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A Long-Term Survey of Spring Monarch Butterflies in North-Central Florida

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A Long-Term Survey of Spring Monarch Butterflies in North-Central Florida Journal of Natural History ISSN: 0022-2933 (Print) 1464-5262 (Online) Journal homepage: http://www.tandfonline.com/loi/tnah20 A long-term survey of spring monarch butterflies in north-central Florida Lincoln P. Brower, Ernest H. Williams, Kelly Sims Dunford, James C. Dunford, Amy L. Knight, Jaret Daniels, James A. Cohen, Tonya Van Hook, Emily Saarinen, Matthew J. Standridge, Samantha W. Epstein, Myron P. Zalucki & Stephen B. Malcolm To cite this article: Lincoln P. Brower, Ernest H. Williams, Kelly Sims Dunford, James C. Dunford, Amy L. Knight, Jaret Daniels, James A. Cohen, Tonya Van Hook, Emily Saarinen, Matthew J. Standridge, Samantha W. Epstein, Myron P. Zalucki & Stephen B. Malcolm (2018) A long-term survey of spring monarch butterflies in north-central Florida, Journal of Natural History, 52:31-32, 2025-2046, DOI: 10.1080/00222933.2018.1510057 To link to this article: https://doi.org/10.1080/00222933.2018.1510057 Published online: 10 Sep 2018. Submit your article to this journal Article views: 59 View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tnah20 JOURNAL OF NATURAL HISTORY 2018, VOL. 52, NOS. 31–32, 2025–2046 https://doi.org/10.1080/00222933.2018.1510057 A long-term survey of spring monarch butterflies in north-central Florida Lincoln P. Browera#, Ernest H. Williams b, Kelly Sims Dunforda, James C. Dunfordc, Amy L. Knightd, Jaret Danielsc, James A. Cohenc, Tonya Van Hookc, Emily Saarinene, Matthew J. Standridgec, Samantha W. Epsteinc, Myron P. Zalucki f and Stephen B. Malcolmg aDepartment of Biology, Sweet Briar College, Sweet Briar, VA, USA; bDepartment of Biology, Hamilton College, Clinton, NY, USA; cMcGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, FL, USA; dFlorida Natural Areas Inventory, Florida State University, Tallahassee, FL, USA; eDivision of Natural Sciences, New College of Florida, Sarasota, FL, USA; fSchool of Biological Sciences, The University of Queensland, Brisbane, Australia; gDepartment of Biological Sciences, Western Michigan University, Kalamazoo, MI, USA Long-term springtime counts of immature and adult monarch but- ARTICLE HISTORY terflies and their Asclepias humistrata host plants in north-central Received 16 October 2017 Florida reveal a close relationship between the milkweed’sphenol- Accepted 1 August 2018 ogy and the butterfly’s spring remigration from Mexico. Remigrant KEYWORDS adults arrive after most frosts occur and as the milkweeds are flour- Population decline; spring ishing but before the plants begin to senesce. The peak of adult remigration; phenology; arrival is during the first few days in April; the eggs that are laid milkweeds; monitoring during this peak develop through April, leading to a second peak in adult abundance in early May. These newly emerged adults continue the migration northward. In addition to assessing the phenology of annual events, our long-term survey, with regular monitoring from 1994–2017, has enabled us to evaluate long-term trends. Both adults and immatures have declined in abundance from 1985 to 2017; since 2005, both have declined by around 80%. This decline has occurred concurrently with the decline in the number of monarchs at their Mexican overwintering sites. This paper presents the results of a long-term survey of early spring breeding by monarch butterflies (Danaus plexippus L., Lepidoptera, Nymphalidae) on Asclepias humis- trata Walt. (Apocynaceae, Asclepiadoideae) in north-central Florida. Early observations from 1981–1993 were followed by standardized surveys from 1994–2017. Consistent long-term monitoring of changes in monarch spring breeding are rare for this well- studied species, even though recolonization of the upper USA and Canada depends on spring breeding in southern states. Previous work on spring migration found a delayed return and smaller geographic coverage of the initial wave of spring remigration (Howard and Davis 2015), with both results likely a result of declining overall abundance. Here we add to an understanding of spring migration by addressing questions about the CONTACT Ernest H. Williams [email protected] #Deceased © 2018 Informa UK Limited, trading as Taylor & Francis Group Published online 10 Sep 2018 2026 L.P. BROWER ET AL. changing abundance of monarchs in north Florida and the phenological matching of milkweed host plants with monarch arrival and reproduction. Considerable research has supported the hypothesis that the early spring monarchs collected from Texas to north Florida are individuals returning from overwintering sites in Mexico (Lynch and Martin 1987, 1993; Malcolm et al. 1987, 1993; Martin and Lynch 1988; Cockrell et al. 1993; Riley 1993; Howard and Davis 2004). Monarchs are widely reported to reach the southern tier of states, including north Florida, during March (data for 2000–2017, Journey North 2017). However, the geographic origin of the spring monarchs in north-central Florida continues to be debated because, in order to arrive in this area in late March and early April from their Mexican overwintering sites, the monarchs would have to either fly from Mexico across the Gulf of Mexico, which has not been documented and seems unlikely for several reasons (Brower 1985, 1995), or, after reaching the Gulf coast in Texas, fly east along the coast and then south into north- central Florida. Alternatively, they could be monarchs that overwintered in colonies along the Gulf Coast and migrated inland in late March and early April. While some monarchs may have flown in from coastal overwintering sites, the distribution and occurrence of these sites is historically haphazard and their overwinter- ing numbers small (Brower 1995). Recent evidence supports the minimal influence of coastal sites; Journey North data on overwintering butterflies in nine coastal states over eight years, 2002–2009, summarized by Howard et al. (2010), reported 254 individuals but no overwintering clusters, with 80% of the observations in Texas and Florida. Furthermore, our extensive cardenolide fingerprinting data (Malcolm et al. 1993) together with over a century of natural history observations reviewed in Brower (1995) strongly support the hypothesis that the majority of these April Cross Creek monarchs bred in the north the previous summer on A. syriaca, overwintered in Mexico, and flew to Cross Creek in the subsequent early spring. Here we present the results of a long-term spring monitoring programme in which we recorded the phenology of a large population of Asclepias humistrata plants and the arrival time, abundance, and reproduction of monarch butterflies on this milkweed in north-central Florida. Early observations from 1983–1994 were followed by regular and consistent monitoring from 1994 to 2017. We give evidence that the abundance of the spring remigrant population has diminished through time, as have both the eastern and western North American overwintering populations (Brower et al. 2012; Xerces Society 2016; Pleasants et al. 2017; Schultz et al. 2017; Malcolm 2018), and that the arrival of monarchs is timed to coincide with the availability of their host plant A. humistrata. Methods Host plant and study site Asclepias humistrata, the sandhill or pinewoods milkweed, is the common milkweed in our study area; it is largely a south-eastern US coastal plains species occurring from south-central Florida at approximately 28°N latitude northwards to North Carolina at approximately 37°N latitude and westwards to eastern Louisiana at approximately 90°W longitude. It is a decumbent, herbaceous perennial plant that blooms from April to July, and its habitat includes well-drained soils on sand dunes, dry oak woods, and pine-scrub JOURNAL OF NATURAL HISTORY 2027 habitats (Woodson 1954). In north-central Florida A. humistrata is frequently abundant in overgrazed pastures and along disturbed roadsides. Based on its known US distribution (Woodson 1954; Kartesz 2015), we estimated the species range, all in the south-eastern USA, as 254,000 km2 (= 98,070 square miles). In contrast, Asclepias syriaca L., the principal summer foodplant of monarchs in eastern North America and Canada, has a much more extensive distribution (3,375,000 km2 = 1,303,095 square miles) (Woodson 1954; Crolla and Lafontaine 1996; Kartesz 2015). Thus, the distribution of A. humistrata is only 7.5% that of A. syriaca. The study area consisted principally of a cattle pasture (ca. 9 ha) at Cross Creek in Alachua Co., FL (N 29°31.75′, W 82° 11.86′), with more than 1000 A. humistrata plants (Figures 1, 2 and 3). According to the owner, Zane Hogan, the pasture had not been treated with herbicide for several years prior to and throughout the duration of our study. This allowed the survival of milkweed as well as other species undesirable for cattle, including prickly pear cactus, Opuntia humifusa Raf. (Cactaceae), tread-softly, Cnidoscolus stimulosus (Michx.) Engelm. & Gray (Euphorbiaceae), and yellow thistle, (Cirsium horridulum Michx. (Asteraceae). Grazing cattle avoided eating or damaging the milkweeds, which contain large amounts of latex and high concentrations of toxic cardiac glycosides (Martin et al. 1992). The pasture was surrounded on the north, east, and west by ‘improved’ pasture that contained few milkweeds or other nectar sources and to the south by County Road 346 and a commercially planted slash pine forest, Pinus elliotti Engelm. (Pinaceae). Most land in this region is either pasture or commercial slash pine plantation. There were several large longleaf pines, Pinus palustris Mill. Pinaceae, scattered through the pasture, and a ca. 20 m wide strip of live oaks, Quercus virginiana Mill. (Fagaceae), separated the study site from the neighbouring Figure 1. Satellite view of Hogan’s pasture (X) and the roadsides in Cross Ck, Alachua Co., FL, where the density, health, stage and number of Asclepias humistrata plants and stems and the spring breeding populations of monarch butterflies were assessed. 2028 L.P. BROWER ET AL. Figure 2. View across Hogan’s pasture showing numerous flowering Asclepias humistrata plants. 28 April 2010.
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