commons Attribution 4.0 License. 38   Smith et al. stopover sites in North America. ENSO events cause vari- habitat. We captured birds in shrub-dominated habitats ation in precipitation and temperature in parts of South through both the spring migratory period and into the rd nd America and the Caribbean-Central America region [31, breeding season (from the 3 week of April through the 2 32] both of which influence habitat quality for wintering week of June). We delineated the endpoint of spring migra- migrant songbirds [33, 34]. Further, both Hüppop and tion for each species by examining histograms of first cap- Hüppop [35] and Tøttrup et al. [36] found en route temper- tures, determining the date when number of first captures ature to be associated with mean spring passage timing dropped to zero. We operated between 10 (2004, 2011 – in a majority of short and long distance migrants moving 2019, 2021) and 16 (2005 – 2010) permanently positioned through Europe while in North America Marra et al. [22] mist-nets, checking nets at 30 minute intervals. Nets were found median passage dates to be inversely related to en opened by sunrise, remaining open until late-morning. route temperatures in the majority of species examined. We did not capture birds in the event of excessively low Finally, Balbontín et al. [21] found an advance in timing temperatures or rain. For each individual captured we of arrival at breeding grounds for barn swallows (Hirundo recorded species, age and when possible to determine rustica) when ecological conditions en route improved by plumage, sex [39], mass and tarsus length along with while Smith et al. [6] demonstrated relationships between capture date and time. Birds were banded with a U.S. annual variation in environmental conditions and timing Geological Service aluminum leg band and all recaptures of arrival at northerly breeding grounds in American Red- measured without reference to previous records. In all starts. subsequent analyses we only use data collected the first Even as a large body of literature exists linking extrin- time within a year an individual was captured and include sic factors to en route passage timing [6, 21-23, 37] less is only those species for which we captured 10 or more indi- known about how these factors influence en route condi- viduals within a year for at least 80% of the years encom- tion and especially if and how ENSO events taking place passed by this study (veery – 82%, Gray Catbird – 100%, on the wintering grounds influence arrival timing and Common Yellowthroat – 100%). For all individuals within condition of birds at northerly stopover and breeding each species we estimated body condition using the areas [28]. Our objective was to document passage timing Scaled Mass Index of Peig and Green [40], using tarsus and size-corrected body mass at first capture using a long- length to adjust mass as if all individuals within a species term (17 yr) dataset collected during spring migration in were the same size. northeastern Pennsylvania. We tested the hypotheses that We estimated environmental conditions on the win- 1) extrinsic factors encountered both on the wintering tering grounds following the methodology of Paxton et al. grounds (as indicated by ENSO) as well as during migra- [41]. We identified years between 2004 and 2021 as El Niño, tion (en route temperatures), and 2) intrinsic factors (age La Niña or non-ENSO during winter using the National and when possible sex) influenced condition and timing Weather Service Climate Prediction Center (ONI; https:// of arrival at our site in northeastern Pennsylvania, USA. www.cpc.ncep.noaa.gov/products/precip/CWlink/MJO/ enso.shtml mean monthly values of the standardized ENSO conditions (January to March). This index is a measure of departure from long-term average sea surface 2 Methods temperature (SST) in the east-central Pacific Ocean within o o 0 o the Nino 3.4 region (5 N-5 S, 120 -170 W). The index clas- We collected data describing arrival phenology and sifies El Niño and La Niña events based on a threshold size-corrected body mass of the above species in Lacka- of 3 consecutive months above the 0.5 C SST anomaly (El wanna County, Benton Township, northeastern Pennsyl- Niño) or below the -0.5 C SST anomaly (La Niña). vania at both Lackawanna State Park and private lands We investigated the effect of en route conditions on immediately adjacent to the Park from 2004 – 2019, 2021. arrival timing and condition by downloading temperature All netting locations were within 1.8 km of each other. data collected from The National Centers for Environmen- Shrub habitat was approximately 25-35 years post agricul- tal Prediction (NCEP)/National Center for Atmospheric ture and was a mix of exotic (primarily honeysuckle [Lon- Research (NCAR) Reanalysis data set [42] using R package icera spp.]) and native shrubs (primarily dogwood [Cornus RNCEP, version 1.0.7 [43]. This data set is high-quality, well spp.] and prior to infestation by exotic Viburnum Leaf documented, freely available and is increasingly being Beetle (Pyrrhalta viburni) in 2008, arrowwood viburnum used in ecological research [43], including bird migration (Viburnum dentatum) (see 38) as well as a small number [44, 45]. To estimate annual variation in spring tempera- of saplings of most tree species found in nearby forested tures encountered by migrating birds we selected temper- Landbird arrival in northeastern Pennsylvania    39 ature data from areas south of our study site bounded by sex on passage timing. We treated capture day, April and 30.0 to 39.9 N and -85.0 to -78.0 W for April and May, 2004- May temperatures as continuous variables and age, sex 2021 (Figure 1, En route Stations). We selected this area to and ENSO as factors (Table 1). To evaluate size-corrected maximize the likelihood that temperature data reflected body mass at arrival we used a similar modeling approach what the individuals captured at our site experienced though because mass may change with date we included en route as most populations of migrant landbirds tend ordinal date as a numerical variable in the model (Table to follow a north-south axis [46, 47]. Further, modeling 1). For Veery we omitted data from 2004, 2012 and 2018 (Gray Catbird, Common Yellowthroat; 48], geolocator data because we captured less than 10 birds in these years. (Veery; 49) geolocator and recapture data (Gray Catbird; We evaluated assumptions for each model by following 49) and genetic data (Common Yellowthroat; 48, 50, 51) the methodology of Zuur et al. [57]. Using a lme approach support our assumption that our temperature data reflect allowed us to account for heterogeneity and temporal what birds experienced prior to arrival at our site. autocorrelation as necessary [57]. We screened data for Because ENSO events may influence temperature and multicollinearity using function vif from package ‘usdm’ precipitation in the southeastern United States [52, 53] we [58] and used the package ‘emmeans’ [59] to calculate evaluated another NCEP data set, using temperature data parameter estimates and where necessary Tukey post hoc from stations within an area bounded by 25.0 to 32.5 N and tests. While we performed post hoc tests using all three -93.7 to -78 W (Figure 1, Gulf Coast stations). We looked for an influence of winter ENSO on environmental conditions during migration by comparing yearly April and May tem- peratures between El Niño, La Niña and non-ENSO years using climate data obtained from en route weather stations (En route stations, Figure 1) using Kruskal-Wallis tests. Similarly, we compared El Niño, La Niña and non-ENSO years within the southeastern United States (Gulf Coast stations, Figure 1) to look for evidence of ENSO effects on the Gulf Coast in April and May. We also used Spearmans’ correlations to look for evidence of relationships between yearly April and May temperatures and ENSO index using both temperature data sets (en route and Gulf Coast). We performed all statistical analyses in R version 4.1.2 [54]. We used a linear mixed modeling approach similar to Rockwell et al. [55] where we controlled for individual by entering band number as a random factor because we captured some individuals across multiple years. We used Figure 1. Areas for which we downloaded NCEP temperature data to evaluate en route temperature (En route Stations) effects on timing lme in R package ‘nlme’ [56] to assess the influence of age, and condition of birds arriving at our site in northeastern Pennsylva- April and separately, May temperatures south of the study nia (Study Site) and to evaluate ENSO effects on temperatures along area, ENSO, and in the case of Common Yellowthroat, the Gulf Coast (Gulf Coast Stations). Table 1. Mixed models used to evaluate winter (El Niño Southern Oscillation, ENSO) and en route conditions (April and May temperature) on arrival day and size-corrected body mass. Band number was included as a random effect to control for repeated captures across years. Species Response Variable Model Veery Arrival day Arrival day ~ age + April temperature + May temperature + ENSO Gray Catbird Arrival day Arrival day ~ age + April temperature + May temperature + ENSO Common Yellowthroat Arrival day Arrival day ~ age + sex + April temperature + May temperature + ENSO Veery Size-corrected mass Size-corrected mass ~ age + April temperature + May temperature + ENSO Gray Catbird Size-corrected mass Size-corrected mass ~ age + April temperature + May temperature + ENSO Common Yellowthroat Size-corrected mass Size-corrected mass ~ age + sex + April temperature + May temperature + ENSO 40   Smith et al. age categories we only report parameter estimates for P = 0.48) temperatures using data collected from en route age categories second year (SY – entering first breeding stations. Further, examination of temperature data from season) and after second year (ASY – entering at least the only stations in the southeastern United States (Gulf Coast second breeding season hence older than SY) as birds cat- Stations) revealed no difference in average April (χ = 0.71, egorized as after hatch year (AHY) could have been either df = 2, P = 0.70) or May (χ = 0.43, df = 2, P = 0.81) tem- SY or ASY individuals. peratures between El Niño, La Niña or non-ENSO years. Finally, we found no relationship between ENSO index and April (Spearman’s r = 0.02, n = 18, P = 0.95) nor May (Spearman’s r = 0.18, n = 18, P = 0.48) temperatures. 3 Results 3.1 ENSO 3.2 Extrinsic factors We identified 11 years as either El Niño (2005, 2010, 2016, We found no evidence that temperatures experienced 2019) or La Niña (2006, 2008, 2009, 2011, 2012, 2018, 2021) south of our site during April influenced arrival timing in over the 17 years of this study (2004 – 2019, 2021). We found any species nor did May temperatures south of our study no evidence that ENSO events influenced average temper- area appear to influence arrival timing in Veery (Table 2). atures birds experienced during migration. For example, We did find that Gray Catbird and Common Yellowthroat we found no difference in average April (χ = 0.53, df = 2, migrating in warmer Mays arrived earlier at our site (Table P = 0.772) or May (χ = 0.05, df = 2, P = 0.98) temperatures 2). Within a year later arriving individuals had higher between El Niño, La Niña or non-ENSO years nor did we size-corrected body mass in Gray Catbird and Common find relationships between ENSO and April (Spearman’s r Yellowthroat though arrival day was not associated with = 0.02, n = 18, P = 0.95) or May (Spearman’s r = 0.18, n = 18, size-corrected body mass in Veery (Table 2). Finally, both Table 2. Mixed-model results of the influence of extrinsic factors on arrival day (Ordinal) and size-corrected body mass in landbird species, using habitat in and around Lackawanna State Park, Lackawanna County, northeastern Pennsylvania, during spring migration, 2004 – 2019, 2021 (years 2004, 2012 and 2018 omitted for Veery due to within-year sample sizes - see Methods) Species Response Variable N Β SE F df P Veery April temperature Arrival day 260 -0.268 0.788 0.12 1,43 0.74 May temperature Arrival day 260 -0.700 0.838 0.699 1,43 0.41 Arrival day Size-corrected mass 255 0.018 0.016 1.28 1,42 0.27 April temperature Size-corrected mass 255 -0.169 0.172 0.97 1,42 0.33 May temperature Size-corrected mass 255 0.170 0.180 0.90 1,42 0.35 Gray Catbird April temperature Arrival day 1,158 -0.518 0.316 2.69 1,111 0.104 May temperature Arrival day 1,158 -2.268 0.297 58.17 1,111 < 0.001 Arrival day Size-corrected mass 1,111 0.026 0.001 10.90 1,93 0.001 April temperature Size-corrected mass 1,111 0.109 0.083 1.73 1,93 0.19 May temperature Size-corrected mass 1,111 0.347 0.078 19.72 1,93 < 0.001 Common Yellowthroat April temperature Arrival day 689 -0.144 0.332 0.19 1,68 0.67 May temperature Arrival day 689 -1.727 0.285 36.75 1,68 < 0.001 Arrival day Size-corrected mass 674 0.012 0.004 7.72 1,64 0.007 April temperature Size-corrected mass 674 0.102 0.038 7.05 1,64 0.01 May temperature Size-corrected mass 674 -0.020 0.038 0.28 1,64 0.60 Landbird arrival in northeastern Pennsylvania    41 Gray Catbird (May) and Common Yellowthroat (April) 3.3 Intrinsic factors migrating in warmer temperatures arrived with higher size-corrected body mass (Table 2) though en route tem- Older birds arrived earlier than younger in Veery, Gray peratures were not associated with size-corrected mass in Catbird and Common Yellowthroat (Table 3). We also Veery (Table 2). found older individuals arrived with higher size-cor- We found evidence that Veery arrived later follow- rected mass in Gray Catbird though found no age effect ing La Niña years (F = 6.30, P = 0.004, Figure 2) while on size-corrected mass in Veery nor Common Yellowthroat 2,43 Common Yellowthroat arrived earlier following El Niño (Table 3). Finally, we found evidence that males arrived events relative to non-ENSO years (F = 5.20, P = 0.008, before females in Common Yellowthroat (Table 3) though 2,68 Figure 3). We also found a possible effect of ENSO on Gray found no effect of sex on size-corrected mass at first Catbird (F = 2.98, P = 0.055) with birds arriving fol- capture in Common Yellowthroat (Table 3). 2,111 lowing El Niño winters on ordinal day 137 ± s.e. 0.59, La Niña winters on ordinal day 139 ± se 0.482 and following non-ENSO years on ordinal day 139 ± s.e. 0.55. Finally, we 4 Discussion found no evidence of ENSO effects on size-corrected mass in any species: Veery (F = 2.05, P = 0.14); Gray Catbird 2,42 Timing and condition upon arrival at northerly breed- (F = 0.99, P = 0.37); Common Yellowthroat (F , = 1.52, 2, 93 2 64 ing grounds, both of which may be influenced by events P = 0.23). encountered prior to arrival at the migratory destination, may have reproductive consequences [1-3] such that indi- Table 3. Estimated arrival day and size-corrected body mass by status (age ASY - after second year, SY – second year and in Common Yel- lowthroat sex), for landbird species using habitat in and around Lackawanna State Park, Lackawanna County, northeastern Pennsylvania, during spring migration 2004 – 2019, 2021 (years 2004, 2012 and 2018 omitted for Veery due to within-year sample sizes - see Methods). Parameter estimates are arrival day (Ordinal) or size-corrected body mass (g), post hoc P represents results of Tukey post hoc tests compar- ing three age groups (see Methods). Species Response Variable F df P Status N Parameter SE Post hoc Estimate P Veery Arrival day 15.70 2,43 < 0.001 ASY 124 135 0.983 <0.001 SY 97 143 1.025 Size-corrected mass 3.38 2,42 0.04 ASY 121 30.5 g 0.265 0.129 SY 95 29.7 g 0.267 Gray Catbird Arrival day 28.31 2,111 < 0.001 ASY 341 137 0.612 < 0.001 SY 596 141 0.388 Size-corrected mass 16.81 2,93 < 0.001 ASY 315 35.9 g 0.192 < 0.001 SY 583 34.9 g 0.099 Common Arrival day 22.94 2,68 < 0.001 ASY 282 135 0.483 < 0.001 Yellowthroat SY 185 140 0.513 Size-corrected mass 1.29 2,64 0.28 ASY 274 9.9 g 0.063 0.364 SY 180 9.8 g 0.074 Arrival day 15.49 1,613 < 0.001 Female 238 139 0.449 < 0.001 Male 451 136 0.416 Size-corrected mass 3.11 1,601 0.08 Female 232 9.8 g 0.065 0.079 Male 442 9.9 g 0.049 The overall effect of age was significant though post-hoc comparisons revealed no difference between ASY and SY birds. 42   Smith et al. Figure 2. Arrival day (estimated marginal means) by ENSO effect Figure 3. Arrival day (estimated marginal means) by ENSO effect occurring the prior January – March of the wintering period for Veery occurring the prior January – March of the wintering period for using habitat in and around Lackawanna State Park, Lackawanna Common Yellowthroat using habitat in and around Lackawanna County, northeastern Pennsylvania, during spring migration 2005 – State Park, Lackawanna County, northeastern Pennsylvania, during 2011, 2013 - 2017, 2019, 2021. Whiskers represent ± 1 s.e. Bars with spring migration 2004 – 2019, 2021. Whiskers represent ± 1 s.e. different letters are significantly different at P < 0.02. Bars with different letters are significantly different at P < 0.01. viduals arriving early and/or in superior condition realize cause of these carry over effects [55] because differences higher seasonal reproductive performance. For Veery, in food resource abundance between wintering habitats Common Yellowthroat and possibly Gray Catbird, we are associated with multiple measures of overwintering found that arrival timing at our site was associated with fitness correlates [e.g., 33, 55, 64, 65, 66]. events likely encountered on the wintering grounds. We Precipitation is one ecological factor which plays a key also found that environmental conditions experienced role in determining phytophagous insect abundance [67, en route, as measured by average monthly temperatures 68]. These insects are important food resources for winter- south of our study area, were related to arrival timing ing landbirds [17, 33, 34] and have been identified as one of (Gray Catbird, Common Yellowthroat) and condition (Gray the most important factors determining habitat quality for Catbird, Common Yellowthroat) of birds captured at our wintering migrant songbirds [33, 34]. Relative to xeric hab- northerly site. Finally, we found intrinsic factors such as itats, mesic areas have higher food abundance, permitting sex (Common Yellowthroat) and age to be associated with birds using these areas to more quickly acquire and store arrival time (Veery, Gray Catbird, Common Yellowthroat) energy and nutrients for migration as well as departing and condition (Gray Catbird). earlier from their wintering grounds than individuals from Our results suggesting ENSO effects on arrival timing more xeric habitats [17, 69]. at our study site add to the increasing evidence that eco- ENSO events cause regional variation in precipitation logical factors operating during the non-breeding season and temperature during the winter throughout much of may carry over, affecting individuals and population pro- northern South America, areas within the Caribbean as cesses in subsequent seasons. For example, measures of well as Florida and coastal areas of several Gulf Coast states winter precipitation and primary productivity in Africa [32, 53, 70]. This variation, in turn, may influence habitat have been related to both timing of spring arrival and quality, differentially affecting birds depending upon onset of nesting in European migrants [26, 60]. Further, where they overwinter and what ENSO phase they expe- poor winter habitat quality has been associated with rience. Further, evidence suggests that these differential poor body condition in en route migrants [19, 61] as well effects may influence birds during migration and as they as delayed arrival and poor arrival condition at northerly arrive at the breeding grounds. For example, Paxton et al. stopover and breeding areas [15, 30, 55, 62], later breed- [41] provided evidence that condition of South American ing dates [3, 26], reduced reproductive success [3, 55, 63] wintering species migrating through coastal Alabama was and lower return rates [17]. Evidence suggests variation in influenced by ENSO events encountered on the wintering resource abundance on the wintering grounds as a major grounds while González-Prieto and Hobson [30] found Landbird arrival in northeastern Pennsylvania    43 that condition of both American Redstarts and Palm War- the eastern seaboard (Common Yellowthroat - North and blers (Setophaga palmarum) was enhanced upon arrival South Carolina; Gray Catbird – South Carolina through in Manitoba, Canada during springs preceded by La Niña Connecticut) evidence suggests that Gray Catbirds and events. Further they found that arrival day in Yellow War- Common Yellowthroats breeding in the mid-Atlantic likely blers (Setophaga petechia), Wilson’s Warblers (Cardellina winter in southern Florida, islands in the Caribbean [48, pusilla) and Alder Flycatchers (Empidonax alnorum) was 73] and for Common Yellowthroat, possibly the eastern advanced in years preceded by La Niña events. Horn et al., coast of Central America [74]. El Niño effects in the Gulf [28] found limited evidence that winter ENSO events influ- Coast region [52], Florida and islands in the Caribbean enced body condition (1 of 7 species) and more evidence [70] include above normal precipitation and below normal that ENSO events influenced arrival timing (5 of 7 species) temperatures [53, 75] though the effects seem to end by of landbirds passing through Long Point Bird Observatory March [53, 70]. We unfortunately do not know specifically in southern Ontario. where birds using our site overwinter. However, if they While we found no evidence ENSO influenced body did overwinter in the Gulf region, Florida or in localized condition we did find that Veery delayed arrival in north- areas within the Caribbean-Central American region more eastern Pennsylvania after experiencing La Niña events strongly affected by ENSO, we hypothesize that during the prior winter. Evidence suggests Veery breeding in El Niño years birds experienced higher food abundance, the mid-Atlantic (which includes Pennsylvania) winter enhancing body condition which permitted early winter in central South American [49] which, combined with departure, in turn advancing arrival timing at our north- our results suggest birds captured at our site experienced erly site. Overall, our results are consistent with prior areas in South America where La Niña reduced habitat studies suggesting that local winter environmental condi- quality. We also found evidence that ENSO influenced tions influence condition and wintering ground departure arrival timing in Common Yellowthroat and possibly Gray in migratory birds [17, 30, 69] which in turn carry over to Catbird. Interestingly, even with considerable overlap other phases of the avian annual cycle. of the wintering ranges of the species influenced by La Timing of arrival at our site by Gray Catbird and Niña in the González-Prieto and Hobson [30] study and Common Yellowthroat, as well as arrival condition in Gray Common Yellowthroat and Gray Catbird, our findings Catbird and Common Yellowthroat appeared influenced suggest that birds arriving at our site wintered in areas by factors encountered en route. Our results suggesting where ENSO effects were different than birds captured in that Gray Catbird and Common Yellowthroat migrating in the Gonzalez-Prieto and Hobson [30] study. For example, warmer Mays arrived at our site earlier are similar to the we found that Common Yellowthroat arrived earlier fol- findings of Marra et al. [22] who found, in these species, lowing El Niño winters compared to non-ENSO years. that birds experiencing warmer temperatures en route Regardless, our results, combined with those of Paxton arrived earlier at Long Point Bird Observatory, Ontario et al. [41], Gonzalez-Prieto and Hobson [30] and Horn et Canada. Moreover, our arrival timing results support the al., [28] suggest that ENSO effects encountered during the hypothesis that some species adjust their migration rate to winter influence landbird migrants both early in migra- match annual variation in en route temperatures or asso- tion, when birds are closer to the wintering grounds and ciated factors such as leaf emergence and resource avail- later in migration, when birds are closer to and arriving at ability [6, 21, 22]. Low temperatures restrict emergence northerly breeding grounds and further, that ENSO effects of arthropods at northerly latitudes [6, 30, 76], a primary differentially influence landbirds dependent upon over- food resource during spring migration and low food avail- wintering location. Further, our results (no effect of ENSO ability during colder springs increases thermoregulatory on body condition) support the hypothesis that negative costs during migration [77]. ENSO effects on condition are strongest during the earlier Timing and rate of migration may not be directly stages of spring migration and that birds may be able to regulated by temperature but by factors, such as leaf compensate for poor departure condition while en route emergence and associated resource availability, as well [28]. as other climatic factors resulting from the northward While both Gray Catbird and Common Yellowthroat advancing vernal front [22]. The northward progression have wide wintering ranges [71, 72] encompassing much of spring leaf-out is strongly associated with temperature and may provide important information about habitat of Mexico (Common Yellowthroat) or Caribbean coastal for spring migrating landbirds [22]. Leaf-out is correlated Mexico (Gray Catbird) and the Caribbean-Central America with increases in herbivorous arthropods [6, 78], impor- region as well as throughout the Gulf Coast region of the tant prey for insectivorous landbird migrants and foliage United States, Florida as well as a number of states along 44   Smith et al. may offer camouflage to migrating landbirds from pred- intrinsic factors influenced arrival timing and condition ators [22]. Given the energetic requirements of migration in northeastern Pennsylvania, there are several possi- [79] presence of adequate food resources is critical to max- ble reasons for differential arrival by sex and age. For imizing fat deposition rate [80, 81]. Further, temperature example, evidence suggests males generally precede is associated with an assortment of climatic conditions females to breeding areas (protandry) in an attempt to important to the timing and rate of migration, including obtain the highest quality territories and/or to maximize winds, cold fronts and late season snow storms [37], all of mating opportunities [11, 83, 84] hence males suffer a which may influence migration speed [22, 82]. greater cost than females from arriving late [11]. Further, Our results indicating that size-adjusted body mass younger birds have more time left for future reproduction at first capture of both Gray Catbird and Common Yel- relative to older individuals hence are expected to be more lowthroat was higher for those individuals migrating in risk-averse, delaying arrival [83]. These ultimate causes years with warmer April (Common Yellowthroat) or May for differential arrival may manifest via classes of indi- (Gray Catbird) temperatures are sensible as one would viduals departing wintering areas at different times and/ expect migrants experiencing better conditions en route or migrating different distances [46, 85-87] or at different to be in better condition as warmer temperatures reduce speeds [46, 88, 89]. Further, there is an increasing body of thermoregulatory costs while also providing more arthro- evidence suggesting that age- and sex-related behavioral pods. asymmetries, via influencing winter habitat occupancy, The El Niño-Southern Oscillation may also influence influence departure time [15] and condition from the win- precipitation and temperature patterns within North tering grounds [17] as well as arrival time and condition of America, notably in the southeastern United States [52, arrival to northerly breeding grounds [3, 15, 16]. Another 53]. El Niño events tend to induce above normal precip- reason might be due to age-related experience or domi- itation and below normal temperatures [53, 75] while La nance asymmetries experienced en route. For example, Niña events tend to induce above average temperatures younger, inexperienced individuals (SY birds on their first [52] in the southeastern U.S. Hence, our findings suggest- spring migration) may be less adept at dealing with en ing that ENSO events influenced arrival timing might be route exigencies [2, 14, 55]. Spending more time deposit- a consequence of ENSO events influencing precipitation ing fat may come at a cost with respect to migratory timing and temperature patterns within habitats along the north- [2], in turn delaying arrival at our northerly site. Further, ern Gulf Coast. younger individuals forced into suboptimal habitats [14, We feel our results generally support the effect of 90] may experience conditions, e.g., fewer food resources, ENSO encountered on the wintering grounds rather than that suppress individual condition, both delaying arrival en route for a number of reasons. First, we found no differ- and negatively influencing condition at northerly stopo- ence between La Niña, El Niño and non-ENSO years with ver and breeding locations. Taken together, our results respect to mean temperature, in either April or May (when suggest species-specific responses to how birds deal with most individuals of both species migrate), from weather temperatures en route and that more work examining stations throughout the entire range from which we col- species differences in condition upon arrival at the breed- lected temperature data nor from those stations limited ing grounds is warranted. to the southeastern United States. Further, we found no association between ENSO index and April or May tem- peratures throughout the entire range from which we Acknowledgements collected temperature data nor from those stations in the southeastern United States. Finally, Veery pass through Funding was provided by the Pennsylvania Department the Gulf region in late April to early May [49] while Gray of Conservation and Natural Resource Conservation Catbird and Common Yellowthroat begin migration or Program, Penn State University Scranton and the Univer- pass through the Gulf region in late March to mid-April [72] sity of Scranton. We thank Ms. A. Bushko and Lackawanna and prior work suggests ENSO effects in the Gulf region State Park for permission to capture birds on their proper- typically end in March [but see 53, 75]. ties and are grateful to M. Carey for providing insight into We found that male Common Yellowthroats arrived the ecology of this system. We thank Dr. Andrew Davis before females and that older birds arrived earlier (Veery, and 2 anonymous reviewers whose comments signifi- Gray Catbird, Common Yellowthroat) and with higher cantly improved the manuscript. We thank The University size-corrected body mass (Gray Catbird) than younger of Scranton and Penn State Scranton for supporting our birds. While our results are insufficient to evaluate how work. Finally, B. Smith and numerous University of Scran- Landbird arrival in northeastern Pennsylvania    45 [15] Marra PP, Hobson KA, Holmes RT. Stable-carbon isotopes ton undergraduates contributed to this project. All proce- link winter and summer events in a migratory bird. Science. dures performed in this study complied with the laws and 1998;282:1884-6 regulations within the United States of America, and the [16] Norris DR, Marra PP. Seasonal interactions, habitat quality, and population dynamics in migratory birds. Condor. 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Journal

Animal Migration – de Gruyter

Published: Jan 1, 2022

Keywords: arrival timing; carry-over effects; landbird; spring migration

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