VOLUME 72, NUMBER 2 127
Journal of the Lepidopterists’ Society
72(2), 2018, 127–144
CITIZEN SCIENTIST TAGGING REVEALS DESTINATIONS OF MIGRATING MONARCH BUTTERFLIES, DANAUS PLEXIPPUS (L.) FROM THE PACIFIC NORTHWEST
DAVID G. JAMES*1, TANYA S. JAMES1, LORRAINE SEYMOUR1, LINDA KAPPEN2
TAMARA RUSSELL3, BILL HARRYMAN4, AND CATHY BLY5
1Department of Entomology, Washington State University, Irrigated Agriculture Research and Extension Center, 24105 North Bunn Road, Prosser, Washington 99350
2 4427 Humbug Creek Road, Applegate Oregon 97530
3 Alaska Psychiatric Institute, 3700 Piper Street, Anchorage, Alaska 99508
4Idaho Fish and Game, Bayview Field Office, PO Box 806, Bayview, Idaho 83803
5 Washington State Penitentiary, 1313 North 13th Avenue, Walla Walla, Washington 99362
*Corresponding author e-mail: david_james@wsu.edu
ABSTRACT. The fall migration of Monarch butterflies (Danaus plexippus) in the Pacific Northwest (PNW) was studied in an un funded citizen science project during 2012–16 by tagging 13778 reared and 875 wild Monarchs. More than a third of these Monarchs were reared by inmates of the Washington State Penitentiary (WSP) in Walla Walla, Washington. Sixty (0.41 %) tagged Monarchs were recovered from distances greater than 10 km (mean: 792.9 ± 48.0 km) with most found in California, SSW of release points. One WSP-reared Monarch was found 724 km to the SE in Utah. Monarchs tagged in Oregon flew SSE to California. No Idaho-tagged Monarchs were found in California but two were recovered at locations due south. No wild tagged Monarchs from Washington, Ore gon or Idaho were recovered. Monarchs from Washington and Oregon were found during October-February at 24 coastal California overwintering sites spanning 515 km from Bolinas to Carpinteria. A single wild spring Monarch tagged in May in northern California was recovered 35 days later and 707 km ENE in Twin Falls, Idaho. This study provides compelling evidence that many Monarchs in southern and central parts of Washington and Oregon migrate south in the fall to overwintering sites along the California coast. It also provides some evidence for southerly and south-easterly vectoring of migrating Monarchs from eastern Washington and Idaho, indi cating the possibility of migration to Arizona or Mexico overwintering sites. In addition to improving our understanding of Monarch migration in the PNW, this study also contributed to conservation by adding nearly 14000 butterflies to the population. The incredi ble involvement of incarcerated and non-incarcerated citizen scientists generated much community and media interest which in turn led to greater involvement by citizens. Increased awareness of Monarchs, their biology and conservation in the PNW has been an un expected but important spin-off of this study.
Additional key words: overwintering sites, migration directions, conservation, incarcerated citizen scientists.
It has long been assumed that all Monarch butterflies, Danaus plexippus (L.), seen in the summer in the Pacific Northwest (PNW) (British Columbia, Washington, Idaho, Oregon) originate from California and return in the autumn to that state for overwintering (e.g. Urquhart 1960, Urquhart and Urquhart 1977, Brower 1995, Dingle et al. 2005, Stevens and Frey 2010). Pyle (2015) in a review on Monarchs in the PNW noted that this was an assumption based on good sense rather than an abundance of good scientific evidence. However, some limited data do exist showing movement of wild Monarchs from the PNW to California. Essentially, these comprise about a dozen records of tagged Monarchs flying from Boise, Idaho to the California coast in the 1970’s, and a single individual from Roosevelt, Washington found on the California coast near Santa Cruz in 1997 (Pyle 2015). Tagged Monarchs used in Urquhart and Urquhart (1977) and by a Monarch enthusiast (Paul Cherubini, pers. comm.)
provided other records of migration to California but are not considered to be an accurate and reliable representation of what PNW Monarchs do because they involved Monarchs obtained in California and Ontario, shipped to the PNW for release. See Pyle (2015) for a detailed explanation why transferred Monarchs cannot be considered as surrogates for naturally occurring Monarchs in the location of release. Therefore, the data emanating from the transfer experiments of Urquhart and Urquhart (1977) were instrumental in building the potentially flawed canon that all PNW Monarchs migrate to and from California. Moreover, there are some data and observations suggesting that autumn PNW Monarchs may also migrate in directions other than those which would take them to California. Although some of the wild Monarchs that were tagged in Boise in the 1970’s (by school teachers Faye Sutherland and Mary Henshall) flew to California, others were recovered from distances of up to 750 km in
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Washington State Penitentiary (WSP), Walla Walla, WA
reared and tagged Monarchs in each year of the five
year program. Eggs and first instar larvae produced by
female D. plexippus obtained from breeding
populations in far northern California (Trinity River
National Forest) or central Washington, were provided
to WSP inmates who then reared the larvae to
adulthood (James 2016). Adults were tagged by inmates
and released in the Penitentiary grounds or taken to
nearby locations for release by Penitentiary staff. In
2012, white tags showing an email address and serial
number were manufactured at WSP but in subsequent
years customized white tags were obtained from
MonarchWatch.org. Rearing of Monarchs was adopted
by WSP as a way of improving the mental health of long
term inmates by giving them a purpose and focus to
FIG. 1. Monarch butterfly with tag used in this study.
a south easterly direction in Utah (Pyle 1999, 2015). Pyle (1999) in his autumn journey from British Columbia to Mexico encountered migrating Monarchs in the inland PNW and the Great Basin and noted their vanishing bearings. Fifty three of 62 Monarchs in this region gave vanishing bearings toward the south and south east (Brower and Pyle 2004). These bearings would lead migrating western Monarchs to Mexico, a hypothesis supported by Pyle’s observations in New Mexico of Monarchs flying south to Mexico just 16 km away (Pyle 1999, 2015). Indirect evidence of the origin of Monarchs in California overwintering colonies based on spatial analyses of stable isotope ratios was recently provided by Yang et al. (2015). While these data supported the hypothesis that some PNW Monarchs migrate to California they did not exclude the possibility that others may overwinter elsewhere. Morris et al. (2016) presented data showing Monarchs tagged in Arizona migrate to both California and Mexico overwintering sites, the first hard evidence that western Monarchs may overwinter in Mexico.
Given the history of efforts to understand the seasonal movements of Monarchs in the PNW and the equivocal nature of conclusions drawn from limited or flawed experiments, we commenced a program of tagging in 2012. Conducted primarily on PNW-sourced Monarchs both reared and wild, we present the results here from our first five years.
MATERIALS AND METHODS
Annual rearing of Danaus plexippus for tagging was conducted in multiple locations throughout the Pacific Northwest (British Columbia, Washington, Idaho, Oregon and Nevada) during 2012–2016. Inmates at the
their lives (James 2016). Rearing was also conducted at Prosser and Yakima, WA (DGJ, TSJ, LS) and in Bayview, ID (BH, Bill Ament). During 2014–16, citizen scientists in south west Oregon, north and central Oregon, British Columbia and Idaho reared and tagged Monarchs. The vast majority of Monarchs were reared from locally-obtained livestock in WA, OR and ID. However, in the first two years of the project ~ 500 Monarchs released in WA and ID were reared from stock originating from a mid-California butterfly farm (Table 1). All rearers were aware of the importance of cleanliness in preventing disease. Gravid females from which livestock were obtained in WA and ID were checked during 2014–16 for the presence of protozoan parasite Ophryocystis elektroscirrha (OE) spores (Altizer and de Roode 2015) and not used for egg-laying if they tested positive. The incidence of OE-affected wild adults was low (< 5 %). Mass-rearing was conducted only once in each location each season to prevent disease and OE build-up. Tags were placed on the mitten-shaped or discal cell on the ventral surface of a hindwing (Fig. 1). All Monarchs were tagged with a single tag except for 264 (2014) and 631 (2015) reared/released at Yakima, WA which were ‘double tagged’, ie they had a single tag on each hindwing. None of these double-tagged Monarchs was recovered. From 2013–2016, wild Monarchs in Washington, Idaho, Oregon and northern California were also opportunistically tagged during May–September. For all tagged Monarchs, the date of tagging/release, name of tagger, tag number, location of release and the sex of the butterfly were recorded.
Tagged Monarchs were recovered by citizens sighting and/or photographing the butterfly and emailing the address on the tag. Two of the authors (DGJ & TSJ) conducted eight visits of four to eight days each to a total of 37 north and central California overwintering
VOLUME 72, NUMBER 2 129
TABLE 1. Coastal California Monarch overwintering sites visited and searched by DGJ and TSJ for tagged Monarchs during 2013–16. Period of Visit Monarch Overwintering Sites Visited
Santa Cruz (Natural Bridges, Lighthouse Field, Moran Lake), Pacific Grove, Ardenwood, Bolinas (Kale
November 27–30 2013 (4 days) November 26–29 2014 (4 days)
December 28–January 3 2014/15 (7 days)
November 22–28 2015 (7 days)
December 27–January 1 2015/16 (6 days)
February 2–5 2016 (4 days)
November 19–25 2016 (7 days)
December 27–January 1, 2017 (6 days)
& Juniper, Agate Ck.)
Santa Cruz (Natural Bridges, Lighthouse Field, Moran Lake), Pacific Grove, Andrew Molera SP, Monterey,
Goleta (Ellwood, Maria Ygnacio Ck. San Jose Ck,), Ventura (Ocean Ave. Park), Gaviota Beach, Pismo (Halcyon Hill, Oceano CG., State Beach), Andrew Molera SP., Pacific Grove, Monterey, Santa Cruz (Natural Bridges, Lighthouse Field, Moran Lake)
Goleta (Ellwood, Maria Ygnacio Ck. San Jose Ck,), Carpinteria Creek, Ventura (Ocean Ave. Park, Camino Real Park, Arrundel Barranca), Gaviota Beach, Nipomo-Trilogy, Pismo (Halcyon Hill, Oceano CG., State Beach), Morro Bay CG., Los Osos, Andrew Molera SP., Pacific Grove, Monterey, Santa Cruz (Natural Bridges, Lighthouse Field, Moran Lake), Ardenwood Farm, Bolinas (Kale & Juniper, Agate Ck.)
Gaviota Beach, Nipomo-Trilogy, Pismo (Halcyon Hill, Oceano CG., State Beach), Morro Bay CG., Andrew Molera SP., Pacific Grove, Monterey, Santa Cruz (Natural Bridges, Lighthouse Field, Moran Lake), New Park Mall (Newark), Ardenwood Farm, Stinson Beach, Bolinas (Kale & Juniper, Agate Ck.)
Santa Cruz (Escalona, Potbelly, Sunscape), Bolinas (Kale & Juniper, Agate Ck.), Albany Hill, Berkeley Aquatic Park, San Leandro G.C., New Park Mall (Newark)
Goleta (Ellwood, Tecolote), Carpinteria (Creek, Dump), Ventura (Ocean Ave, Park), Santa Barbara (Padaro Lane 2 sites), Gaviota Beach, Nipomo-Trilogy, Pismo (Halcyon Hill, Oceano CG., State Beach), Morro Bay (CG., Del Mar Park, Toro Creek), Cayucos (Villa Creek), Andrew Molera SP., Pacific Grove, Santa Cruz (Natural Bridges, Lighthouse Field, Moran Lake, Escalona, Aptos GC, Branciforte Avenue), New Park Mall (Newark), San Leandro G.C., Albany Hill, Berkeley Aquatic Park, Stinson Beach, Bolinas (Kale & Juniper, Agate Ck.)
Pismo (Halcyon Hill, Oceano CG., State Beach), Morro Bay (CG., Del Mar Park, Toro Creek), Cayucos (Villa Creek), Esalen (Big Sur), Cambria (Hamlet), Santa Cruz (Lighthouse Field, Moran Lake, Escalona, Branciforte Avenue), New Park Mall (Newark), San Leandro G.C., Albany Park, Muir Beach, Stinson Beach, Bolinas (Purple Gate, Kale & Juniper, Agate Ck.)
RESULTS
colonies during 2013–16 (Table 1). In 2016, substantial searching of overwintering colonies during October to January 2017 for tagged Monarchs was also conducted by Joe Billings, an Arizona-based naturalist with MonarchQuestAZ. John Dayton, a Monarch biologist located in Santa Cruz frequently visited local overwintering sites (primarily Lighthouse Field, Natural Bridges, Moran Lake) during October–February in each year and reported tags. Most citizen and author recoveries of tagged Monarchs were supported by a confirmatory photograph. Recoveries of Monarchs within 10 kilometers of the release point were considered ambiguous and are not reported here. Date, location and tag number were recorded for all recovered Monarchs and straight-line distances from release to recovery locations, calculated.
A total of 13778 Monarchs were reared and tagged in the PNW during summer and fall 2012–16 with the number tagged each year ranging between 1756 and 3863 (Table 2). In addition, 875 wild Monarchs were tagged during 2013–16 (Table 3). A total of 60 tagged Monarchs were recovered during this study representing 0.41% of the butterflies tagged (Tables 4–6, Figs. 2–4). All but three recoveries were of butterflies reared from locally-sourced livestock (Table 4). All recoveries were of reared butterflies; no wild Monarchs tagged in Washington, Idaho or Oregon were recovered. A substantial increase in recoveries occurred in later years of the project with 80% of the recoveries occurring in 2015 and 2016 (Fig. 5). The majority (36.7%) of Pacific Northwest fall migrants were found in California, SSW of release points (Figs. 2, 4 & 6).
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TABLE 2. Numbers of Monarchs reared and tagged by organizations and citizen scientists in the Pacific Northwest during 2012-16 (WSP = Washington State Penitentiary, WSU = Washington State University, CCC = Cowiche Canyon Conservancy, CWI = College of Western Idaho, IDFG = Idaho Fish and Game, IMNH = Idaho Museum of Natural History, USFWS = United States Fish and Wildlife Service). Origin (state) of reared Monarchs shown in parentheses after totals (CAF = California farm-sourced). Names of citizen taggers are shown in acknowledgments.
2012-
Location 2012 2013 2014 2015 2016 2016
Washington
WSP (Walla Walla, Pasco)
WSU (Yakima, Prosser) CCC (Yakima)
Citizens (Redmond, Vancouver)
1310 (472 N CA, 834 WA)
Jul 7-Oct 4
620 (408 CAF, 158 WA, 54 N CA) Jul 7-Oct 15
1307 (482 N CA, 815 WA)
Jul 1-Sep 23
449 (333 WA, 87 CAF, 29 N CA) Jul 31-Oct 25
880 (419 N CA, 461 WA)
Jul 1-Oct 13
1035 (884 WA, 151 N CA)
Aug 22-Oct 19
1390 (738 N CA, 652 WA)
Jul 6-Oct 20
1097 (656 WA, 441 N CA)
Jul 1-Oct 21
60 (WA)
Sep 11-29
136 (CAF)
Aug 27-Oct 16
242 (128 N CA,
114 WA)
Jul 4-Sep 2 5129
1254 (WA)
Aug 17-Oct 20 4455
152 (WA)
Aug 26-Sep 14 212
40 (WA)
Sep 18-22 176
Total (WA) 1930 1756 1915 2683 1688 9972 Idaho
IDFG (Bayview)
451 (CAF) Aug 9-Sep 13
241 (ID)
Aug 11-Sep 25
252 (ID)
Aug 27-Oct 14
116 (CAF)
Aug 6-18 1060
CWI (Nampa) 30 (ID) Oct 6-26 30 27 (ID)
IMNH
Citizens (Boise, Blackfoot, Nampa)
5 (ID)
Jul 4-Sep 12
Jul 7-Sep 12 27
75 (ID)
Jul 5-Oct 26 80
Total (ID) 451 0 241 257 248 1197
S. Oregon & N. CA
Citizens (Applegate,
Eugene, Medford, Port
Orford, Brookings, Talent,
Elkton, Jacksonville,
Williams, Gold Hill, Wonder, Redding (CA)
USFWS
60 (OR)
Jun 10-Oct 6
849 (OR) Jul 1-Nov 1
1211 (OR)
Jul 1-Oct 26 2120 221 (OR)
Jul 1-Oct 18 221
Total (S. OR & N. CA) 0 0 60 849 1432 2341
N & C Oregon
Citizens (Corvallis,
Portland, Bend, Newberg,
Crow) 44 (OR) Jul 1-27 British Columbia
120 (OR)
Jul 13-Aug 30 164
Citizen (Naramata) 53 (BC) 53 Nevada
Citizens (Reno, Verdi, Sparks)
30 (NV)
Jul 10-Aug 18
21 (NV)
Aug 23-Oct 6 51
Total (PNW) 2381 1756 2269 3863 3509 13778
VOLUME 72, NUMBER 2 131
FIG. 2. Directions traveled and destinations of Monarchs reared and tagged in Washington during July-September 2012- 16.
FIG. 3. Directions traveled and destinations of Monarchs reared and tagged in Oregon during August-October 2014-16.
FIG. 4. Directions traveled and destinations of Monarchs reared and tagged in Idaho during August-October 2012-15.
One tagged Monarch was recovered in Brigham City, Utah, 724 km south east of its release point in Walla Walla, WA (Fig. 2). One male traveled 20 km east from Prosser to Benton City in WA while a female from Brookings, OR was found 84 km to the ENE in Williams, OR. All but one of the remaining 13 (23.3%) Monarchs that had an easterly component to their flight (SSE) originated in central or southern OR and their slight easterly orientation was likely influenced by the NW–SE oriented CA coastline (Fig. 3). The exception was released in Yakima, WA but had only traveled 32 km SSE before recovery. The majority of migrants from
Washington (63.3%) were recovered in locations SSW of their release points (Figs 2 & 6). In contrast, the majority of migrants from Oregon (46.1%) were recovered SSE of release points (Figs. 3 & 6). Two of three recovered Idaho Monarchs were found due south of release points at 93 and 124 km (Figs. 4 & 6). Forty nine recoveries of Washington and Oregon-released Monarchs were made at 24 coastal California overwintering sites during October–February, 22 of these occurred in 4 Santa Cruz overwintering colonies (Table 7, Figs. 2–3). Two individuals were recovered at more than one overwintering site (Table 6, see below).
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In Washington the recovery rate was higher for
Yakima-released Monarchs (0.40 %) than Monarchs
released 180 km further east at Walla Walla (0.12 %). A
single wild Monarch tagged as it was migrating in late
spring along the Trinity River in far northern California,
was recovered 35 days later and 707 km ENE in Twin
Falls, Idaho (20.2 km/day) (Table 9, Fig. 7).
Residency and inter-colony movement of
tagged monarchs at overwintering sites.
Seventeen tagged Monarchs sighted at overwintering
colonies were re-sighted two to six times in the same
colonies 3 to 123 days (Mean 28.5 ± 6.7) later (Table
10). Two tagged Monarchs were re-sighted in different
overwintering colonies. One of these (A4853 released
FIG. 5. Number of PNW-tagged Monarchs recovered annu ally during 2012-16. Numbers above and below each bar rep resent percentage recovery rate and the number of Monarchs reared and tagged in each year, respectively.
Twelve tagged Monarchs were found at the Lighthouse Field overwintering site in Santa Cruz during the study. Remarkably, seven of these migrated from the same location in Yakima, WA (2 in 2014, 5 in 2016). The second most popular location for recoveries (7) was Bolinas and 5 were recovered at three Morro Bay overwintering sites (Table 8). In 2016, 10 of 15 recoveries of Yakima, WA Monarchs were released within a 48 hour period (20–22 August). These butterflies were found at overwintering sites in Santa Cruz (4), Bolinas (2), San Leandro (1), Goleta (1), Morro Bay (1) and en route at Anapolis, CA (1), north of San Francisco. Thirteen tagged Monarchs were recovered during the fall migration before arrival at overwintering sites, showing a mean (± SE) travel rate of 35.1 ± 4.3 km/day (Table 8). No Idaho Monarchs were recovered at California overwintering sites but three were reported during migration at distances of 93-209 km south or south-west of release points (Fig. 4). The greatest straight line release point to recovery point distance was 1360 km from Yakima, WA to Tecolote Canyon near Goleta, CA (Table 6). The mean distance traveled by the 60 recovered butterflies was 792.9 ± 48.0 km. Washington-released Monarchs traveled further (994.9 ± 67.4 km, n = 29) than central Oregon (945.0 ± 180.5 km, n = 2), southern Oregon (823.2 ± 191.8 km, n = 26) or Idaho (142 ± 34.7 km, n = 3) Monarchs. Of the 60 tagged Monarchs that were recovered, 29 (48.4%) originated in Washington, 26 (43.3%) in southern Oregon, 3 (5.0%) in Idaho and 2 (3.8%) in north-central Oregon. The percentage recovery rate was higher in southern (1.1%) and northern-central (1.2%) Oregon Monarchs than Washington (0.29%) or Idaho (0.25%) Monarchs.
in Corvallis, OR, August 30 2016) was seen at three Santa Cruz overwintering sites during October 11–December 30 following its original sighting in North Beach, San Francisco (Table 6, Fig. 8). A6504 was seen at two Carpinteria overwintering sites (Table 6).
DISCUSSION
This study provides definitive evidence that many Monarchs in Washington and Oregon migrate south in late summer and fall to a range of overwintering sites along the California coast from Marin County, north of San Francisco to Carpinteria in Santa Barbara County just north of Ventura. The rate of recovery of tagged Monarchs increased substantially during the project. The number of butterflies we tagged also increased annually but not to the same extent as recoveries. It is possible or even likely that the recent increased awareness of Monarchs among the general public and the ubiquity of cell phone cameras have conspired to make the reporting of tagged Monarchs easier and more frequent.
Forty nine reared and tagged Monarchs from Washington and Oregon were recovered at California overwintering sites while 13 were recovered during migration, most heading south to south west towards California. One Washington-released Monarch was recovered at an I-84 rest stop just north of Brigham City, Utah, 724 km south east of its release point at Walla Walla. If this individual arrived at this location under its own power, then it clearly was not heading towards the California coast and may have been orienting towards Arizona and Mexico. Interestingly, no Monarchs reared and tagged in Idaho were recovered in California. Only three Idaho Monarchs were recovered and two of these were heading south following routes that if continued would lead them towards Mexico rather than California. These records of south-easterly and southerly-vectoring Monarchs
VOLUME 72, NUMBER 2 133
FIG. 7. Spring remigration. Direction traveled and destination (Twin Falls, ID) of a wild Monarch tagged on May 24 2015 in the Trinity National Forest, CA.
from eastern Washington and Idaho support the 1970s data of Idaho schoolteachers Faye Sutherland and Mary Henshall as well as the suggestions of Dingle et al. (2005) and observations of Pyle (1999, 2015), that postulate at least some Monarchs in the eastern portion of the PNW do not migrate to California.
FIG. 6. Proportional representation of directions traveled by tagged Monarchs released in the Pacific Northwest during 2012-16 and recovered at distances > 10 km from release points.
In our study we reared and tagged 1197 Monarchs in Idaho and the recovery rate of 0.25 % was comparable to that from the 9972 Monarchs we reared and tagged in Washington (0.29 %). Morris et al. (2015) also obtained a 0.2 % recovery rate (in Mexico and California) from 12,088 wild Monarchs tagged in Arizona during an 11 year period. However, Joe Billings (MonarchQuest AZ) recorded a 1.44 % recovery rate (in Mexico and California) for wild-tagged Monarchs in Arizona during 2014-16 (J. Billings, pers. Comm.). The recovery rate for Oregon-reared Monarchs was 1.1–1.2 %, similar to the recovery rate for tagged eastern US Monarchs (1.2%) during 1998–2015 (http:// monarchwatch.org/blog/2016/11/02/tagging-results-and the-monarch-decline/). In Virginia, a difference in recovery rates in Mexico for Monarchs tagged in a coastal (1.29 %) and a more inland location (0.16 %), <200 km apart, was reported by Brindza et al. (2008). At two locations in Washington (Yakima, Walla Walla) where substantial numbers of tagged Monarchs were released, the recovery rate for the more eastern site (Walla Walla: 180 km east of Yakima) was lower (0.12 %) than for Yaklma (0.40 %). Lower recovery rates for eastern Washington and Idaho Monarchs possibly indicate that a proportion of the Monarch population in these areas migrate through landscapes less populated by people with a reduced chance of being sighted and reported. Migration south or south-easterly would take Monarchs for more than 2000 km through sparsely settled areas of Idaho, Utah and Arizona. Clearly, far
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TABLE 3. Numbers of wild Monarchs tagged by citizen scientists and agency personnel (WSU, IDFG, CWI, USFWS) in the Pacific Northwest during 2013–16
Location 2013 2014 2015 2016 2013–16 Washington
Yakima 1 0 0 2 3 Vantage 24 163 79 70 336 Roosevelt 0 2 0 0 2 Moses Lake 0 0 5 0 5 Prosser 0 0 1 0 1
Total (WA) 25 165 85 72 347 Idaho
Pocatello 0 0 5 1 6 Boise 0 0 13 51 64 Nampa 0 0 1 28 29 Glenn’s Ferry 0 0 0 1 1 Horseshoe Bend 0 0 0 6 6 Payette 0 0 0 3 3 Aberdeen 0 0 0 10 10 Salmon 0 0 0 19 19 Montpelier 0 0 0 6 6 Jerome Co. 0 0 0 5 5 Twin Falls Co. 0 0 0 21 21 Wendell 0 0 0 1 1 Jefferson Co. 0 0 0 11 11 Bingham Co. 0 0 0 18 18 Oneida Co. 0 0 0 8 8 Owyhee Co. 0 0 0 27 27 Gooding Co. 0 0 0 2 2 Total (ID) 0 0 19 218 237
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TABLE 3. Continued.
Location 2013 2014 2015 2016 2013-16 Southern Oregon
Southern Oregon
Applegate 0 19 43 43 105 Talent 0 0 16 1 17 Medford 0 0 17 8 25 Malheur 0 0 2 0 2 Elkton 0 0 0 2 2 Klamath Falls 0 0 0 1 1
Total (S. OR) 0 19 78 55 152 Northern & Central Oregon
Umatilla 0 0 12 0 12 Corvallis 0 0 0 12 12 Portland 0 0 0 4 4 Bend 0 0 0 5 5 Clarno 0 0 0 6 6 Total N & C Oregon 0 0 12 27 39 Nevada
Reno 0 0 4 42 46 N. California
Eagle Ck, Trinity NF 0 0 21 32 53 Total (PNW) 25 184 219 447 875
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TABLE 4. Release, recovery and distance data for Monarchs reared, tagged and released in the PNW during 2012-14. Distance
Tag # Sex
Release
Location Walla Walla,
Release Date & Tagger
Recovery Location
Recovery Date
(kilometers &
direction) Finder
1837* Female
WA July 13 2012 WSP Brigham City, UT August 3 2012 724 SE Michael Sell August 15 2012
260** Female Post Falls, ID
Bill Harryman Potlatch, ID August 18 2012 93 S Judi Rohn
3172** Female Yakima, WA Walla Walla,
September 17 2012 David James
September 18
Agate Ck., Bolinas, CA (OS)
Milton-Freewater,
October 30
2012 998 SW Rose Poulsen October 26
1076 Female
WA
2013 WSP
September 4 2014
OR
2013 11 S Gayle Blomme September 7
6949 Male Prosser, WA 7024 Male Yakima, WA
Lorraine Seymour Benton City, WA
September 4 2014
David James Glen Ellen, CA September 25
2014 20 E Stephen Zetz
September 27
2014 1030 SSW David Hamilton
5741 Female Stateline, ID
2104 Bill Ament Walla Walla, WA October 3 2014 209 SW Esther Schluter September 30
0683 Female Applegate, OR
2014 Linda
Kappen San Mateo, CA Lighthouse Field,
October 10
2014 531 S Albert Fong
3524 Male Yakima, WA
August 25 2014 David James
Santa Cruz, CA (OS)
November 21
2014 1085 SSW John Dayton
5925 Female
Walla Walla, WA
September 5 2014 WSP
August 29 2014
Maria Ygnacio Ck., Goleta, CA (OS)
Lighthouse Field, Santa Cruz, CA
November 22
2014 1336 S November 28
Charis van der Heide
6416 Male Yakima, WA
David James
September 7 2014
(OS)
Pacific Grove, CA
2014 1085 SSW Jasmine James
6996 Male Prosser, WA
Lorraine Seymour
(OS) January 3 2015 1094 SSW David James
*Source: reared from wild-collected N California female (May 2012)
**Source: reared from CA commercial butterfly farm stock. All other recoveries derived from local (to release point) brood stock. OS = overwintering site. WSP = Washington State Penitentiary
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TABLE 5. Release, recovery and distance data for Monarchs reared, tagged and released in the PNW during 2015. OS = over wintering site
Tag# Sex
Release Location
Release Date & Tagger
August 28 2015 Bill
Recovery Location
Recovery Date (additional
sighting)
Distance
(kilometers
& direction) Finder
A3328 Male Rathdrum, ID
Harryman Troy, ID September 9 2015 124 S Gerry Queener
A3001 Female Brookings, OR
August 31 2015
Vicki Mion Williams, OR September 17 2015
September 10
2015 84 ENE
Michelle Scherer
A3223 Male Brookings, OR
Vicki Mion Aptos, CA October 16 2015 612 SSE Deb Ospina
A0578 Male Walla Walla, WA
September 8 2015 WSP
October 5 2015
Morro Bay SP
CG CA (OS) October 21 2015 1247 SSW Regena Orr
A3771 Male Applegate, OR
Tyler Kappen Boonville CA October 23 2015 365 SSW Beth Swehla Villa Ck.,
A2099 Male Medford, OR
September 9 2015 Tiffany Wyatt
Cayucos, CA
(OS) October 28 2015 801 S Paul Cherubini Natural
A2067 Female Applegate, OR A2062 Male Applegate, OR
August 19 2015 Linda Kappen
August 19 2015 Linda Kappen
September 17 2015
Bridges SB,
Santa Cruz, CA (OS)
Moran Lake, Santa Cruz, CA (OS)
Pismo Beach
November 16 2015
(November 28
2015) 604 SSW
November 10 2015
(November 26
2015) 605 SSW
November 24 2015
(December 28
Andrea &
Denise, David James
John Dayton, David James
A3264 Male Applegate, OR
Linda Kappen September 7 2015
SP (OS)
Morro Bay SP
2015) 861 SSE David James
A0417 Male Pasco, WA
WSP
CG CA (OS) November 24 2015 1247 SSW David James
A0508 Male Pasco, WA 4789 Female Talent, OR
September 7 2015 WSP
August 19 2015 Aleece Townsend
September 13 2015
Moran Lake, Santa Cruz, CA (OS)
Kale Road,
Bolinas, CA (OS)
Kale Road,
Bolinas, CA
November 26 2015
(December 30
2015) 1078 SSW
November 28 2015
(December 10
2015) 486 S
David James, Tanya James & John Dayton
David James, Paul Cherubini
A3712 Female Brookings, OR A0354 Male Touchet, WA
Vicki Mion
September 6 2015 WSP
September 26 2015
(OS) November 28 2015 483 SSE Tanya James
New Park Mall,
Newark, CA
(OS) December 31 2015 991 SSW David James Kale Road,
A2045 Male Medford, OR
Robert & Simone Coffan
October 4 2015
Bolinas, CA (OS)
Agate Ck., Bolinas CA
January 1 2016
(February 3 2016) 502 S David James
A3063 Male Talent, OR
Aleece Townsend
(OS) January 1 2016 486 S David James
138138 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
TABLE 6. Release, recovery and distance data for Monarchs reared, tagged and released in the PNW during 2016. OS = over wintering site.
Tag # Sex Release Location
Release Date & Tagger
Recovery Location
Recovery date
(additional sighting)
Distance
(kilometers & direction)
Finder
A6093 Male Ashland, OR Aug 28 2016
Steven Johnson Vacaville, CA Sept 5 2016 457 S Kathy Keatley
Garvey
A7079 Female Brookings, OR Aug 30 2016
Patsy Haggerty Windsor, CA Sept 6 2016 438 SSE Mike & Barbara
Schumacher
B2838 Male Yakima, WA Aug 22 2016
David James Anapolis, CA Sept 14 2016 917 SSW Jane Simmonds
San Francisco, CA
A 4853 (seen at 4 locales)
Female Corvallis, OR
Aug 30 2016
Molly Monroe & Amelia Jebousek
(1) Lighthouse Field, Santa Cruz, CA, (OS) (2)
Natural Bridges SB, Santa Cruz CA (OS) (3)
Moran Lake,
Santa Cruz, CA (OS) (4)
Lighthouse Field,
Sept 18 2016 (1) (Oct 11 2016 (2) Nov 25 2016 (3) Dec 30 2016 (4))
Sept 30 2016 (1)
765 S
103 SSE
(from location 1) 2.6 W
(from
location 2) 7.4 E (from location 3)
Lisa de Angelis, John Dayton, Aleece
Townsend,
David James
B2701 Male Yakima, WA Aug 21 2016 David James
B4112 Female Yakima, WA Oct 2 2016
Santa Cruz, CA (OS)
(Jan 6 2017 (2) Jan 9 2017 (3) Feb 1 2017 (4))
1085 SSW John Dayton, David James
David James Wapato, WA Oct 8 2016 32 SSE Heinz Humann
A2705 Female Talent, OR
Aug 17 2016 Aleece
Townsend
Natural Bridges SB, Santa Cruz, CA (OS)
Oct 10 2016 591 SSE Steve Cary
B2174 Female Yakima, WA Sept 8 2016 Cindy Dunbar
Morro Bay SP CG
CA (OS) Nov 1 2016 1275 S Regena Orr
A6935 Female Brookings, OR
Aug 25 2016 Andrea
Christensen Sept 20 2016
Lighthouse Field, Santa Cruz, CA (OS)
Lighthouse Field,
Nov 5 2016 (1) (Nov 28 2016 (2), Nov 30 2016 (3))
Nov 5 2016 (1)
618 SSE John Dayton, Joe Billings
John Dayton,
B2452 Male Redmond, WA
Connie
Granberg
Santa Cruz, CA (OS)
(Nov 25 2016 (2)) 1207 S
Aleece
Townsend
B2729 Male Yakima, WA Aug 21 2016 David James
Natural Bridges SB, Santa Cruz, CA (OS)
Nov 7 2016 1085 SSW John Dayton Nov 13 2016 (1) (Nov
B2742 Male Yakima, WA Aug 21 2016 David James
Lighthouse Field, Santa Cruz, CA (OS)
Carpinteria Ck.,
23 2016 (2), Nov 30 2016 (3), Dec 12 2016 (4), Dec 30 2016 (5), Jan 6 2017 (6))
Nov 11 2016 (1)
1085 SSW John Dayton, David James
A6504
(seen at 2 locales)
Male Sisters, OR Sept 17 2016 Susie Werts
CA (OS) (1) Dump Road, Carpinteria CA (OS) (2)
(Nov 22 2016 (1), Jan 11 2017 (2), Jan 23 2017 (2), Jan 26 2017 (2))
1126 SSE, 0.6 E (from location 1)
Joe Billings, David James
VOLUME 72, NUMBER 2 139 TABLE 6. Continued.
Tag # Sex Release Location
Release Date &
Tagger Recovery Location
Recovery date (additional
sighting)
Distance
(kilometers & direction)
Finder
B3554 Male Elkton, OR
Sept 23 2016
Barbara Slott/Kris Hendricks
Ardenwood Historic Farm, Fremont, CA (OS)
Nov 11 2016 (1)
(Nov 16 2016 (2) 694 SSE
Paul
Cherubini, Christine Garcia
B2603 Male Yakima, WA Aug 20 2016 David James
B3587 Female Talent, OR Sept 27 2016 Aleece Townsend
B3202 Male Yakima, WA Sept 9 2016 David James
Toro Ck., Cayucos CA
(OS) Nov 13 2016 1247 S Joe Billings Del Mar Park, Morro
Bay (OS) Nov 19 2016 798 SSE Joe Billings
Lighthouse Field,
Santa Cruz, CA (OS) Nov 23 2016 1085 SSW David James
B2766 Male Yakima, WA Aug 21 2016 David James
Sept 2 2016
Lighthouse Field, Santa Cruz, CA (OS)
Branciforte Drive,
Nov 23 2016 (1) (Nov 25 2016 (2), Nov 30 2016 (3))
1085 SSW
David James, Aleece
Townsend, John Dayton
A6195 Male Elkton, OR
Barbara Slott/Kris Hendricks
Santa Cruz, CA (OS) Nov 23 2016 768 SSE David James
B2604 Female Yakima, WA Aug 20 2016 David James
B2682 Male Yakima, WA Aug 21 2016 David James
B2837 Female Yakima, WA Aug 22 2016 David James
San Leandro GC, CA
(OS) Nov 24 2016 1006 SSW David James Kale Road, Bolinas,
CA (OS) Nov 25 2016 998 SSW David James Kale Road, Bolinas,
CA (OS) Nov 25 2016 998 SSW David James
A6205 Female Elkton, OR
Sept 2 2016 Barbara Slott, Kris Hendricks
Sept 2 2016
Lighthouse Field, Santa Cruz, CA (OS)
Esalen Institute, Big
Nov 30 2016 (1) (Dec 30 2016 (2), Jan 6 2017 (3))
753 S John Dayton, David James
A6189 Male Elkton, OR
Barbara Slott, Kris Hendricks
Sur, CA (OS) Dec 2 2016 850 S Joe Billings
B3038 Male Yakima, WA Sept 2 2016 David James
Las Varas Ranch, Goleta, CA (OS)
Dec 14 2016 (1)
(Jan 7 2017 (2)) 1357 S Joe Billings
B3390 Female Yakima, WA Sept 18 2016 David James
B2832 Female Yakima, WA Aug 22 2016
Lighthouse Field, Santa Cruz, CA (OS)
Tecolote Canyon,
Nov 30 2016 (1) (Dec 9 2016 (2), Dec 30 2016 (3))
1085 SSW John Dayton, David James
A7393 Male Applegate, OR
David James
Aug 27 2016 Kirsten & Jakob Shockey
Goleta, CA (OS) Dec 14 2016 1360 S Jessica Griffiths
Cambria, CA (OS) Dec 29 2016 (1)
(Jan 1 2017 (2)) 761 SSE David James,
Joe Billings
A6879 Female Wilderville OR Aug 21 2016 LaDawn Wilhelm
Sept 2 2016
Purple Gate, Bolinas,
CA (OS) Jan 1 2017 502 S David James Ellwood, Goleta, CA
A6188 Male Elkton, OR B3555 Male Elkton, OR
Barbara Slott/Kris Hendricks
Sept 23 2016
Barbara Slott/Kris Hendricks
(OS) Jan 9 2017 1071 SSE Joe Billings
Dump Road,
Carpinteria, CA (OS) Jan 20 2017 1085 SSE Joe Billings
B3510 Male Applegate, OR Sept 25 2016 Linda Kappen
Lighthouse Field,
Santa Cruz, CA (OS) Feb 9 2017 597 S Bill Henry
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TABLE 7. California overwintering sites at which Pacific Northwest-tagged Monarchs were recovered during 2012–16
North
TABLE 8. Distance and daily rate of travel shown by 13 reared and tagged fall-migrating Monarchs.
Release
Overwintering site
Santa Cruz
Washington -released
Southern Oregon released
central
Oregon
released Total
Tag #
Release Recovery locations
Walla-Walla
Distance traveled (km)
Recovery period
(days)
Rate of travel
(km/day)
Lighthouse
Field 8 3 1 12 Santa Cruz
1837
WA – Brigham City UT
724 21 34.5
Natural Bridges 2 2 1 5
Santa Cruz
Moran Lake 2 1 1 4
Santa Cruz
Branciforte 0 1 0 1 Bolinas-Kale 2 3 0 5 Bolinas-Agate 0 1 0 1
260 Post Falls ID
– Potlatch ID 93 3 31.0
3172 Yakima WA –
Bolinas CA 998 43 23.2 Yakima WA –
Bolinas-Purple
Gate 0 1 0 1
Morro Bay
SPCG 3 0 0 3
Morro Bay-Del
Mar Park 0 1 0 1
Morro Bay-Toro
Creek 1 0 0 1 Carpinteria
7024 5741 0683
Glen Ellen CA
Stateline ID – Walla Walla WA
Applegate OR – San Mateo CA
1030 23 44.8 209 9 23.2 531 11 48.3
Dump Road 0 1 1 2
Carpinteria
Creek 0 0 1 1
Cayucos-Villa
Creek 0 1 0 1 Cambria 0 1 0 1 Esalen-Big Sur 0 1 0 1
A3328 Rathdrum ID
– Troy ID 124 12 10.3
A3223 Brookings OR
– Aptos CA 612 29 21.1 Applegate OR
A3771
Pacific Grove 1 0 0 1
– Boonville CA
365 18 20.3
Pismo Beach 0 1 0 1 Goleta-Ellwood 0 1 0 1
Goleta-Maria
Ygnacio Creek 1 0 0 1 Goleta-Las
Varas 1 0 0 1
Goleta-Tecolote
Creek 1 0 0 1 San Leandro
GC 1 0 0 1
A6093 Ashland OR –
Vacaville CA 457 8 57.1
A7079 Brookings OR
= Windsor CA 438 7 62.6
B2838 Yakima WA –
Anapolis CA 917 23 39.8 Corvallis OR –
Ardenwood
Farm 0 1 0 1
New Park Mall
Newark 1 0 0 1
A4853 MEAN
San Francisco CA
765 19 40.3
TOTAL 24 20 5 49
(±) SE 558.7 ± 88.0 17.4 ± 3.0 35.1 ± 4.3
VOLUME 72, NUMBER 2 141 TABLE 9. Release, recovery and distance data for a wild Monarch tagged in northern California in late spring 2015.
Tag # Sex Release location Release date Recovery location Recovery date
Eagle Creek Trinity
Distance
(kilometers) & Direction
707 ENE
Finder Kendra
4130 Male
River N CA May 24 2015 Twin Falls ID June 28 2015
(spring migrant)
Hathaway
TABLE 10. Tagged Monarchs seen multiple times at California overwintering sites during 2015–16. Number of
Residency at site
Tag # Overwintering site
times sighted First sighted Last sighted
(days)
A2067 Santa Cruz-Natural Bridges 2 Nov 16 2015 Nov 26 2015 10 A2062 Santa Cruz-Moran Lake 2 Nov 10 2015 Nov 26 2015 16 A0508 Santa Cruz-Moran Lake 2 Nov 26 2015 Dec 30 2015 34 A3264 Pismo Beach 2 Nov 24 2015 Dec 28 2015 34 4789 Bolinas-Kale 2 Nov 28 2015 Dec 10 2015 12
A2045 Bolinas-Kale 2 Jan 1 2016 Feb 3 2016 34 Santa Cruz-Lighthouse
B2701 A6935 B2452 B2742 B2766 A6205 B3390
Field 4 Sept 30 2016 Feb 1 2017 123
Santa Cruz- Lighthouse
Field 3 Nov 5 2016 Nov 30 2016 25
Santa Cruz-Lighthouse
Field 2 Nov 5 2016 Nov 25 2016 20
Santa Cruz-Lighthouse
Field 6 Nov 13 2016 Jan 6 2017 54
Santa Cruz-Lighthouse
Field 3 Nov 23 2016 Nov 30 2016 7
Santa Cruz-Lighthouse
Field 3 Nov 30 2016 Jan 6 2017 37
Santa Cruz-Lighthouse
Field 3 Nov 30 2016 Dec 30 2016 30
B3554 Ardenwood Farm 2 Nov 11 2016 Nov 17 2016 7 B3038 Goleta: Las Varas 2 Dec 14 2016 Jan 7 2016 24 A7393 Cambria 2 Dec 29 2016 Jan 1 2016 3
A6504 Carpinteria Creek 2 Nov 11 2016 Nov 26 2016 15 Carpinteria Creek
A6504
(Dump Rd) 3 Jan 11 2017 Jan 26 2017 15
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Yakima, WA were recovered either en route (1) or at
overwintering sites (9). This exceptional recovery rate
suggests that weather conditions during this narrow
window of time may have been extremely beneficial for
migration. The weather in southern Washington and
Oregon during the four week period following this
release (https://www.wunderground.com/) was mostly
warm (daily maxima 25–30 °C) and dry, conditions
highly suitable for migration. Prevailing winds during
the first two weeks of September in central and
northern Oregon were predominantly from the north
which clearly would have been favorable for tagged
Monarchs like B2838 released at Yakima on August 22
and seen at Anapolis in northern California on
September 14 2016.
FIG. 8. A Corvallis-Oregon-tagged Monarch (Tag A4853) dur ing its migration visiting flowers on the rooftop garden of Lisa de Angelis’ apartment in North Beach, San Francisco on Sep tember 18 2016. This Monarch was reared and tagged in Cor vallis, Oregon on August 30 by Molly Monroe and Amelia Je bousek (Photo: Lisa de Angelis).
more Monarchs in eastern areas of the PNW need to be tagged to obtain a sufficient number of recoveries to provide a better insight into the directions taken by migrating Monarchs in this region as well as their ultimate destinations. The lack of recovery of any wild tagged late summer/fall PNW Monarchs in this study is curious. A total of 822 wild Monarchs were tagged in Idaho, Washington and Oregon and if a 0.4% return was expected, then 3–4 individuals should have been recovered. Tagging greater numbers of wild Monarchs presumably should result in recoveries as it did in Arizona (Morris et al. 2015).
Yang et al. (2015) using spatial analyses of stable isotope ratios in the wings of 114 Monarchs overwintering at four California overwintering sites (Santa Cruz (2), Pismo Beach, Goleta) in 2009, showed great variation in the likely natal origins of these Monarchs. However, overall these data suggested an estimated 40 % of overwintering Monarchs originated in the eastern PNW (eastern WA and OR, ID). Our tagging data certainly support the notion of significant recruitment of California overwintering populations from the PNW.
Thirteen tagged Monarchs were reported 3–43 days after release and before they reached overwintering sites (Fig. 8). A mean travel rate of 35.1 km a day was recorded by these Monarchs which is a little lower than the estimated mean of 45 km/day provided by Brower et al. (2006) for a sample of 38 migrating Monarchs in the eastern US. In 2016, ten tagged Monarchs (3.7 %) from 271 released during a two day period (August 20–22) at
Washington and Oregon-tagged Monarchs were recovered in 24 coastal California overwintering colonies with most (33, 67.3 %) found in the San Francisco-Monterey area (Bolinas to Pacific Grove) sites. Twenty two (44.9%) of these recoveries were found in four Santa Cruz overwintering sites making this town the favored destination of PNW Monarchs. However, this was at least partly the result of regular inspection of colonies throughout each winter by local Monarch biologist John Dayton. Eleven (50 %) of the Monarchs found in Santa Cruz were reported by Dayton. One Santa Cruz site, Lighthouse Field, accounted for 12 tagged Monarchs with seven originating from the same backyard location in Yakima, WA. Nine tagged Monarchs were recorded in the mid coast area (Big Sur to Pismo Beach) and a similar number (7) were recovered from the Goleta Carpinteria area. Only three Monarchs were reported from overwintering colonies by people not associated with the project. In contrast, all 13 tagged Monarchs recovered during migration were reported by members of the public, unaware of the project. It is very likely that numbers of recoveries in the future could be increased further if volunteers living in the vicinity of overwintering sites could be recruited to regularly inspect overwintering colonies in the manner that John Dayton did in Santa Cruz during this project.
Seventeen tagged Monarchs were seen at overwintering sites on multiple occasions demonstrating residency at the sites for periods ranging from 3–123 days. However two Monarchs were sighted at more than one overwintering site and one of these was seen at three overwintering sites in the Santa Cruz area between October 11 and December 30 2016. Limited data on overwintering site fidelity and movement between sites has been reported (Nagano et al. 1993) but temporary residence at three overwintering sites in California by a single Monarch has not to our
VOLUME 72, NUMBER 2 143
knowledge been previously documented. Although our data on residency are limited they suggest that both types of behavior, high site fidelity and wandering occur in overwintering populations. High winds and storms likely influence inter-colony movement and this appeared to be a factor in the movement of A6504 from Carpinteria Creek to the nearby Dump Road site in 2016/17. The entire population at Carpinteria Creek dispersed in early January 2017 following a series of storms and at least some of the population (including A6504) reformed at Dump Road.
Fifty three wild spring migrants were tagged in the Trinity National Forest (along the Trinity River) in northern California in late May 2015 and 2016 and one (1.9 %) was recovered at Twin Falls in Idaho. Given the low number of individuals tagged, this recovery was unexpected and either indicates exceptional good luck or spring tagging of migrating Monarchs in northern California may yield more recoveries than tagging wild fall migrants. This is the first published record of a California-tagged spring Monarch recovered in the PNW. Nagano et al. (1993) tagged more than 50,000 Monarchs in 10 California overwintering colonies and recovered 100 at distances up to 465 km NW or E from release points, all within California and mostly during March–April. The migrants we tagged in northern California were the progeny of overwintered Monarchs. As with overwintered Monarchs it is likely that the late spring generation of migrants also migrates in northerly and easterly directions (as in Malcolm et al. (1993) for the eastern US population) with our single recovery an example of the latter. This broad range of compass directions would facilitate expansion of the Monarch population into Idaho, Utah and Montana as well as Oregon, Washington and British Columbia.
The research presented here supports the traditional assumption that Pacific Northwest Monarchs overwinter in California. Our data obtained over five years on recovery of tagged Monarchs released in the central and southern parts of both Washington and Oregon suggest that populations in these areas migrate to at least 24 overwintering sites along a 500 km section of the California coast. It seems likely that PNW Monarchs utilize the entire geographic range of coastal overwintering in California, certainly encompassing the span of noticeably large overwintering colonies from the San Francisco region to the Ventura region. Further tagging may extend this range to its northern limit in southernmost Mendocino County and/or further south to the San Diego area. We also obtained limited data indicating fall movement of some eastern Washington and Idaho Monarchs towards Utah and Arizona, reinforcing the earlier observations and suggestions of
Brower and Pyle (2004), Dingle et al (2005) and Pyle (2015) that some populations of Monarchs in eastern areas of the PNW may migrate to Mexico for overwintering. Increased emphasis on tagging reared and wild Monarch populations in eastern parts of British Columbia, Washington, Idaho, Nevada and Utah will be a priority of continuing citizen scientist-aided research on Monarch migration in the PNW. Recent interest by penal institutions in Idaho and Nevada to offer Monarch rearing opportunities to inmates similar to the Washington State Penitentiary model may enable rapid ramping up of tagged Monarch numbers in these states.
In addition to an improved understanding of Monarch migration in the PNW, this study also contributed meaningfully to the conservation of Monarchs by adding nearly 14000 butterflies to the population over five years. Penitentiary, University and private rearing in this study achieved survival rates way above the estimated 5–10% level that occurs in wild populations (Oberhauser et al. 2015). When survival from egg to adult was recorded in the Penitentiary mass rearings, rates generally ranged between 50 and 85%. The involvement of citizen scientists in this study rearing, catching and tagging Monarchs throughout the PNW generated much community interest in the study which in turn led to greater involvement by citizens. A Facebook page was established in 2012 and now has a following of more than 3800 people https://www.facebook.com/MonarchButterfliesInThePac ificNorthwest/. Increased awareness of Monarchs, their biology and conservation in the PNW has been an unexpected but welcome side-effect of this study.
ACKNOWLEDGMENTS
This research was made possible by an ‘army’ of citizen scien tist volunteers, including those citizens incarcerated at Washing ton State Penitentiary, Walla Walla. No specific financial support was provided or expended in this research except for unsolicited donations made by John Bittenger, Dick T. Price and Dave Hop per for the purchase of tags, for which we are deeply grateful. Our heartfelt thanks go to the following citizen scientists who reared and tagged Monarchs: Jennifer Andrew, Eva Antonijevik, Belinda Barnes, Norma Benson, Brenda Blakk, Kjersten Braaten, Maia Black, Betsy Bloomfield, Eddie Burke, Chris Carvalho, Andrea Christensen, Robert & Simone Coffan, Mavis Connor, Heather Crunchie, Roxie Crunchie, Maria Dietrich, Daniel De Bud, Dee de Bud, David Dube, Cindy Dunbar, Martha Effler, Patti Ensor, Patti Farris, David Fast, Lee Finney, Bud Gagner, Jeri Gleiter, Connie Granberg, Brad Grimm, Patsy Haggerty, Ling Helphand, Kris Hendricks, Linda High, Lauren Hisatomi, Celisa Hopkins, Scott Huber, Paul Huffman, Amelia Jebousek, Steve Johnson, Cynthia Kimel, Judith Kjellman, Jim & Bonnie Kizer, Gerry Lauby, Laila Lienesch, Tasha Leonard, Jacqueline Milikien, Amy Miller, Vicki Mion, Molly Monroe, Michael Moritz, Ann Mary Myers, Gary Pearson, Stephanie Reilly, Dana Ross, Lydia Roys, Jessica Roys, Joseph Roys, Vir ginia Rivers, Linda Ramsey, Brian Ruf, Statia E. Ryder, Suzie Savoie, Charity Shirley, Jakob Shockey, Kirsten Shockey, Bar bara Slott, Aleece Townsend, Dennis Triglia, Elzi Volk, Sandy
144144 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Wasson, Melanie Weiss, LaDawn Wilhelm, Susie Werts, Marty Witt and Tiffany Wyatt. Some of these citizen scientists also tagged wild Monarchs. We also thank the following personnel from Federal and State agencies for tagging wild and reared Monarchs: Shilah Allen (USFWS-intern), Brett Amdor, Bill Ament (IDFG), Tyler Archibald, LaDel Bonham (USFWS), G. Burak (USFWS), Ann Brueck, Becky Hansis-O’Neill (Idaho Museum of Natural History), Tracy Hart (USFWS), Leith Edgar (USFWS), Kat Findlay (IDFG), Sandi Fisher (USFWS), Jacky Friedman (USFWS-volunteer), Abby Goszka (BLM), Molly Hayes (USFWS-intern), Dave Hopper (USFWS), Marissa Jag ger (BLM), Chris Jensen (FS), Jennifer Jones (USFWS), B. Ki bler (USFWS), Akimi King (USFWS), Dea Kiss (USFWS-in tern), Kristin Lohr (USFWS), Russ Holder (USFWS), Hollie Leavitt (College of Western Idaho), Rose Lehman (FS), Vance McFarland (College of Western Idaho), Casey Mitchell, Jordan Mitchell (IDFG), Jeremy Moore (USFWS), Dan Nolfi (US FWS), Evan Ohr (USFWS), Ian Packham (IDFG), Dusty Perkins (College of Western Idaho), John Proctor (USFWS), Cody Prouse (IDFG), Melissa Reilly, Nate Richardson (US FWS), Dean Rose (IDFG), Barbara Schmidt (USFWS), Colleen Trese, Aaron Utz (Idaho Power Co.), Beth Waterbury (IDFG), Lydia Winton (IDFG), Ross Winton (IDFG), Kelly Weimer, Dwayne Winslow (USFWS), Josh White (IDFG) and Austin Young (IDFG). We also thank all the people who sighted our tagged Monarchs and reported them (credited individually within Tables 4-6). Special thanks must go to John Dayton of Santa Cruz and Joe Billings of MonarchQuestAZ for reporting 11 and 9 tagged Monarchs, respectively. Remarkably, between them they contributed a third of the recovery data reported in this paper! We also thank John for providing information annu ally on the Santa Cruz overwintering colonies. We thank Paul Cherubini, Bill Shepard and Craig Wakamiya for providing valu able information on overwintering colonies and Adrienne De Ponte kindly arranged for us to visit the San Leandro golf course colony. We are grateful to Mia Monroe for providing informa tion annually on the overwintering populations in Marin Co., CA and facilitating visits to some of the sites. Permits were obtained annually from the Washington Department of Fish and Wildlife to rear and tag Monarchs in that state and we thank Ann Potter for her guidance and direction. In many respects, the writings of and conversations with Robert Michael Pyle were the stimulus and guidance for this research and I (DGJ) humbly thank him for sharing his wisdom. Finally, we (DGJ, TSJ) thank our daugh ters, Jasmine, Rhiannon and Annabella for being an integral part of this study with their enthusiastic participation in our annual Thanksgiving and Christmas searches for tagged Monarchs in California. They also participated in the rearing of many Mon archs in our home.
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Submitted for publication 14 April 2017; revised and accepted 21 September 2017.