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Flower-Feeding by Two Species of Mydas Fly (Diptera: Mydidae) in Florida, with Notes on Tree-form Licania michauxii (Gopher Apple)
Alexander B. Orfinger, Sandor L. Kelly, and Derek A. Woller

Southeastern Naturalist, Volume 17, Issue 2 (2018): N19–N27

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N19 2018 Southeastern Naturalist Notes Vol. 17, No. 2 A.B. Orfinger, S.L. Kelly, and D.A. Woller Flower-Feeding by Two Species of Mydas Fly (Diptera: Mydidae) in Florida, with Notes on Tree-form Licania michauxii (Gopher Apple) Alexander B. Orfinger1,*, Sandor L. Kelly1, and Derek A. Woller2 Abstract - We present an account of flower-feeding by 2 species of mydas flies in Florida. We observed adults of Mydas clavatus and Mydas maculiventris feeding on flowers of both typical and unusually large (tree-form) Licania michauxii (Gopher Apple; Chrysobalanaceae). We also observed M. maculiventris feeding on flowers of Serenoa repens (Saw Palmetto; Arecaceae) in Florida scrub habitat. We discuss these observations following a brief review of feeding ecology in mydids, and include a checklist of the 12 known Florida mydids with a summary of our current knowledge of adult feeding in these species. We also provide additional sites of tree-form Gopher Apple in Martin County, FL, with a discussion of this unique form and the location of a new record-size specimen in Jonathan Dickinson State Park. Introduction. Mydas flies (Diptera: Mydidae) are a cosmopolitan insect family of modest diversity with an estimated 471 described species in 66 genera and 11 subfamilies (Calhau et al. 2014, Lyons and Dikow 2010). Adult mydids are medium- to large-bodied (9–60 mm long) and often appear to be Batesian mimics of stinging wasps (McAlpine 1981). Adult mydas flies are short-lived and seldom collected despite their global distribution and relatively large size (Lyons and Dikow 2010). Consequently, little is known of the natural history of adult mydids (Williams, 1995). Larval mydas flies are predaceous, often consuming Coleoptera larvae (McAlpine 1981, Papavero and Artigas 2009). However, the literature addressing adult feeding habits is mixed and many knowledge gaps exist for particular taxa. Whether adult mydids are predators, flower-visitors, or do not feed at all has been discussed for decades (Cole 1969, Kingsley 2002, Larson et al. 2001, Malloch 1915, Papavero and Knutson 1975, Papavero and Wilcox 1968, Proctor and Yeo 1973, Richter and Zaitzev 1988, Triplehorn and Johnson 2005, Wilcox 1981, Wilcox et al. 1989, Williams 1995). Olivier (1811) was the first to report adult mydids as predators, a notion reinforced by Paramonov (1950) and Genung (1959). However, Zikan (1944) observed male adult Gauromydas heros (Perty) attempting to copulate with males of its own species, other mydas flies, and hymenopterans of similar appearance, suggesting that such confused behavior may be misconstrued as predation. Zikan (1944) instead hypothesized, based on his observations of atrophied mouthparts of multiple genera,that mydids either do not feed as adults or, in the case of male G. heros, feed on flowers. Malloch (1915) collected adults on flowers but did not directly observe feeding. Davis (1921) reported being bitten by Mydas clavatus (Drury), which would indicate mouthparts adapted for biting, but was likely pinched with the femorae and tibiae (Williams 1995). Zaitlin (1978) and Zaitlin and Larsen (1984) further supported the idea of mydids as flower-feeders based on a lack of piercing mouthparts as adults, the fact that males and females have identical head morphologies, and, most importantly, because both sexes of multiple species were seen feeding on fl owers (Table 1). 1University of Central Florida Collection of Arthropods, Department of Biology, University of Central Florida, Orlando, FL 32816. 2USDA, APHIS, PPQ, CPHST Phoenix Lab, 3645 East Wier Avenue, Phoenix, AZ 85040-2931. *Corresponding author - aborfinger@gmail.com. Manuscript Editor: JoVonn Hill Notes of the Southeastern Naturalist, Issue 17/2, 2018 2018 Southeastern Naturalist Notes Vol. 17, No. 2 N20 A.B. Orfinger, S.L. Kelly, and D.A. Woller Focusing on Namibian mydids, Wharton (1982) speculated that it was possible that species of Afroleptomydas could nectar, based on Hesse’s (1969) observations of these flies collected on flowering Mesembryanthemum spp. (ice plants; Aizoceae). Wharton (1982) also suggested that the developed mouthparts of Parectyphus namibiensis Hesse might enable some species to feed as adults. Wharton (1982) further noted that many examined genera exhibit vestigial mouthparts and that these same genera were active during times outside of the flowering season in the studied region. Cannings’ (2006) thoughts on mydasfly feeding mirrored Wharton’s (1982) postulations, which were logical, but not conclusive. Proctor and Yeo (1973) and Larson et al. (2001) mentioned that some members of the family appear to nectar. Those authors provided no direct evidence; rather, they stressed that trophic ecology and pollinating abilities are poorly known and that direct observations are rare. Williams (1995) collected a female Mydas clavatus while it was feeding on flowers of multiple plant species (Table 1), confirming such feeding in adults of this species. Kingsley (2002) then described the endangered Rhaphiomidas terminatus abdominalis Cazier feeding on flowers of Eriogonum fasciculatum Benth. (California Buckwheat). Finally, Deyrup and Deyrup (2012) found 4 species of mydids visiting and feeding on flowers of Serenoa repens (Bartram) Small (Saw Palmetto, Arecaceae) in Florida (Table 1). Given the evidence of at least some mydids feeding on flowers, and the paucity of evidence supporting predation, it is no surprise that the Manual of Nearctic Diptera (Wilcox 1981:537) states that “the adult mydid is essentially a flower feeder”. Contradictorily, though, other textbooks and guides readily present adult mydas flies as predatory: for example, Borror and DeLong’s Introduction to the Study of Insects (Triplehorn and Johnson 2005) presumes that adult mydids are predaceous. The 2015 printing of Field Guide to Insects and Spiders (National Audubon Society 1980) goes even further, claiming that M. clavatus adults eat “caterpillars, other flies, bees, and true bugs”. To add to our still-sparse collective knowledge of mydid feeding behavior, we herein describe observations of 2 Florida species (M. clavatus and M. maculiventris Westwood) visiting flowers of a plant not previously noted in the literature as a food source: Licania michauxii Prance (Gopher Apple; Chrysobalanaceae). We also corroborate the findings of Deyrup and Deyrup (2012) with additional observations of M. maculiventris feeding on Saw Palmetto flowers. We include a checklist of the 12 known Florida mydas flies with a summary of their floral hosts as adults, and briefly discuss the exceptionally large (i.e., treeform) variety of Gopher Apple we observed because it is an evergreen species that does not normally exceed 6 to 12 inches (Ward and Taylor 1999). Methods. We observed M. clavatus and M. maculiventris feeding on Gopher Apple flowers on 16 and 17 May 2015, respectively, during trips to collect xerophilic grasshoppers for unrelated research in Florida scrub habitat. The first trip was to Jonathan Dickinson State Park (JDSP), Martin County, FL (WGS84 27°0'13.7268"N, 80°6'6.0942"W) and the second was near Frostproof, Polk County, FL (WGS84 27°39'57.9954"N, 81°33'33.7248"W). We were unable to collect the flies directly from the flowers, but we captured members of the same species with an aerial net within an hour of the observed flower-visiting at each site during the same outings. We made the observations of M. maculiventris individuals visiting flowers of Saw Palmetto on 27 May 2016 in Pinus clausa (Chapm. ex Engelm.) Sarg. (Sand Pine)–Quercus (oak) scrub habitat on the University of Central Florida (UCF) campus, Orlando, Orange County, FL (WGS84 28°36'39.6714"N, 81°12'19.2954"W), at which time we collected a voucher specimen with an aerial net while the fly was visiting the flowers. We made taxonomic identifications by referencing previously identified specimens in the UCF Collection of Arthropods (UCFC) and Florida State Collection of Arthropods (FSCA), and by using taxonomic keys. Mydas clavatus is distinct and cannot be confused N21 2018 Southeastern Naturalist Notes Vol. 17, No. 2 A.B. Orfinger, S.L. Kelly, and D.A. Woller Table 1. Checklist of the 12 species of Mydidae known to occur in Florida (in alphabetical order by genus and then specific name) and all known floral host plants of adults (if any). ABS = Archbold Biological Station, Highlands County, FL; * = reported as M. carbonifer (synonym); and ** = revision lists label data from several paratypes stating “at blooms of Sabal etonia”. Species Floral host References Mydas atratus Macquart Saw Palmetto Deyrup and Deyrup (2012)* Mydas brunneus Johnson None currently known N/A Mydas clavatus (Drury) Aloysia virgata (Ruíz & Pavón) (Sweet Almond Verbena; Verbenaceae) BubGuide (2017) Gopher Apple Present study Saw Palmetto Deyrup and Deyrup (2012) Asclepias syriaca L. (Common Milkweed; Asclepiadaceae) Zaitlin and Larsen (1984) Asclepias verticillata L. (Swamp Milkweed; Asclepiadaceae) Zaitlin and Larsen (1984) Asclepias spp. (milkweed; Asclepiadaceae) Malloch (1915), Williams (1995) Monarda punctata L. (Spotted Beebalm; Lamiaceae) Zaitlin and Larsen (1984) Pycnanthemum virginianum (L.) Rob. & Fernald (Common Mountain Williams (1995) Mint; Lamiaceae) Spiraea alba Du Roi (Meadowsweet; Rosaceae) Williams (1995) Mydas floridensis Wilcox, Papavero Saponaria officinalis L. (Soapwort; Caryophyllaceae) Zaitlin and Larsen (1984) & Pimentel Mydas fulvifrons (Illiger) Teucrium canadense L. (Canada Germander; Lamiaceae) Zaitlin and Larsen (1984) Mydas maculiventris (Westwood) Verbena hastata L. (Swamp Verbena; Verbenaceae) Zaitlin and Larsen (1984) Eupatorium mohrii Greene (Mohr’s Throughwort; Asteraceae) Unpubl. data from ABS Licania michauxii Prance (Gopher Apple; Chrysobalanaceae) Present study; unpubl. data from ABS Sabal etonia Swingle ex Nash (Scrub Palmetto; Arecaceae) Unpubl. data from ABS Mydas weemsi Wilcox, Papavero Saw Palmetto Deyrup and Deyrup (2012) & Pimentel Nemomydas jonesii (Johnson) Sideroxylon tenax L. (Tough Bully; Sapotaceae) Unpubl. data from ABS Nemomydas melanopogon Steyskal Sabal etonia Swingle ex Nash (Arecaceae) Wilcox et al. 1989** Phyllomydas parvulus (Westwood) None currently known N/A Phyllomydas quercus Wilcox None currently known N/A Phyllomydas weemsi Wilcox Saw Palmetto Deyrup and Deyrup (2012) 2018 Southeastern Naturalist Notes Vol. 17, No. 2 N22 A.B. Orfinger, S.L. Kelly, and D.A. Woller with any other species. Mydas maculiventris has multiple morphs, one of which is similar in appearance to the sympatric M. weemsi Wilcox, Papavero, & Pimentel. To verify the identity of this second species, we followed Wilcox et al. (1989) to key our specimens and confirmed them to be M. maculiventris. We generated the checklist of Florida mydids by consulting the scientific literature and identifying specimens in the UCFC and FSCA. We also conducted and summarized a review of all available scientific literature referencing flower-visiting in adult mydids that occur in Florida. We documented 2 new occurrences of tree-form Gopher Apple in JDSP, the first at the same site as the M. clavatus individual that we observed feeding and the second just to the southeast across the main park road (WGS84 27°2'6.3702"N, 80°6'52.65"W). We used a standard tape measure to determine length along the ground, width along the ground, and height at tallest point for the second Gopher Apple specimen. Imaging of the curated voucher specimen (Fig. 1) at UCFC was performed using a Visionary Digital imaging system equipped with a Canon EOS 6D DSLR camera combined with a 100-mm lens to take multiple images at different focal lengths. The resulting files were converted from RAW to TIFF format using Adobe Lightroom (v.4.4), and stacked into a single composite image using Zerene Stacker (v.1.04). Photographs of the live specimens (Fig. 2) were taken with a Pentax K5, a Pentax AF-540 flashgun through a soft box, and an old Kiron 105-mm 1:1 manual-focus macro lens. The photograph of the record size JDSP Gopher Apple plant (Fig. 3) was taken with a Canon PowerShot D20. For all images, as needed, we employed Adobe Photoshop CS6 Extended to add a scale bar, adjust light levels, background coloration, and sharpness. Results. In all observations included in this study, we observed individual mydas flies visiting multiple flowers on each plant, and using their mouthparts in a deliberate way that indicated feeding. The flies were most likely nectaring, but feeding on pollen cannot be ruled out. Either way, we documented individuals of M. clavatus and M. maculiventris feeding on flowering Gopher Apple on 2 separate occasions. During the first, on 16 May 2015, we observed M. clavatus feeding in Florida scrub habitat within JDSP. In this case, the flowering Gopher Apple patch was very dense, had a tree-form, and was groomed by park staff because it was located along the west edge of a parking lot just north of the park entrance. Flower-visitation occurred between 3 PM and 4 PM, at which time tthe weather conditions were hot and somewhat overcast. Figure 1. Mydas maculiventris (female) voucher, collected while visiting Saw Palmetto flowers on University of Central Florida campus, Orlando, Orange County, FL. (A) Lateral habitus. (B) Dorsal habitus. N23 2018 Southeastern Naturalist Notes Vol. 17, No. 2 A.B. Orfinger, S.L. Kelly, and D.A. Woller The second occasion happened the next day (17 May 2015) near Frostproof, FL. We observed M. maculiventris feeding on Gopher Apple, but this particular flowering patch was semi-dense, low to the ground, and on the outskirts of a scrubby area that appeared to be in the process of converting into an oak hammock. This site is on the outskirts of a fairly densely forested area set within a matrix of orange groves and connected to Saddle Blanket Lakes Preserve to the west. We observed the fly feeding between 11:30 AM and 12:30 PM; weather conditions were hot and sunny with clear skies. The following year, on 27 May 2016, we documented individuals of M. maculiventris feeding on flowers of Saw Palmetto in Sand Pine–oak-scrub habitat on the UCF campus. We decided to visit this site for exploratory purposes based on previous collection events of the species derived from the UCFC database (http://osuc.biosci.ohio-state.edu/UCFC/inst-full. html?id=146). We noted multiple individuals visiting several Saw Palmetto plants along a sandy trail between 11:00 AM and 12:00 PM; weather conditions were again hot and sunny with clear skies. An image of a voucher specimen—a female collected while feeding—is depicted in Figure 1. Flower-visiting by both of these Florida mydid species is depicted in Figure 2. Included in Table 1 is a checklist of all known Florida mydas flies and their known floral hosts, to which our observations have been added. We found 2 patches of tree-form Gopher Apple (Ward and Taylor 1999) in JDSP. The first is the plant where we observed M. clavatus feeding, but, unfortunately, a prescribed Figure 2. Live mydas flies on flowers: (A) and (B) Mydas clavatus on Aloysia virgata (Ruíz & Pavón) (Sweet Almond Verbena) in Sebring, Highlands County, FL; (C) and (D) M. maculiventris on Gopher Apple at Archbold Biological Station, Highlands County, FL. 2018 Southeastern Naturalist Notes Vol. 17, No. 2 N24 A.B. Orfinger, S.L. Kelly, and D.A. Woller burn destroyed this patch shortly after we became aware of it; thus, we made no formal measurements of patch size. Measurements of the second patch are as follows: 3.66 m long, 1.32 m wide, and 1.77 m tall (Fig. 3). Discussion. The observations provided here add further evidence of flower-feeding in adult Mydidae. However, the ecology of mydas flies remains poorly understood. Based on our findings, additional images of probable Florida flower-feeding on Bug Guide (2017) (e.g., Fig. 2A, B), and observations from Mark Deyrup (Insect Ecologist, Archbold Biological Station, Highlands County, FL), it is likely that mydas flies, at least in Florida, prefer to feed on a variety of flowers, possibly with a preference for light colored (or UV), open flowers. Future studies should further review publicly contributed mydas fly images on platforms like Bug Guide (2017) because they might reveal further insights into potential feeding trends of adult mydids as well as new plant records. We hope that further studies on these flies will aid in filling the plethora of existing gaps in our collective knowledge. For example, we believe the fact that at least some species feed on flowers has now been well-established, but feeding questions remain, such as: are the adults nectaring, eating pollen, or both? Furthermore, reports in the literature that adult mydids are predators must be investigated and verified or refuted. Our discovery of additional sites of tree-form Gopher Apple was quite serendipitous. The atypically large morph of this plant was not described until 1999 (Ward and Taylor 1999) when a small stand was found in Merritt Island National Wildlife Refuge near Titusville, FL. The tallest plant in this stand was 1.65 m tall. Until now, no additional sites have been noted in the scientific literature. The 2 JDSP sites recorded here are, therefore, unique Figure 3. The extant tree-form Gopher Apple patch, in foreground, within Jonathan Dickinson State Park, Martin County, FL. This unusually large specimen represents a new size record for the species at 3.66 m long, 1.32 m wide, and 1.77 m tall. N25 2018 Southeastern Naturalist Notes Vol. 17, No. 2 A.B. Orfinger, S.L. Kelly, and D.A. Woller and potentially merit further scrutiny. The extant site represents a new size record based on its height of 1.77 m (and assuming that the previous record has not grown taller since). Ward and Taylor (1999) provided 2 hypotheses to explain the existence of tree-form Gopher Apple in the refuge. The simpler of the 2 suggests that the plants are simply the result of existing in a fortuitous, fire-protected location. The second possibility is that the size is genetic in origin. The latter prediction is supported by the absence of similarly large individuals in nearby Gopher Apple patches that are also in unburned localities (see Ward and Taylor [1999] for a more thorough discussion). The proximate cause of tree-form Gopher Apple is unresolved and presents interesting questions for future investigation. In 2014, S.L. Kelly and D.A. Woller stumbled upon a third location that may also contain tree-form Gopher Apple, but measurements were not taken. The area was semi-developed, just outside and along the western perimeter of Railhead Scrub Preserve (in Collier County, FL, just south of Bonita Springs: WGS84 26°18'19.71"N, 81°47'29.6838"W), and possesses unusually dense patches that are definitely taller than average. No plants were observed to be higher than a knee (~0.5 m), but the area warrants further exploration and documentation. There is still much to learn about the biology of Florida mydids and tree-form Gopher Apple. Both taxa exist in imperiled Florida scrub habitat (Deyrup 1990, Olson and Dinerstein 2002, Sharpe 2010, Turner et al. 2006, Weekley et al. 2008). Therefore, it seems necessary to encourage relevant ecological studies to bolster our scant knowledge of the natural history of these intriguing organisms. Finally, just as protective measures were taken for the tree-form Gopher Apple stand in Merritt Island (Ward and Taylor 1999), it may be advisable to carefully manage the newly recorded extant stand in JDSP, given its unique physical (and potentially genetic) attributes. Acknowledgments. The authors wish to thank Mark Deyrup for providing unpublished flowervisiting records of Mydidae at Archbold Biological Station; Brian Silverman, of the UCFC, for imaging the voucher specimen of M. maculiventris; Charles Whitehill, Paul Skelley, and Gary Steck for providing access to the FSCA mydid collection; Tim Lethbridge for providing the wonderful live images of both species; JDSP Park Biologist Rob Rossmanith, who provided measurements and photos of the extant tree-form Gopher Apple patch; I. Jack Stout, of UCF, who first told the authors about the existence of tree-form Gopher Apple; Aileen Perilla and Ryan Ridenbaugh for being additional observers of feeding at the Frostproof site; and JoVonn G. Hill (Mississippi State University) and an anonymous reviewer for manuscript improvements. Literature Cited BugGuide. 2017 Mydidae. Available online at https://bugguide.net/node/view/695634/bgimage. Accessed 1 November 2017. Calhau, J., S.S Nihei, and C.J.E. Lamas. 2014. Taxonomic revision of Pseudorhopalia Wilcox & Papavero, 1971 (Insecta, Diptera, Mydidae, Rhopaliinae), with description of a new species from the Brazilian Amazon. Zootaxa 3884(4):333–346. Cannings, R.A. 2006. A review of the distribution and natural history of Apiocera barri and Nemomydas pantherinus (Diptera: Apioceridae and Mydidae), two rare asiloid flies from the southern Interior of British Columbia. Journal of the Entomological Society of British Columbia 103:55–60. Cole, F.R. 1969. 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