Distribution and foraging patterns of common loons on Lake Michigan with implications

for exposure to type E avian botulism

Kevin P. Kenow a,*, Steven C. Houdek a, Luke J. Fara a, Brian R. Gray a, Brian R. Lubinski b,

Darryl J. Heard c, Michael W. Meyer d, Timothy J. Fox a, Robert J. Kratt a

a
U.S. Geological Survey, Upper Midwest Environmental Sciences Center, 2630 Fanta Reed

Road, La Crosse, WI 54603, USA

b
U.S. Fish and Wildlife Service, 5600 American Blvd. West, Suite 990, Bloomington, MN

55437, USA

c
College of Veterinary Medicine, University of Florida, 2015 SW 16th Ave, Gainesville, FL

32610, USA

d
Wisconsin Department of Natural Resources, 107 Sutliff Avenue, Rhinelander, WI 54501,

USA

*
Corresponding author. kkenow@usgs.gov; Tel: 1 608 781 6278; Fax: 1 608 783 6066.

© 2018. This manuscript version is made available under the Elsevier user license
http://www.elsevier.com/open-access/userlicense/1.0/
Abstract

Common loons (Gavia immer) staging on the Great Lakes during fall migration are at risk to

episodic outbreaks of type E botulism. Information on distribution, foraging patterns, and

exposure routes of loons are needed for understanding the physical and ecological factors that

contribute to avian botulism outbreaks. Aerial surveys were conducted to document the

spatiotemporal distribution of common loons on Lake Michigan during falls 2011-2013. In

addition, satellite telemetry and archival geolocator tags were used to determine the distribution

and foraging patterns of individual common loons while using Lake Michigan during fall

migration. Common loon distribution observed during aerial surveys and movements of

individual radiomarked and/or geotagged loons suggest a seasonal pattern of use, with early fall

use of Green Bay and northern Lake Michigan followed by a shift in distribution to southern

Lake Michigan before moving on to wintering areas. Common loons tended to occupy offshore

areas of Lake Michigan and, on average, spent the majority of daylight hours foraging. Dive

depths were as deep as 60 m and dive characteristics suggested that loons were primarily

foraging on benthic prey. A recent study concluded that round gobies (Neogobius

melanostomus) are an important prey item of common loons and may be involved in

transmission of botulinum neurotoxin type E. Loon distribution coincides with the distribution

of dreissenid mussel biomass, an important food resource for round gobies. Our observations

support speculation that energy transfer to higher trophic levels via gobies may occur in deep-

water habitats, along with transfer of botulinum neurotoxin.

Key words: archival geolocator tag, avian botulism, common loon, foraging patterns, Lake
Michigan, satellite telemetry

2
Introduction

The Great Lakes afford an important resource to migrating and wintering waterbirds by

providing abundant resting and foraging opportunities. However, little information exists on the

pelagic distribution and abundance of waterbirds on Lake Michigan during migration and

wintering periods. Resource managers seek information on Great Lakes waterbird

concentrations during migration and winter as they deal with several priority conservation issues,

including impact assessment of near-shore and off-shore wind turbine placement,

characterization of sea duck wintering distribution and population status, and elucidating factors

that influence the outbreak of type E avian botulism caused by exposure to Clostridium

botulinum neurotoxin type E (BoNT/E). All of these issues and more require a better

understanding of the distribution, abundance, and temporal use patterns of waterbirds.

Episodic outbreaks of type E botulism have resulted in die-offs of waterbirds in the Great Lakes

since at least the 1960s, but outbreaks have become more common and widespread since 1999,

particularly in Lakes Michigan and Erie (Riley et al., 2008). Type E botulism has likely been

responsible for nearly 100,000 waterbird mortalities on the Great Lakes (Chipault et al., 2015),

and extensive bird mortality in northern Lake Michigan near the Sleeping Bear Dunes National

Lakeshore has caused great concern among resource managers and the public. Common loons

(Gavia immer) made up a substantial proportion of these waterbird mortalities associated with

avian botulism outbreaks (Brand et al., 1988; Chipault et al., 2015). The actual sites of BoNT/E

exposure for birds remain unknown. The physical and ecological factors that lead to type E

botulism outbreaks are poorly understood; but invasive species, such as dreissenid mussels

(Dreissena bugensis and D. polymorpha) and round gobies (Neogobius melanostomus), may

serve to mobilize the toxin from lake-bed sediments to fish-eating waterbirds and other aquatic

3
species (Getchell and Bowser, 2006; Lafrancois et al., 2011). An understanding of feeding

patterns and exposure routes of sentinel waterbird species historically at risk to type E botulism

die-offs, such as the common loon, is central to developing ecological studies to assess pathways

of BoNT/E exposure through aquatic food chains in the Great Lakes and identification of

physical and biological linkages that drive botulism outbreaks.

To address the information needs of resource managers, we 1) documented the fall distribution

and relative abundance of common loons using selected areas of Lake Michigan via aerial

surveys during the falls of 2011 through 2013 and 2) monitored movements and foraging

patterns of a sample of common loons breeding in Wisconsin, Minnesota, and Michigan, fitted

with satellite transmitters and/or archival geolocator tags, to determine spatial and temporal

patterns in their distribution and abundance during fall migration with an emphasis on staging on

the Great Lakes.

Methods

Aerial surveys

We surveyed fixed-width transects to determine waterbird distribution and estimate relative

density. Survey areas were established to document the fall distribution and foraging patterns of

waterbirds at-risk to type E botulism on areas of Lake Michigan that have been associated with

past type E botulism waterbird mortality events and are also based on partner (U.S. Fish and

Wildlife Service and Great Lakes Commission) information needs relevant to potential wind

energy development and wintering sea duck distribution. Fixed-width transects were spaced at

4.8 km intervals and extended up to 32 km offshore so as to include waters with depth up to 80

4
m. Transects generally paralleled shorelines to maximize efficiency and safety. Transects were

established using snapPLAN software (TRACK’AIR Aerial Survey Systems, The Netherlands).

Surveys were flown at an average ground speed of about 220 km/h at an altitude of about 61-76

m above the water using a U.S. Fish and Wildlife Service fix-winged aircraft (Partenavia P68

Observer 2). Two trained observers, one on each side of the plane, identified and tallied

waterbirds within 200 m-wide strip transects on either side of the plane. Each observation was

linked to a GPS waypoint and recorded using an integrated GPS voice recording system (Hodges

and Thorpe, 2002).

Waterbird survey data were transcribed from audio files to Excel spreadsheets. Excel files

containing location coordinates and waterbird counts were imported into ArcGIS and converted

into an ESRI Shapefile of observation points. The Batch Kernel Density Tool (automates

density estimates by species and by search radii; Fox, 2018a) was used to create surfaces of

density of observed loons based on a kernel distribution function estimation (ArcGIS, ESRI,

Redlands, CA, USA). The analysis parameters used were search radius of 10 km, output cell

size of 100 m, density units of observed birds per square kilometer, and mask layer of open water

within the study area. It is important to note that the distribution maps do not reflect absolute

density of loons, but rather provide a relative density based on observed birds along the transect

observation area (i.e., loons that occur between the transect strips are not included in the density

estimate). Surveys were typically conducted roughly once per month during fall migration

(September through December).

Tracking movements of individuals using satellite telemetry and archival geolocator tags

5
Breeding common loons tracked for the movement and foraging pattern study were obtained

from lakes in central and northern Minnesota and Wisconsin and the Upper Peninsula of

Michigan during summers 2009-2012 (Figure 1). Lakes with loon territories considered for

inclusion in the study were selected in consultation with Minnesota Department of Natural

Resources (DNR) and Wisconsin DNR personnel. Loon territories were monitored for nesting

activity and breeding success. Adult loons that successfully produced chicks and the resulting

juveniles were captured using night-lighting techniques (Evers, 1993). Each loon was marked

with an aluminum numbered U.S. Fish and Wildlife Service band and a unique combination of

colored leg-bands to aid with future field identification of individuals. The sex of individual

loons of each territorial pair was determined by noting which loon yodeled, a call made only by

male loons (McIntyre, 1988). Both adults of a territorial pair were fitted with archival geolocator

tags (Model LAT 2500; 34.6 x 8.3 mm, 4.4 g; Lotek Wireless Inc.). A combination of adhesive

and plastic cable ties were used to affix the geolocator tag to a modified lock-on aluminum leg

band (Figure 2). Geolocator tags were attached to 176 common loons (Table 1). Tags were

programmed to collect 1) daily location estimates for up to two years, 2) temperature (0.02 °C

accuracy, ≤ 0.05 °C resolution) at 30-min intervals, and 3) pressure data (±1% accuracy, 0.05%

resolution) at 20-sec intervals during daylight hours to document foraging patterns (dive profiles)

during fall migration and during the first few weeks after arriving on the wintering grounds.

Data stored on geolocator tags were not transmitted, requiring that the marked loon be recaptured

to recover the tag and download the data. The geolocator tags were capable of storing data for

several years before the devices needed to be downloaded.

Radiomarked loons were incorporated into the study to provide fine-resolution location data for a

sample of the adult male common loons fitted with geolocator tags. Satellite transmitters (e.g.,

6
Model PTT-100, Microwave Telemetry, Inc) were implanted in 31 adult male common loons

that were captured on breeding lakes in Minnesota, Wisconsin, and the Upper Peninsula of

Michigan during July 2010 and July 2011 (Table 1). Transmitters were surgically implanted in

the abdominal cavity following procedures developed by Korschgen et al. (1996). Surgical

techniques and the handling and care of loons were done under approval of the Animal Care and

Use Committee of the Upper Midwest Environmental Sciences Center and complied with the

Animal Welfare Act (Public Law 99-198 and 9 CFR Parts 1, 2, and 3). The surgical procedures

were conducted in a portable laboratory by Dr. Darryl Heard, College of Veterinary Medicine,

University of Florida, Gainesville, Florida. After surgery, loons were held until demonstrating

control of head and neck and an ability to assume an alert posture.

The satellite transmitters were programmed to transmit on a variable schedule based on the

anticipated stage of migration during the loon’s annual cycle - 8 hours on: 72 hours off during

the breeding season, 8 hours on: 24 hours off while migrating, 6 to 8 hours on: 96 hours off on

the wintering grounds, 8 hours on: 24 hours off during spring migration, and 8 hours on: 96

hours off for the remaining life of the transmitter. Loon locations were estimated from the

Doppler shift in the transmitter carrier frequency and provided by the Argos system (CLS

America, Lanham, Maryland). Location estimates were acquired using Argos Standard Service

Processing (Argos Location Classes [LC] 3, 2, 1, and 0) and Auxiliary Location Processing (LC

A, B, and Z). One standard deviation of nominal accuracy for location estimates with LC 3, 2, 1,

and 0 are <250, 250 to 500, 500 to 1,500, and >1,500 m, respectively (Argos, 2016). We utilized

the most accurate locations per 8-hour transmission period for each individual to describe the

daily location of that adult loon.

7
Geotagged adult loons were recaptured during subsequent breeding seasons using diurnal lift-net

trap and night-light nest-capture techniques (Kenow, 2009) and night-lighting during chick

rearing (Evers, 1993). Geolocator tag data collected over the previous year(s) were downloaded

from tags using LAT Viewer Studio software (Lotek Wireless Inc.). Geolocator tag location

estimates were determined by light-based geolocations using the template-fitting approach

(Exstrom, 2004), in combination with tag temperature (water surface temperature) and pressure

(dive depth) data. Template-fit error estimates were used to filter aberrant geolocation estimates.

Sea surface temperature (derived from NASA Moderate Resolution Imaging Spectroradiometer

[MODIS] imagery) across North America inland lakes, Atlantic coastal waters, and the Gulf of

Mexico, coupled with diving depth information were used to improve or obtain location

estimates and timing of migration movements when light-based geolocation estimates were

unreliable. The Composite Raster and Divergence Tool (Fox, 2018b) was used to facilitate this

process. Geotag temperature data also provided indication of major flights (characterized by

prolonged drop in temperature). Flight times were used to set bounds on the distances of

migration events. Location estimates were used to determine gross movement patterns and

generalized location (i.e., Green Bay, northern Lake Michigan, southern Lake Michigan) of

loons while using Lake Michigan. Green Bay is separated from the rest of Lake Michigan by the

Door Peninsula (WI), the Garden Peninsula (MI), and the chain of islands between them; we

designated the separation of northern and southern Lake Michigan by a line extending from

Rawley Point (WI) to Big Sable Point (MI) as depicted in the third panel of Figure 3.

Interpretation of dives from geotag pressure data

Attributes of dives (i.e., proportion of time underwater, depth of dive, ascent and descent rates,

duration of dive, time duration within 2 m of maximum depth, post-dive surface interval) were

8
extracted from pressure data, where depth (m) = pressure (dbars) * 1.019716

(http://www.seabird.com/document/an69-conversion-pressure-depth), using SAS (SAS, 2015)

software code developed by the first author (Kenow, 2018). Dives were considered to be

submergence below the water surface, which we define as recorded depth of >0.8 m (typical

body length of adult common loons in this study). Because pressure data were collected at 20-sec

intervals, we interpolated the times of both dive initiation and surfacing. Average ascent and

descent rates were estimated individually for each loon and used to interpolate these times,

where rates were based on median values derived from dives >47.25 m (n = 1,214). The descent

rate used for interpolation was 1.056 m/sec and the ascent rate was 1.605 m/sec.

Previous studies of common loon foraging behavior on breeding lakes have considered a break in

foraging bouts to occur when the post-dive surface interval exceeded 120 sec (Strong and

Bissonette, 1989; Nocera and Burgess, 2002). While this criterion may have been applicable to

loons foraging on breeding lakes, a cursory examination of common loon dive data while

foraging in deep waters (>20 m) of Lake Michigan indicated that post-dive surface interval of

120 sec did not provide an acceptable bout ending criteria. Consequently, we defined dive bouts

when sequential dives were separated by post-dive surface interval >360 sec (based on

inspection of dive profiles and evaluation of bout ending criteria estimates for a subsample of

loons per sequential differences analysis [per program diveMove; Luque, 2007]).

Dive profile patterns provided information on the vertical foraging patterns of loons marked with

geolocator tags and, in the case of radiomarked loons, were coupled with bathymetric data

(National Geophysical Data Center, 1996) using ArcGIS software (ESRI 2015). Spot

measurements of water depth and distance to shore were determined for each radiomarked loon

location. Only location estimates for LC qualities 1-3 were used in the analyses.

9
Statistical Analyses

Associations of depth of dive, dive duration, log-transformed duration at maximum depth, and

log-transformed post-dive surface interval with the factors area within Lake Michigan and sex ×

presence/absence of a radio transmitter were estimated using linear models. Models were fitted

using restricted maximum likelihood and SAS’ generalized linear mixed modelling procedure

(PROC GLIMMIX; SAS, 2015). Reported means represent marginal means for balanced

populations while geometric means represent exponentiations of marginal means. Unless

otherwise specified, means for duration at maximum depth and post-dive surface intervals

represent geometric means. Error rates for multiple comparison tests were addressed using

Scheffe’s method (Scheffé, 1953; Snedecor and Cochran, 1989).

Results

Aerial waterbird surveys

Common loon distribution and abundance among selected portions of Lake Michigan are

illustrated in Figure 3. The areas of Lake Michigan covered during each survey varied

considerably depending on objectives of the particular survey, weather conditions, and

availability of pilot and plane. To better illustrate patterns of common loon occurrence, the

distribution maps represent a consolidation of data by month over a three-year period, 2011-

2013. We observed that common loons generally used Green Bay and the northern end of Lake

Michigan during the early portion of fall migration (September through early October). Highest

densities of loons during this early period occurred in central Green Bay and in the main basin of

Lake Michigan off the northeast end of Door County, Wisconsin, stretching north along the

Garden Peninsula and into Manistique Bay. As the season progressed, the distribution of

10
common loons shifted to the southern end of the lake by mid- to late-November. The highest

densities of loons in southern Lake Michigan occurred in waters between 20 and 60 m deep.

Few common loons were observed during surveys conducted during early- to mid-December and

were primarily located in southern Lake Michigan.

Most observations of common loons during aerial surveys consisted of single loons (1,743 of

2,564 observations; 68%). Observed flock size varied from 1 to 35 loons; 90% of the flocks

contained ≤ 3 loons, and 95% of the flocks contained ≤ 5 loons. Flock sizes of >5 loons

occurred primarily in Green Bay and northern Lake Michigan. Of the total number of common

loons tallied, 37% (1,743/4,766) occurred as singles.

Movements of marked common loons and their use of Lake Michigan

Twenty-seven radiomarked loons were tracked during all or a portion of their migration to

wintering grounds (Electronic Supplementary Material (ESM) Table S1); one radiomarked loon

died of unknown causes on its breeding lake (Loon ID 107260), and three transmitters failed

prior to migration (Loon ID 107276) or provided limited location data during migration (Loon

IDs 55482, 107265). Of these 27 loons, 26 used some portion of Lake Michigan during

migration. Among loons using Lake Michigan, one loon succumbed to aspergillosis (Loon ID

55482) on Green Bay, and signals were lost from the transmitters of two loons (Loon IDs 55485,

55487).

We recovered 94 geolocator tags from among 114 of the geotagged loons through recapture or

carcass recovery. Hence, loon recapture/recovery and geolocator tag retention percentages were

65% and 82%, respectively. Sixty-two tags recovered from adult loons without radio

transmitters (28 male, 33 female, and 1 unknown sex) provided data to document movement

11
during fall migration (ESM Table S2). Of these 62 loons, 60 used one or more of the Great

Lakes and 59 used some portion of Lake Michigan during migration.

Departure dates from lakes on which the radiomarked and geotagged loons held breeding

territories were variable among the 89 loons that provided migration data, and varied from 24

July to 26 November (Figure 4). Upon departure from breeding lakes, several radiomarked

and/or geotagged loons were documented at Minnesota, Wisconsin, and Michigan lakes before

migrating to the Great lakes (ESM Tables S1 and S2).

The migration route of most loons (85 of 89) included Lake Michigan. Common loons using

Lake Michigan initially arrived in Green Bay (22), northern Lake Michigan (22), or southern

Lake Michigan (41). Most of the loons arriving at Green Bay or northern Lake Michigan (25 of

44) made their way to southern Lake Michigan before departing to wintering areas (Figure 5).

Green Bay was used by 22 of the marked loons, northern Lake Michigan by 34 loons, and

southern Lake Michigan by 66 loons. The average length of stay on Lake Michigan (including

Green Bay) was about 27 days and varied from <1 to 108 days. Length of stay was negatively

associated with arrival date on Lake Michigan (linear model: slope = -0.81 [SE 0.05], r2 = 0.77,

p < 0.0001, n = 81). For example, those loons arriving to Lake Michigan before 1 October

stayed an average of 68 days (SD 30 days, n = 20), whereas those arriving after 1 November

stayed an average of 8 days (SD 6 days, n = 42).

The median date of departure of marked common loons from Lake Michigan was 23 November,

range was 4 September to 11 December (Figure 4). Of the 85 loons using Lake Michigan, 66

(78%) wintered in the Gulf of Mexico, 12 (14%) wintered in the Atlantic Ocean, two (2%)

wintered in reservoirs in Indiana or Kentucky, signals of three (4%) were lost en route, and two

12
(2%) were confirmed to have died en route to wintering areas. Of 84 loons that completed fall

migration to wintering areas (this includes 80 that used Lake Michigan and 4 that did not use

Lake Michigan during fall migration), 69 (82%) wintered in the Gulf of Mexico, 13 (15%)

wintered in offshore waters of the Atlantic Ocean, and two (2%) wintered in Indiana or Kentucky

reservoirs (Figure 5).

Location estimates (LC ≥ 1) for radiomarked common loons on Lake Michigan were distributed

1.7 to 39.2 km offshore (Figure 6). Most (95%) location estimates were greater than 2.8 km

offshore while 50% of locations were greater than 10.5 km offshore. The average distance to the

nearest shore varied from 7.8 km in Green Bay to 9.5 km in northern Lake Michigan to 19.6 km

in southern Lake Michigan (Table 2). Water depths at sites occupied by radiomarked loons

varied from 6.2 to 250.3 m while depth was greater than 32.4 m at 50% of locations.

Foraging patterns of common loons using Lake Michigan during fall migration

Dive analyses were based on 133,256 dives of 53 geotagged loons that used Lake Michigan for ≥

1 day during fall migration (Table 3). Proportion of time loons were recorded as underwater

during the course of the day reflects a pattern of foraging activity during daylight hours (Figure

7). While the proportion of time does not appear to vary appreciably among locations within

Lake Michigan in non-radiomarked loons, radiomarked loons (all male) appeared to spend less

time underwater; most markedly in the main part of Lake Michigan (both northern and southern

areas). The median hourly proportion of time spent underwater among radiomarked loons during

midday (0900 to 1600 hr) was 0.25 (0.07, 0.48 [5th and 95th quantiles, respectively]), compared

to median values of 0.34 (0.15, 0.57) for non-radiomarked females, and of 0.34 (0.11, 0.59) for

non-radiomarked males. Among non-radiomarked loons, median proportions of time underwater

13
were 0.37 (0.08, 0.58) in Green Bay, 0.34 (0.15, 0.61) in northern Lake Michigan, and 0.34

(0.14, 0.58) in southern Lake Michigan. Over the course of the fall season, proportions of time

underwater increased among loons using northern Lake Michigan in all three sex-radio groups

(Figure 8). A depressed level of time spent underwater was again apparent in radiomarked

loons.

Common loon dives were as deep as 43 m in Green Bay, 56 m in northern Lake Michigan, and

60 m in southern Lake Michigan and exhibited bimodal (southern Lake Michigan) and multi-

modal (Green Bay and northern Lake Michigan) distributions (Figure 9). Mean dive depths

varied significantly among Lake Michigan area × sex-radio categories (F4,10 = 144.17, p <0.001).

Mean dive depth for females in Green Bay was shallower than that for dive depth of non-

radiomarked males in Green Bay while mean dive depths by males in Green Bay did not clearly

differ by radiomarking status (Table 4). Mean dive depths of females were deeper in southern

Lake Michigan relative to Green Bay and northern Lake Michigan, while mean dive depths of

non-radiomarked males were shallower in northern Lake Michigan than in Green Bay and

southern Lake Michigan (Table 5). The mean dive depth of radiomarked males was deeper in

southern Lake Michigan than in Green Bay and northern Lake Michigan. Averaged over sex-

radio effects, mean dive depth among loons using southern Lake Michigan exceeded those in

Green Bay and northern Lake Michigan (Table 4).

As with dive depths, mean duration of dives varied significantly among Lake Michigan area ×

sex-radio categories (F4,10 = 138.72, p < 0.0001). Mean dive durations did not vary significantly

among sex-radio categories within or among areas of Lake Michigan (Table 4). For all sex-radio

categories, mean dive duration differed between southern Lake Michigan and Green Bay and

northern Lake Michigan (Table 5). Mean dive duration was longer in southern Lake Michigan

14
than in Green Bay and northern Lake Michigan (averaged over sex-radio effects; Table 4). Dive

duration was strongly associated with dive depth (Figure 10). For example, the mean duration of

40 (± 5)-m dives was roughly 1.6-times longer than the mean duration of 20 (± 5)-m dives

among non-radiomarked male loons using southern Lake Michigan (mean = 136 sec [133, 140]

versus 87 sec [83, 90]).

Common loon dive shapes were characterized by a relatively long duration at maximum depth

(here defined as within 2 m of maximum depth) relative to the duration of the entire dive. The

dive shape of most dives (82%) met the criterion of Schreer et al. (2001) of a “square” shape,

where time at maximum depth was ≥50% of the dive duration. Mean log-transformed duration

of dive at maximum depth varied significantly among Lake Michigan × sex-radio categories

(F4,10 = 236.78, p < 0.0001). For non-radiomarked loons, mean durations at maximum depth

were longer in southern Lake Michigan than in Green Bay, and, for females only, than in

northern Lake Michigan (Tables 4 and 5). Mean durations at maximum depth were significantly

longer among radiomarked loons in southern Lake Michigan and Green Bay than in northern

Lake Michigan (Tables 4 and 5). Mean durations at maximum depth for males did not vary

significantly by radiomarking status. Averaged over sex-radio effects, mean dive duration at

maximum depth among loons using southern Lake Michigan (52 sec) was longer than in Green

Bay (43 sec) and northern Lake Michigan (41 sec; Table 4). Duration at maximum depth

generally increased with depth of dive (Figure 11). For example, the mean duration at maximum

depth of 40 (± 5)-m dives was 1.4-times longer than the mean duration of 20 (± 5)-m dives

among non-radiomarked male loons using southern Lake Michigan (mean = 71 sec [65, 76]

versus 52 sec [48, 56]).

15
Mean log-transformed duration of post-dive surface intervals also varied significantly among

Lake Michigan × sex-radio categories (F4,10 = 88.39, p < 0.0001). Within sex-radio categories,

post-dive duration means were variable among areas of Lake Michigan (Table 4). For females,

post-dive surface interval means in southern Lake Michigan exceeded those in northern Lake

Michigan and Green Bay (Table 5). For males without transmitters, mean post-dive surface

intervals in Green Bay > southern Lake Michigan > northern Lake Michigan (Table 5). For

radiomarked males, mean post-dive surface interval in southern Lake Michigan > Green Bay >

northern Lake Michigan. Mean post-dive surface interval for radiomarked males in southern

Lake Michigan was longer than non-radiomarked males in northern Lake Michigan (differences

in means = 36.1 sec, with ratio of means 1.68 [1.06, 2.66]). Post-dive surface interval means

varied significantly with both depth of dive (Figure 12) and dive duration. For example, mean

post-dive surface interval of 40 (± 5)-m dives with a dive duration of 150 sec was about 2.2-

times longer than the mean post-dive surface interval of 20 (± 5)-m dives with a dive duration of

120 sec among non-radiomarked male loons using southern Lake Michigan (mean = 105 sec [95,

115] versus 48 sec [44, 53]).

Discussion

Common loon distribution on Lake Michigan

Lake Michigan appears to be a critical staging area for common loons that breed among lakes in

the Northern Lakes and Forests and North Central Hardwood Forests ecoregions of the United

States (U.S. EPA, 2013) based on the large proportion of marked loons that used Lake Michigan

during fall migration. Distribution observed during aerial surveys (Figure 3) and movements of

individual radiomarked and/or geotagged loons (Figures 4 and 5) suggest an analogous seasonal

16
pattern of use, with early fall use of Green Bay and northern Lake Michigan followed by a shift

in distribution to southern Lake Michigan. Common loons occupied fairly deep areas of Lake

Michigan (Figure 13) and generally occurred in waters well offshore (Figure 14 and Figure 6).

Median water depths at locations of radiomarked loons were 27.3 m among loon locations within

Green Bay and about 50 m among loons using the main portion of Lake Michigan. Median

distance from shore varied from 7.1 km in Green Bay to 20.4 km in southern Lake Michigan.

Of common loons tallied during aerial surveys of Lake Michigan, about 37% occurred as singles.

This percentage of single loons may be biased high and flock sizes biased low, as about 34% of

loons were likely underwater during a near instantaneous sampling during the survey plane

overpass. The proportion of loons occurring as singles appeared to be higher than that

documented during late October on Mille Lacs Lake, Minnesota where about 8.4% of loons

observed occurred as singles (McIntyre and Barr, 1983). Flock sizes tended to be larger on Mille

Lacs Lake, given aggregations of >50 individuals, although most groups contained <20

individuals.

Common loon foraging patterns

While staging on Lake Michigan, typical common loon foraging patterns varied by time of day

(Figure 7) and seasonally (Figure 8). McIntyre and Barr (1983) characterize loons as visual,

diurnal predators and associated foraging activity to light levels. During midday, loons were

underwater about 34% of the time in this study. Considering the post-dive surface interval as

part of foraging activity, roughly 68% of the midday period was spent engaged in foraging

activity. In comparison, McIntyre and Barr (1983) reported that total daily foraging time among

common loons on Mille Lacs Lake occupied 46.3% and afternoon (1300-1700 hrs) foraging

17
activity was 78.5% of behaviors. Daily foraging activity of common loons wintering off

Assateague Island, Virginia, was reported as 55.3% of activities (McIntyre, 1978); and foraging

accounted for 23% to 38% of recorded activity among three years of observations of loons

wintering in coastal waters of Weekapaug, Rhode Island (Daub, 1989; Ford and Gieg, 1995).

Foraging among common loons on breeding areas comprised 53-57% of their time during pre-

nesting (but was reduced during the nesting [34-36%] and post-nesting [15-19%] periods) in the

Upper Peninsula of Michigan (Evers, 1994) and 56% of the time among loons with failed nest

attempts and non-breeding loons in Nova Scotia and New Brunswick, Canada (Nocera and

Taylor, 2000).

Mean dive duration among common loons foraging in Lake Michigan (Table 4) exceeded those

reported from studies on breeding lakes (range of 33 to 43 sec; McIntyre, 1978; Parker, 1988;

Paruk, 1999; Nocera and Burgess, 2002; Gingras and Paszkowski, 2006) or wintering areas

(range of 34 to 40 sec; Robinson, 1923; Dewar, 1924; McIntyre, 1978), which tended to be

shallower systems. Evers et al. (2010) reported that dives longer than 120 sec were uncommon.

However, our findings indicate that the average dive duration of loons foraging at depths of 40 ±

5 m in southern Lake Michigan was 139 sec (95% CLs: 137 and 141 sec). Dive durations were

≥120 sec in about 23% of dives documented in Lake Michigan. The observed relation of longer

dive durations with increasing dive depths among common loons using Lake Michigan is

consistent with that observed in other diving birds (Wilson and Wilson, 1988; Kooyman et al.,

1992; Wilson et al., 2011).

The increasing duration at maximum depth of common loons diving in Lake Michigan with

increased dive depth is consistent with findings in studies of diving seabirds (Wilson and Wilson,

1988; Wilson et al., 1996). We also found that common loon post-dive surface intervals on Lake

18
Michigan were positively associated with both depth of dive and dive duration. No association

was determined between loon post-dive surface interval (dive-pause) and dive duration on

breeding lakes in Nova Scotia and New Brunswick (Nocera and Burgess, 2002), but dive

durations (and likely foraging depths) were considerably less than that observed on Lake

Michigan. Schreer et al. (2001) noted a general increase in post-dive surface interval with

increasing dive duration among 12 species of pinnipeds and seabirds. Mean duration of the post-

dive surface interval of deep diving double-crested cormorants (Phalacrocorax auritus) was

significantly longer than that of shallow diving cormorants (Enstipp et al., 2001); through

experimentation using exposure to hyperoxic and hypoxic gas mixtures, the authors concluded

that surface recovery time was related to time required to refuel O2 stores and behavior was

apparently controlled primarily by blood gas (O2 and CO2) levels.

Dive activity data from geotagged loons indicated that a substantial amount of time was spent

foraging while on Lake Michigan. The dive shape of most dives (82%) met the criteria of

Schreer et al. (2001) of a “square” shape where time at maximum depth ≥ 50% of the dive

duration (also termed ‘U-shaped’ dives; e.g., see Wilson et al. 1996). Dive profile patterns

typically consisted of a series of repeated dives to a consistent depth and consistent duration of

time at or near the maximum depth (e.g., Figure 15). Halsey et al. (2007) suggested that such a

pattern of successive dives is indicative of benthic diving. Wilson and Wilson (1988) also noted

that rapid descent and ascent rates, indicative of steep descent and ascent angles, coupled with

extended time at maximum depth are characteristic of benthic-feeding species. A look at the

relation between recorded dive depths and estimated water depth among high accuracy telemetry

location estimates provide multiple examples of loons foraging at or near the benthic zone

(Figure 16). In five of six cases, dives of Loon ID 55480 were in close proximity to the bottom

19
of southern Lake Michigan as indicated from bathymetric data (the 6th case occurred near

sunrise, when only shallow dives were recorded). In five of seven cases, dives of Loon IDs

55488, 55490, and 107270 were within about 5 m of the estimated bottom depth of Green Bay.

In three of six cases, recorded dives of Loon IDs 55490 and 107272 occurred to depths at or

below the estimated water depth of northern Lake Michigan. The bathymetric pattern of

northern Lake Michigan is highly variable and consists of steep depth gradients, especially

within the area offshore of the Door County Peninsula (Wisconsin), which may have contributed

to the observed discrepancies between dive depth and estimated water depth.

Relation of loon distribution and foraging patterns to avian botulism

The observed dive patterns indicate that common loons forage largely on benthic prey while on

Lake Michigan. Forage trawling data suggest that an abundance of prey fish are present across

the loon’s dive depth range on Lake Michigan, and round gobies have become an important

component of the lake's food web (Madenjian et al., 2015; Bunnell et al., 2017). Round gobies

were the dominant prey (100% frequency of occurrence, ~85% mean abundance among total

number of prey found in gut contents, and sole prey item in 52% of loons examined) identified in

the gastrointestinal tracts of 30 beached common loon carcasses collected in association with

type E avian botulism on Lake Michigan in autumn 2012 (Essian et al., 2016). Dreissenid

mussels, crayfish, and alewife (Alosa pseudoharangus) occurred to a lesser extent in loon gut

contents in the same study. Round gobies were also a common food item found among common

loon carcasses associated with botulism outbreaks on Lake Erie and Lake Huron during 1999-

2003 (Campbell, 2002, Campbell et al., 2002, Hannett et al., 2011). While round gobies prefer

shallow water (0.3-7 m) during the summer spawning season (Kornis et al., 2012), a portion of

the bottom-dwelling goby population tends to migrate offshore in the autumn (C. Madenjian, U.

20
S. Geological Survey, personal communication) and have been recorded overwintering as deep

as 130 m in Lake Ontario (Walsh et al., 2007). The availability of round gobies has increased

dramatically in Lake Michigan since first captured in USGS bottom trawl surveys in 2003, as

indexed by biomass estimates; round gobies made up 40% of the biomass of prey fishes in Lake

Michigan in 2014 (Bunnell et al., 2015). During common loon mortality events in 1963 and

1964, prior to round goby invasion, alewife was the most frequent food item present in the guts

of loon carcasses (Peterson, 1965). Several studies have documented a shift in the diets of Great

Lakes double-crested cormorants from that dominated by pelagic (e.g., alewife, three-spine

stickleback [Gasterosteus aculeatus]) and demersal (e.g., yellow perch [Perca flavescens])

species to the benthic-dwelling round goby with the expansion of round goby populations

(Johnson et al., 2010; Van Guilder and Seefelt, 2013; Hebert et al., 2014; Johnson et al., 2015a;

Johnson et al. 2015b).

Ruetz et al. (2009) described the energy density of invasive round gobies as intermediate among

native benthic prey fish in a Lake Michigan tributary. However, energy density of round gobies

in Lake Erie (Johnson et al., 2005) was lower than the findings of Ruetz et al. (2009), a

phenomenon believed to be related to lower quality diets consisting largely of dreissenid mussels

on Lake Erie. Further, Johnson et al. (2005) reported that round goby energy density was lower

than that among prey fishes in Lake Erie. Applying the length-mass and mass-energy density

relations provided by Johnson et al. (2005), a rough estimate of the energy density of the average

sized goby consumed by Lake Michigan loons (110 mm length; Essian et al., 2016) is 4 kJ/gram

wet mass. Despite a possible lower energy density relative to other benthic prey fish, gobies

reach higher population densities than native prey fish, which could increase the total prey

energy available to common loons as well as other piscivorous predators in localized areas.

21
Assuming that round gobies are the principal forage fish of geotagged common loons, the energy

available in this prey resource is evidently profitable for staging loons in meeting or exceeding

daily energy needs considering the energetic cost of repeated diving to 40+ m, pursuing, and

capturing prey.

Unfortunately, the 20-sec interval at which our geolocator tags were programmed to log pressure

was insufficient to interpret prey ingestion rate based on dive profile undulations (e.g., Simeone

and Wilson 2003). A logging interval of roughly 1-2 sec would be required to identify

undulations associated with prey capture, but would have rapidly filled the memories of the

devices and severely restricted the number of days of dive profile data collection.

The spatial distribution of common loons on Lake Michigan (Figures 3 and 6) coincide with

areas of relatively high dreissenid mussel abundance (Rowe et al., 2015). Nalepa et al. (2010)

determined that in 2008 (note that loon distribution data were collected during 2011-2013)

maximum biomass of quagga mussels (Dreissena bugensis) in southern Lake Michigan occurred

at 31-50 m, with the expectation of population expansion to be most rapid at depths >50 m. As

dreissenid mussels are important in the diet of round gobies in the Great Lakes (see review by

Kornis et al., 2012), one can reasonably assume that gobies exploiting dreissenid mussel beds

provide an abundant supply of forage to loons, thereby driving loon distribution. This food web

link has been suggested as a primary pathway of BoNT/E to piscivorous waterbirds foraging in

shallow waters (Getchell and Bowser, 2006; Yule et al., 2006). The results of our study indicate

that energy transfer from dreissenid mussels to higher tropic levels via gobies may occur in deep-

water habitats as well, along with transfer of BoNT/E (Yule et al., 2006). Brush et al. (2012)

concluded from a study conducted in Lake Ontario, that the importance of dreissenids in goby

diets has been overestimated. We acknowledge that an array of benthic invertebrates may be

22
important in the diet of round gobies, some of which may also serve as a potential pathway for

BoNT/E (Yule et al., 2006).

Chipault et al. (2015) reported a seasonal pattern of common loon mortality, where the relative

number of dead loons tallied on beaches of northern Lake Michigan during 2010-2012 peaked

during October (39% to 78% of loon carcasses tallied within years), followed by November (18-

34%) and September (3-27%). This mortality pattern is consistent with the seasonal pattern of

use observed among radiomarked and geotagged loons using northern Lake Michigan in this

study. High relative densities of common loons were documented in northern Lake Michigan

during aerial surveys conducted during the latter half of September and throughout October

(Figure 3). Arrival dates among marked loons to northern Lake Michigan largely occurred from

late September through mid-November (Figure 4), with departure occurring largely during late

October through mid-November.

Outbreaks of type-E avian botulism have been a common occurrence in northern Lake Michigan

since the early 2000s (La Francois et al., 2011; Chipault et al., 2015), in contrast to Green Bay

and southern Lake Michigan that hosted relatively large numbers of common loons during fall

migration where avian botulism outbreaks have not been documented in recent decades.

Botulism outbreaks were last documented in southern Lake Michigan in 1963 and 1976, and in

Green Bay during 1964-1966 and 1983 (Zuccarino-Crowe, 2009) and occurred prior to

dreissenid mussel and round goby invasions. While common loon use areas within Green Bay

and southern Lake Michigan now appear to have the dreissenid mussel-round goby food web

link, the conditions for production and/or transmission of BoNT/E appear to be lacking in recent

decades. Based on the patterns of common loon use versus mortality on Lake Michigan, we

speculate that there may be spatial and/or seasonal hotspots for BoNT/E production and

23
transmission in northern Lake Michigan. In a companion study, Kenow et al. (2016) used a

carcass drift modeling approach to discern where common loons beached along shorelines in

northern Lake Michigan might have acquired BoNT/E from the prey base. Carcass drift back-

tracing findings, coupled with observed loon distribution and foraging patterns, provided an

opportunity to identify potential offshore source locations and supports speculation about the role

of deepwater depositional areas and dreissenid mussel beds as sites of BoNT/E production.

Research targeted to characterize differences in environmental factors, food web dynamics

(including round goby diets), and BoNT/E availability among areas with varied levels of

apparent risk to BoNT/E exposure (i.e., model-estimated sites of toxin acquisition in northern

Lake Michigan versus loon use sites in Green Bay and southern Lake Michigan) may be a

beneficial approach to further our understanding of avian botulism exposure pathways for use in

the identification and development of management options. It is desirable that this evaluation be

conducted through November (to include the typical peak time of botulism-related mortality;

Chipault et al., 2015) in conjunction with active avian botulism outbreaks to ensure measured

dynamics are temporally relevant.

Lake Michigan served as an important staging area of 96% of the adult common loons followed

over the course of fall migration in this study, spending an average of 27 days before continuing

their migration to wintering areas. Assuming these loons were fairly representative of breeding

adult loons in Minnesota, Wisconsin and the Upper Peninsula of Michigan, the finding implies

that these breeding populations may be at considerable risk of exposure to type E avian botulism

in some years. Loon populations breeding in Manitoba and western Ontario likely also use the

Great Lakes as fall staging areas (Evers et al., 2010) where they too may be susceptible to

botulism-related mortality. Monitoring mortality of common loons during fall staging on Lake

24
Michigan and other Great Lakes, given the importance of this resource in the annual cycle of

mid-continent common loon populations, may allow managers to better assess the implications

of avian botulism on regional loon populations.

Acknowledgements

Funding for this study was provided by Great Lakes Restoration Initiative through an

interagency agreement with the United States Environmental Protection Agency; with additional

support from the Legislative-Citizen Commission on Minnesota Resources’ Environment and

Natural Resources Trust Fund, and U.S. Geological Survey Wildlife Program funds. We thank

the numerous volunteers and field biologists who assisted with field logistics in support of this

study. C. Henderson, L. Naumann, R. Baker, P. Perry, and staff with the Minnesota Department

of Natural Resources lined up local cooperators and provided necessary Minnesota permits.

Local field support was provided by W. Brininger and L. Deede (Tamarac National Wildlife

Refuge), L. Laske and S. Maanum (Big Mantrap Lake Association), M. Hintz (Sportsmen’s Club

of Lake Vermilion, Inc.), R. Mack (Burntside Property Owners’ Association), and J. Zernov. P.

Boma, L. Naumann, J. Wilson, T. Daulton, D. Killian, D. Stockwell, K. Woizeschke, K. Carlyle,

and J. Kohn assisted with capture work. K. Carlyle assisted with transmitter implant surgery.

The USGS National Wildlife Health Center provided diagnostics support of the recovered loon

carcass. L. Robinson assisted with air survey transect design and aviation obstacle maps.

Dispatchers with the Minnesota Interagency Fire Center (MIFC) State Aviation Desk located in

Grand Rapids, Minnesota provided flight following service during survey missions. J. Chipault,

M. Gaikowski, and two anonymous reviewers provided helpful reviews of previous drafts.

Thanks also to J. Sleeman (USGS National Wildlife Health Center) and S. Riley (USGS Great

Lakes Science Center) for coordinating a multi-agency research group that investigated avian

25
botulism. Movement data from the radiomarked loons were served on a web site established to

provide project information to partners and the public

(www.umesc.usgs.gov/terrestrial/migratory_birds/loons/migrations.html). The use of trade,

product, or firm names in the publication is for descriptive purposes only and does not imply

endorsement by the U.S. government.

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35
Table 1. Number of adult common loons radiomarked and geotagged in Minnesota, Wisconsin,
and Michigan during 2009-2012 and subsequent geotag recoveries.

Recaptures-recoveries Number of geolocator

Number of Number of of geolocator-tagged records used in Lake
Year loons loons adults/geotags Michigan analyses
radiomarked geotagged recovered
2009 0 18 14/13 4
2010 10 79 52/46 31
2011 21 37 23/16 10
2012 0 42 25/19 8

Total 31 176 114/94 53

36
Table 2. Characteristics of locations of radiomarked (PTT No.) common loons (Gavia immer) using
Lake Michigan during fall migration, 2010-2011. Locations were based on one location per loon per
day with Location Class ≥ 1. PTT = Platform Transmitter Terminal (satellite transmitter).

PTT Area of Number Dates included in Distance to nearest Water depth (m)
No. Lake of analyses shore (km) mean (SD) min–max
Michigan locations mean (SD) min–max
used in
analyses
55480 Southern 20 18 Oct – 29 Nov 10 18.1 (6.9) 7.1 – 36.3 48.4 (14.4) 23.7 – 70.9
55485 Green Bay 17 28 Jul – 23 Sep 10 8.2 (3.7) 1.7 -14.5 24.8 (6.1) 11.6 – 36.0
Northern 4 25 Sep – 09 Oct 10 6.7 (2.8) 4.1 – 9.6 37.1 (5.1) 30.6 – 42.9
55489 Southern 1 22 Nov 10 20.5 46.3
55490 Green Bay 2 17 – 19 Oct 10 5.5 (1.0) 4.8 – 6.2 23.0 (3.4) 20.6 -25.5
Northern 8 20 – 30 Oct 10 13.2 (4.4) 7.8 – 22.8 97.0 (37.4) 49.2 – 178.8
Southern 12 31 Oct – 26 Nov 10 28.4 (5.6) 21.2 – 39.2 62.1 (25.3) 45.2 – 139.8
55479 Southern 2 21 – 22 Nov 10 22.4 (1.2) 21.6 – 23.2 55.2 (2.7) 53.2 – 57.1
55484 Northern 2 16 – 17 Nov 10 14.5 (1.3) 13.6 – 15.4 92.5 (3.6) 90.0 – 95.0
Southern 3 18 – 24 Nov 10 16.0 (4.5) 12.4 – 21.1 57.2 (6.4) 52.6 – 64.5
55488 Green Bay 30 20 Sep – 16 Nov 10 10.1 (3.2) 2.4 – 14.1 28.2 (6.8) 6.2 – 35.9
Northern 1 17 Nov 10 36.1 250.3
107275 Green Bay 1 31 Oct 11 4.8 24.1
107263 Southern 1 09 Dec 11 12.8 32.5
107267 Green Bay 14 31 Oct – 19 Nov 11 4.7 (1.8) 2.3 – 7.3 22.8 (6.8) 9.9 – 34.0
Southern 2 21 – 22 Nov 11 17.5 (16.7) 5.7 – 29.3 31.2 (16.8) 19.3 – 43.1
107259 Southern 2 07 – 09 Dec 11 17.7 (10.2) 10.5 – 24.9 31.6 (15.4) 20.7 – 42.5
107272 Northern 12 31 Oct – 16 Nov 11 8.5 (3.6) 4.4 – 15.3 52.7 (21.2) 26.6 – 104.8
107266 Green Bay 7 18 – 26 Nov 11 4.0 (1.9) 1.7 – 7.3 27.0 (6.7) 13.6 – 33.5
Southern 2 27 – 29 Nov 11 23.3 (10.7) 15.7 – 30.9 103.9 (65.3) 57.7 – 150.1
107273 Southern 6 19 – 29 Nov 11 5.6 (3.0) 2.9 – 10.6 26.7 (10.1) 17.4 – 46.3
107270 Green Bay 5 24 – 29 Sep 11 8.1 (3.2) 5.7 – 13.0 27.1 (4.4) 20.7 – 32.1
107262 Northern 4 01 – 12 Nov 11 3.8 (0.5) 3.2 – 4.3 19.2 (5.2) 13.6 – 25.8
107261 Northern 2 29 – 30 Nov 11 4.6 (2.0) 3.2 – 6.0 23.3 (9.2) 16.8 – 29.8
107264 Northern 6 23 Sep – 09 Nov 11 8.1 (3.8) 3.3 – 12.6 73.6 (60.7) 20.1 – 157.5
Southern 3 10 – 15 Nov 11 26.7 (14.0) 10.6 – 35.3 59.0 (6.7) 51.3 – 63.9
overall Green Bay 76 7.8 (3.7) 1.7 – 14.5 26.1 (6.6) 6.2 – 36.0
Northern 39 9.5 (6.2) 3.2 – 36.1 65.6 (49.9) 13.6 – 250.3
Southern 54 19.6 (9.4) 2.9 – 39.2 50.8 (23.6) 17.4 – 150.1

37
Table 3. Common loon archival geolocator dive data sample sizes (number of individual loons;
number of dives) by location and sex-radio categories. Dive analyses were based on dive activity of
53 geotagged loons, where some loons occurred at more than one location. PTT = Platform
Transmitter Terminal (satellite transmitter).

Sex-radio category
Location Females Males without PTTs Males with PTTs Total
Green Bay 1; 366 2; 5,603 5; 8,848 8; 14,817
Northern Lake Michigan 8; 32,363 5; 7,552 6; 8,922 19; 48,837
Southern Lake Michigan 12; 28,810 22; 34,239 8; 6,553 42; 69,602
Total 21; 61,539 29; 47,394 19; 24,323 69; 133,256

38
Table 4. Estimates of mean dive depth, dive duration, duration at maximum depth, and post-dive surface interval of geotagged common loons
among areas of Lake Michigan, sex-radio category and interactions, 2009-2012; mean estimates for duration at maximum depth and duration of
post-dive surface interval represent geometric means.

Variable Dive depth Dive duration Duration at maximum depth Duration of post-dive surface
(m; 95% CL) (sec; 95% CL) (sec; 95% CL) interval
(sec; 95% CL)
Green Bay 16.3 (14.0, 18.6) 69.8 (61.9, 77.7) 42.9 (39.4, 46.6) 69.9 (63.9, 76.3)
Northern Lake MI 15.1 (12.8, 17.3) 68.9 (61.3, 76.5) 41.1 (38.0, 44.5) 58.5 (53.8, 63.6)
Southern Lake MI 22.6 (20.3, 24.8) 89.6 (82.0, 97.1) 52.1 (48.2, 56.4) 76.4 (70.2, 83.0)

Female 15.5 (12.1, 19.0) 66.6 (54.9, 78.3) 41.8 (36.9, 47.2) 62.1 (54.6, 70.8)
Male without PTT 20.8 (17.8, 23.9) 82.5 (72.2, 92.7) 48.7 (43.8, 54.3) 67.7 (60.5, 75.9)
Male with PTT 17.6 (13.7, 21.5) 79.2 (66.0, 92.4) 45.2 (39.3, 51.9) 74.2 (64.1, 85.8)

Green Bay × Female 10.3 (6.2, 14.4) 51.8 (37.9, 65.7) 36.7 (31.7, 42.5) 54.9 (46.9, 64.2)
Green Bay × Male without PTT 22.3 (18.9, 25.8) 80.6 (68.9, 92.3) 44.5 (39.3, 50.3) 84.9 (74.5, 96.8)
Green Bay × Male with PTT 16.2 (11.7, 20.7) 76.9 (61.7, 92.1) 48.3 (41.1, 56.6) 73.1 (61.7, 86.7)
Northern Lake MI × Female 12.7 (8.9, 16.6) 58.1 (45.1, 71.0) 37.8 (33.0, 43.3) 60.3 (52.2, 69.5)
Northern Lake MI × Male without PTT 17.4 (14.0, 20.8) 78.9 (67.5, 90.4) 50.6 (44.9, 57.1) 53.2 (46.9, 60.4)
Northern Lake MI × Male with PTT 15.1 (10.7, 19.4) 69.7 (54.9, 84.5) 36.4 (31.1, 42.5) 62.4 (53.0, 73.6)
Southern Lake MI × Female 23.6 (19.7, 27.4) 89.9 (77.0, 102.9) 52.5 (45.9, 60.1) 72.5 (62.9, 83.6)
Southern Lake MI × Male without PTT 22.7 (19.3, 26.1) 87.9 (76.5, 99.2) 51.4 (45.6, 57.9) 68.7 (60.6, 77.9)
Southern Lake MI × Male with PTT 21.5 (17.1, 25.9) 90.9 (76.1, 105.7) 52.5 (45.0, 61.3) 89.3 (75.8, 105.2)

39
Table 5. Differences among dive depth and dive duration means, and ratios of geometric means of duration at maximum depth and of post-dive
surface intervals of geotagged common loons among sex-radio categories among areas of Lake Michigan, 2009-2012. ‘*’ denotes differences at p <
0.05, ‘**’ denotes differences at p < 0.01, ‘***’ denotes differences at p < 0.001; adjusted for multiple comparisons.

Differences among means Ratios of geometric means

Variable Dive depth Dive duration Duration at maximum depth Duration of post-dive surface
(m; 95% CL) (sec; 95% CL) (ratio; 95% CL) interval
(ratio; 95% CL)
Female
Green Bay versus Northern Lake MI -2.4 (-5.7, 0.8) -6.3 (-17.7, 5.1) 0.97 (0.86, 1.10) 0.91 (0.79, 1.06)
Green Bay versus Southern Lake MI -13.3 (-16.6, -10..0)*** -38.2 (-49.7, -26.6)*** 0.70 (0.61, 0.79)*** 0.76 (0.65, 0.88)***
Northern versus Southern Lake MI -10.8 (-11.9, -9.8)*** -31.9 (-35.6, -28.2)*** 0.72 (0.69, 0.75)*** 0.8 (0.8, 0.9)***

Male without PTT

Green Bay versus Northern Lake MI 4.9 (2.7, 7.1)*** 1.7 (-6.0, 9.3) 0.88 (0.81, 0.96)** 1.60 (1.44, 1.76)***
Green Bay versus Southern Lake MI -0.4 (-2.3, 1.6) -7.3 (-14.0, -0.5)* 0.86 (0.80, 0.93)*** 1.24 (1.13, 1.35)***
Northern versus Southern Lake MI -5.3 (-6.3, -4.2)*** -9.0 (-12.6, -5.3)*** 0.99 (0.95, 1.03) 0.77 (0.74, 0.81)***

Male with PTT

Green Bay versus Northern Lake MI 1.2 (-2.3, 4.6) 7.2 (-4.8, 19.2) 1.33 (1.16, 1.52)*** 1.17 (1.00, 1.37)*
Green Bay versus Southern Lake MI -5.3 (-8.7, -1.9)** -14.0 (-26.0, -2.0)* 0.92 (0.80, 1.05) 0.82 (0.70, 0.96)***
Northern versus Southern Lake MI -6.5 (-4.6, -8.3)*** -21.2 (-27.7, -14.8)*** 0.69 (0.64, 0.74)*** 0.70 (0.64, 0.76)***

40
Figure Captions
Figure 1. Distribution of common loons marked with geolocator tags (n=176; all circles and
triangles) and satellite transmitters (n=31; triangles only) in Minnesota, Wisconsin and the Upper
Peninsula of Michigan during summer 2009-2012
Figure 2. Archival geolocator tag affixed to modified leg band and attached to common loon leg
(photo credit: Carrol Henderson, Minnesota DNR).
Figure 3. Distribution and average relative density of common loons (COLO) on Lake Michigan by
month as interpreted from aerial surveys averaged over three years, 2011 – 2013. Kernel density
estimates were averaged by month across years, smoothed, and then clipped to the union of the
surveyed areas

Figure 4. Distribution of dates of common loon departure from breeding lake, and arrival and
departure dates at various areas of Lake Michigan.

Figure 5. Migration patterns of adult common loons (n=89) breeding in Minnesota, Wisconsin, and
the Upper Peninsula of Michigan, with respect to passage through Lake Michigan. Widths of arrows
are proportional to the number of loons represented.
Figure 6. Distribution of location estimates of radiomarked common loons (COLO) on Lake
Michigan during fall staging (where LC ≥ 1).
Figure 7. Proportion of time underwater by location and time of day illustrated using dive data from
53 geotagged common loons (refer to Table 3 for sample sizes) while using Lake Michigan.
Penalized B-spline curves with 95% confidence bands are depicted for radiomarked (platform
terminal transmitter; PTT) male (red), non-radiomarked male (green), and female (blue) loons.

Figure 8. Proportion of time underwater by location and Julian date illustrated using dive data from
53 geotagged common loons (refer to Table 3 for sample sizes) while using Lake Michigan during
mid-day hours (0900-1500 hrs). Penalized B-spline curves with 95% confidence bands are depicted
for radiomarked (platform terminal transmitter; PTT) male (red), non-radiomarked male (green), and
female (blue) loons.

Figure 9. Distribution of dive depths by area of Lake Michigan determined from dive data from 53
geotagged common loons (refer to Table 3 for sample sizes). Scale of y-axes vary among panels.
Figure 10. Relation between common loon dive duration (sec) and depth of dive (m) while using
areas of Lake Michigan (refer to Table 3 for sample sizes). Penalized B-spline curves with 95%
confidence bands are depicted for radiomarked male (red), non-radiomarked male (green), and
female (blue) loons. Confidence bands do not adjust for correlation of measurements within birds.

Figure 11. Relation between common loon duration of dive within 2 m of maximum depth (sec) and
depth of dive (m) while using areas of Lake Michigan (refer to Table 3 for sample sizes). Penalized
B-spline curves with 95% confidence bands are depicted for radiomarked male (red), non-
radiomarked male (green), and female (blue) loons. Confidence bands do not adjust for correlation
of measurements within birds.
41
Figure 12. Relation between common loon post-dive surface interval (sec) and depth of dive (m)
while using areas of Lake Michigan (refer to Table 3 for sample sizes). Penalized B-spline curves
with 95% confidence bands are depicted for radiomarked male (red), non-radiomarked male (green),
and female (blue) loons. Confidence bands do not adjust for correlation of measurements within
birds.

Figure 13. Estimated depth of water at radiomarked common loon location estimates while using
Lake Michigan.
Figure 14. Estimated distance from shore of radiomarked common loon location estimates while
using Lake Michigan.
Figure 15. Dive profile of a geotagged common loon staging on Lake Michigan, 08 November
2010.

Figure 16. Estimated dive depths of radiomarked common loons (‘+’) related to water depth (green
bar) where location estimates were of high accuracy (i.e., Argos Location Class 2 or 3; radius of
error corresponds to one standard deviation of the estimated location error of 250 m or 500 m,
respectively). Dives occurred within 30 min of location estimate, except where designated with ‘*’
(within 60 min) or with ‘**’ (within 90 min).

42
Figure 1 color
Figure 1 bw
Figure 2 color
Figure 2 B&W
Figure 3
Figure 4
Figure 5 color
Figure 5 B&W
Figure 6 color
Figure 6 B&W
Figure 7
Figure 8
Figure 9
Figure 10
Figure 11
Figure 12
Figure 13
Figure 14
Figure 15
Figure 16