1986 Florida Field Trip Report

Ted C. MacRae & Rev. James M. Sullivan1

Fr. James Sullivan (left) and Ted MacRae (right) stand under a banyan tree in Key Largo, Florida, 9 May 1986 (photographer unknown).

In Spring 1986, the Rev. James Sullivan (Fr. Sullivan to those who knew him) and I made a two-week trip to Florida with the objective to explore as much of the state as we could—from the Panhandle to the Keys! My goal, of course, was to collect beetles2, especially woodboring beetles in the families Buprestidae and Cerambycidae (it was only my third trip outside of Missouri for such purpose), while Fr. Sullivan’s was to identify and document as many plant species as possible. This apparent dichotomy in interests was not as clean as it may seem—as an entomologist interested in host plant relationships, the chance to spend time in the field with as accomplished a botanist as Fr. Sullivan was too good to pass up, and Fr. Sullivan’s passion for studying insect associates of the plants he studied greatly aligned our interests. My memories of that trip have faced in the nearly 40 years since, jogged only by the specimens I collected now residing in my cabinet and two trays of 35 mm slides taken with an Olympus OM10 SLR film camera. Fortunately, Fr. Sullivan was more diligent than I in journaling his observations during that trip, a copy of which he gave to me. This report is an attempt to summarize our observations using these materials. In his journal, Fr. Sullivan made the following disclaimer: “Plant determinations in these notes must be regarded as tentative: We have not had the use of a complete flora for any portion of the State of Florida. We have been as precise as possible with the use of several less complete sources. ★This star symbol indicates plant determinations that were later confirmed by the use of the Flora of Tropical Florida or by other adequate keys.”

[Note: names for most of the plants included in this report were confirmed by Fr. Sullivan, the primary exceptions being those indicated with question marks (?). As a result, I have omitted use of the star symbol in this report. Also note that plant taxonomy has likely changed immensely during the past 39 years. Scientific names, common names, and plant families given are those we used at the time, and only in a few cases have they been harmonized with current nomenclature (my notes in [square brackets]). An asterisk (*) denotes plants and insects that were also photographed.]

We left St. Louis on the morning of April 28 with the goal of spending the night in Montgomery, Alabama. The next morning, true to form, Fr. Sullivan got up early to explore the area around the hotel before continuing the drive south. He noted two plants: Cirsium horridulum and Sapium sebiferum (Euphorbiaceae), the latter a fast-growing deciduous tree known as Chinese tallow and native to eastern Asia. All parts of the tree emit a milky white sap when damaged, which is toxic and can cause gastrointestinal upset if ingested. As we continued south, we noted the first Spanish moss (Tilandsia usneoides) draping the trees along Hwy 281. The growth was very well-developed, leading Fr. Sullivan to speculate that it also probably occurred north of Montgomery along I-65 but that we missed it due to our nighttime arrival.

Later in the day we arrived at our first destination, Torreya State Park in the Florida Panhandle, home of the extremely rare Florida nutmeg (Torreya taxifolia*) tree that grows only on the bluffs along the Apalachicola River. We explored the heavily forested hills and ravines of the area and found examples of this plant alongside the road near the entrance to the campground. However, we documented a diverse list of other plants including Acer barbatum [= A. saccharum ssp. floridanum] (Florida maple), Actaea pachypoda, Amorpha fruticosa, Aralia spinosa, Ascyrum sp., Callicarpa americana, Calycanthus floridus, Calycarpon lyoni, Cnidoscolus stimulosus*, Conopholis americana, Conradina canescens*(Lamiaceae), Croomia pauciflora* (Stemonaceae), Decumaria barbara, Dirca palustris, Erigeron strigosus, Erythrina herbacea*, Euonymus americanus*, Lonicera sempervirens*, Halesia sp., Hydrangea quercifolia*, Ilex opaca, Itea virginica*, Lygodium japonicum, Mitchella repens, Myrica serifera, Onoclea sensibilis*, Opuntia humifusa*, Rhaphidophyllum hystrix (needle palm), Ruellia carolinensis*, Sebastiana fruticosa* (Sebastianbush, Euphorbiaceae), Spiranthes praecox*, Viburnum sp., and Wahlenbergia marginata. Insect collecting consisted primarily of an assortment of longhorned beetles attracted to ultraviolet (UV) lights at night.

After two nights at Torreya State Park, we traveled further down the peninsula along the central spine to Highlands Hammock State Park, one of the oldest state parks in Florida protecting 9,000 acres of old-growth cypress swamp and oak hammock. We first explored the Wild Orange Grove Trail (noting wild orange trees as well as our first alligator!) but moved to other areas of the park over the next two days. We noted the occurrence of three species of palms here: Rhaphidophyllum hystrix (needle palm), which lacks the leaf midrib of and has fewer leaf divisions than Sabal palmetto (cabbage palm), the most common palm and distinguished by a strong leaf midrib, and Serenoa repens (saw palmetto) with its saw-toothed petiole edges. We repeatedly saw the tortoise beetle Hemisphaerota cyanea on the leaves of S. palmetto. A nice variety of longhorned beetles was also collected here, including the Florida endemic Typocerus flavocinctus and several lamiines at UV lights at night. A blister beetle (family Meloidae) similar to our Nemognatha nemorensis was common on the flowers of Aster reticulatus* and Pterocaulon pycnostachyum* (both Asteraceae), and on flowers of Cirsium horridulum we saw the leaf-footed bug Acanthocephala terminalis* (Coreidae). Other plants that Fr. Sullivan noted include Abrus precatorius*, Ardisia crenulata* (crenate berry bush, Myrsinaceae), Asimina pygmaea* (?), Bacopa sp.*, Bidens pilosa*, Callicarpa americana*, Cuthbertia graminea (?), Emilia sonchilfolius, Eriocaulon sp.*, Erythrina herbacea, Hypericum sp.*, Ilex cassine, Ilex glabra, Lachnocaulon anceps, Lygodesmia aphylla*, Lyonia lucida*, Mikania scandens, Oxalis violacea* (?), Persea borbonia, Polygala lutea*, Schrankia microphylla*, Tephrosia chrysophylla, Urena lobata, Utricularia sp.*, and Xyris sp.* Fr. Sullivan also noted in his journal a list of a dozen “rare” birds such (e.g., cardinal, catbird, crow, etc.!).

Our next destination was outside the northwestern limits of Everglades National Park at Collier-Seminole State Park, which lies partly within the great mangrove swamp of South Florida (one of the largest mangrove swamps in the world) and covering one of three original stands of royal palm (Roystonea elata [= Roystonea regia]) in Florida (the park was previously called Royal Palm Hammock). We primarily explored the Royal Palm Hammock Nature Trail and along the water’s edge around the boat basin, where Rhizophora mangle* (red mangrove) lined the edges of the salt marsh. Two species of Solanum were observed, primarily S. erianthum (potato tree) but also S. donianum*, and we noted the pleasant fragrance of a Eugenia sp. that escaped identification. Bursera simaruba (“tourist tree”), with its distinctive peeling bark, was also common here. We noted Baccharis halimifolia heavily infested with the leaf beetle Trirhabda bacharidus*, saw Heliconius charitonius butterflies on the wing, and observed a cluster of young seed bug nymphs* (Lygaeidae), likely one of the milkweed-associated species, on a vining species of milkweed. Deer flies (family Tabanidae) were a real problem for both of us, and we had to use head nets (Fr. Sullivan even resorted to wearing his London Fog jacket!). Other plants documented included Acrostichum sp., Alternanthera sp. poss. philoxeroides (Amaranthaceae), Batis maritima, Bidens pilosa var. radiata, Blechum brownei, Borrichia frutescens*(sea daisy), Commelina diffusa, Dicliptera assurgens*, Dicromena sp.* (white-bracted sedge, Cyperaceae), Eugenia sp.*, Ipomoea alba (moon flower), Ipomoea sagittata*, Passiflora pallens, Pithecellobium unguis-cati (cat claw), Pluchia odorata (camphor weed), Polygala grandiflora var. angustifolia, Psychotria undata*, Solidago sp.*, Triodanus sp., Urena lobata*, and Zanthoxylum fagara (lime prickly ash). Similar to previous localities, a diversity of longhorned beetles were attracted to UV lights at night.

After two days at Collier-Seminole, we drove east along the Tamiami Trail, noting the magnificent stands of bald cypress (Taxodium distichum) in the Big Cypress Swamp Preserve and seeing the first water hyacinths (Eichhornia crassipes) in bloom. We also saw Australian pines (several species in the genus Casuarina)—angiosperms rather than gymnosperms. Its needles are much longer than true pine (genus Pinus), and the trees appeared very dark green as seen from a distance. Eventually we landed at John Pennekamp Coral Reef State Park on Key Largo. Most people visit this park to dive and explore the spectacular living reefs of the Florida Keys; however, we had more terrestrial objectives. We began by exploring the coral limestone woodland along the Wild Tamarind Trail, where Metopium toxiferum *was common along the woodland border (and we took care not to touch!). A large ichneumonid wasp* (Ichneumonidae) was seen perched in the understory, and we noted the impressively oversized female of the spider Nephilia clavipes* being courted by an equally strikingly diminutive male. Lysiloma latisiliqua* was abundant in the woodland, as was Bursera simaruba* its bark red and peeling like a sunburned tourist! Coccoloba uvifera was also abundantly fruiting, and other plants seen include Alternanthera sp., Conocarpus erecta var. sericea, Schinus terebinthifolius*, and an unidentified composite (either Eupatorium villosum or Garberia sp.). Along the Mangrove Trail we saw (of course) not only red mangrove Rhizophora mangle*) but also black mangrove (Avicennia germinans*, Avicenniaceae) and white mangrove (Languncularia racemosa, Combretaceae). These three plants are placed in three unrelated families, yet all show a high degree of fidelity to mangrove ecosystems. At a Persea americana* orchard in the adjunct Shaw Property, we saw Hamelia patens* in bloom and Lysiloma latisiliqua growing around the orchard’s edge. Other plants seen in the area include Abutilon sp., Batis sp.*, Eupatorium villosum* (?), Gaillardia pulchella*, Heliotropium angiospermum, Heliotropium curassavicum, Hibiscus tiliaceus, Melanthera sp., and Rivina humilis*.

Our plan the following day was to continue down the length of the Keys, making stops at a few selected places along the way before spending the night in Key West. At our first stop on Lower Matecumbe Key, we noted Avicennia germinans, Eustoma exaltatum*, and Polygala baldwini (as well as a Great Blue Heron) and then continued southwestward to Long Key State Recreation Area [now Long Key State Park]. Most people visit Long Key for its beaches and fishing, but we came to the preserve to explore the endangered coastal dune ecosystem that it protects. Few insects were seen, but a number of interesting, mostly highly salt-tolerant, plant species were seen. These include Abutilon sp., Argemone mexicana*, Cassasia clusiifolia*, Chrysobalanus icaco, Ipomoea pes-caprae, Lantana involucrate, Manilkara bahamensis* (wild dilly, Sapotaceae), Scaevola plumieri* (inkberry, Goodeniaceae), Solanum diphyllum, and Suriana maritima* (bay cedar). We finished the day at Key Deer Preserve on Big Pine Key, a sanctuary for the smallest subspecies of white-tailed deer in North America. We did not see any deer, but we did see some interesting plants. Byrsonima cuneata* (Malphigiaceae) was common here, as was Croton linearis. Along the Nature Trail we saw the orchid Bletia purpurea* (pale pink) and also recorded Aletris sp., Chrysophyllum oliviforme (Sapotaceae), Dichromena sp., Ernodia littoralis*, Metopium toxiferum, Pithecellobium sp., Polygala verticillate (?), and “thatch palms” (plus a hissing alligator!).

The next two days were spent at Everglades National Park, where we began our visit by exploring the Long Pine Key Nature Trail, where Cladium jamaicensis* (sawgrass) and Taxodium distichum* dominated the landscape. We noted that it was easy to pass one’s fingers over the sawgrass blade edge in one direction, but not so easy in the other! It was here that I found what I considered a real prize—my first ever bumelia borer (Plinthocoelium suavelons)! I also collected the very colorful Trichodes apivorus on the flowers of Sabal palmetto. Fr. Sullivan had even more success with the plants—so much, in fact, that we were only able to explore the east end of the trail. Several plants belonging to largely tropical plant familys were seen, including Dodonaea viscosa (varnish leaf, Sapindaceae), Tatrazygia bicolor* (Melastomataceae), and Dipholis salicifolia (willow bustic, Sapotaceae). Polygala balduinii (or a similar species) and an unidentified Buchnera sp. were common. The recorded list of other plant species seen was diverse: Asclepias lanceolata*, Baccharis sp., Byrsonima cuneata, Calopogon sp.* [likely C. tuberosus var. simpsonii], Croton linearis, Dichromena sp., Heliotropium polyphyllum var, polyphyllum (H. leavenworthii ) (Boraginaceae), Jacquemontia jamaicensis, Lobelia glandulosa*, Melanthera angustifolia (Asteraceae), Myrica cerifera, Myrsine guianensis, Passiflora sp.*, Persea bordonia, Piriqueta caroliniana, Psychotria nervosa, Rhus sp., Sabatia sp. poss. brevifolia*, and Stillingia sylvatica ssp. tenuis (Euphorbiaceae). We returned again to Long Pine Key Nature Trail the following day to explore the west end near Pine Glades Lake, finding many of the same plants recorded the previous day but also Ageratum littorale (?), Bletia purpurea, Justicia ovata var. lanceolata*, Lippia stoechadifolia, and Morinda royoc. From there we moved on to the P.K. Nature Trail, where Cynanchum blodgettii was seen twining over much of the vegetation—including other plants of its own species!. Fr. Sullivan spent a good deal of time studying a plant found growing at the edge of Pine Glades Lake, which he presumed to be a species of Lippia that exhibited pleated leaves with matching teeth (leading him to call it “corduroy lippia” or “pleated lippia”). Eventually he settled (and later confirmed) the species as Lippia stoechadifolia, a Neotropical species limited in the U.S. to south Florida and the Keys. Other plants observed included Angadenia berterii, Urechites lutea, and (my favorite) Zamia floridana* [likely Z. integrifolia var. silvicola].

Back at Key Largo near Tarpon Bay (below our motel), Fr. Sullivan continued exploring the plants, especially the mangroves. He noted that Avicennia has “dewdrops” but that the other mangroves do not. This relates to the processes used by the plant to eliminate excess salt, which in Avicennia involves salt water “perspiration” that dries in the heat of the day (indeed, the residue of salt flecks is useful in distinguishing Avicennia from the other mangroves) but in Rhizophora is done by accumulating salt in the oldest leaves before they turn yellow and drop. He also noted that Avicennia and Rhizophora can be distinguished by color; Avicennia, which normally grow a little farther from the water, are closer to gray-green, while Rhizophora are closer to yellow-green. Tridax procumbens was a common roadside weed around the motel—its flowers and fluffy seedheads rise on long scapes, as if leafless, but are actually attached to the sprawling, hairy stems, which bear many deeply cleft leaves with opposite arrangement. He also noted Morinda royoc growing not only in the woods but also hedgerows. It is like Psychotria [both species are in the Rubiaceae], but without the large, nervy leaves. The fruits aggregate to look like large, yellowish mulberries. Hamelia patens grows right along the highway here, and several large Solanum shrubs with stellate trichomes on the leaves and white flowers were seen that may be a complex of species including S. donianum, S. verbascifolium, S. erianthum, or yet another species.

On May 10, the field visits were over, and it would take two days of driving to return to St. Louis. Even beginning the drive home, however, did not stop Fr. Sullivan from botanizing. During a stop at the drawbridge on Hwy 1 between Key Largo and the mainland, Fr. Sullivan collected Stachytarpheta jamaicensis, its flower tubes emerging from upward pointing, elongate triangular bracts, and its leaves being coarsely dentate. Also, from the highway in northern Florida, we saw what appeared to be the frequent occurrence of blooming Asimina. These were low plants with white flowers and leaves present. Spending several days in south Florida also gave Fr. Sullivan a chance to contemplate the different hammock habitats that we had visited, and he noted the following: “A hammock is basically a hardwood forest. A hammock in Florida has a significance parallel to that of a glade in Missouri: it is a relief from the ordinary situation. In the Everglades hammocks take the form of “hillocks”: The forest seems to build itself above the level of the sawgrass wetlands. In Highlands County, on the other hand, the hammocks occupy depressions in the topography. It is natural for the pinelands to burn with some degree of regularity, but the wet depressions are protected from most fires. Fire actually helps Pinus to have a competitive edge (since the needles make good tinder and the resin burns so hot, the pines contribute to their own survival situation), but in the wet depressions the broadleaf hardwoods are able to take over. The State Parks often feature the hammocks. As we go farther south, the hardwood species become more tropical. We have seen a lot of Bursera simaruba, a hammock feature, but have yet to see Ceratiola ericoides, which is more a representative of the norm for this state. Visiting Mahogany Hammock in the Everglades we learn that it is protected from sawgrass fires by a natural moat surrounding it. The moat results as limestone strata are eaten away by the hardwood-produced acids.”


1 Deceased April 15, 2025.

2 Permits for collecting beetles were obtained from the Florida Department of Agriculture and the National Park Service.

© Ted C. MacRae 2025

Birthday beetles

It’s my birthday, and as I’ve done almost every year of my adult life I did my first “real” collecting trip of the season. Since we’re down in Florida visiting Madam’s sister, I was especially excited for the chance to see new ground and hopefully find a few Florida specialties. The day started by getting up early to watch a spectacular full-moonset over the ocean—something I’ve never seen before.

Full moon over the Gulf of Mexico.
Nearing the horizon.
Touchdown!

I then headed up to the Citrus Wildlife Management Area of Withlacoochee State Forest to hook up with fellow beetle enthusiast (and soon-to-be book chapter coauthor) Kyle Schnepp from the Florida State Collection of Arthropods. Although Kyle and I have previously met at society meetings, it was our first time in the field together, and I was keen to take advantage of his local expertise to help me in my quest for Florida-endemic species. The habitat was dry oak/pine sandhill woodland with a grassy understory dotted with Opuntia australis (Florida pricklypear cactus)—itself an endemic plant.

Opuntia australis (Florida pricklypear cactus).

Temperatures were comfortably cool starting out due to an overnight cold front (who knew there was such a thing in Florida?), and I was filled with anticipation as I arrived at the meeting point and prepared my gear and greeted Kyle and his two companions shortly afterwards. I’d hoped to encounter Brachys leafminers, and we beat the oaks, both Quercus laevis (turkey oak) and Q. geminata (sand oak), exhaustively in search of such, but none were found. At one point, I pulled out the telescoping tropics net to see if I would have better luck accessing the upper canopy, but I had no better luck. Through it all, however, I did get a few other miscellaneous beetles and treehoppers on the former and the same on the latter along with an Anthaxia sp. Kyle also collected a Mastogenius sp. on the latter, which I hoped to encounter as well but never did, while his companions encountered single specimens of Agrilus rubroniger (a very nice find—I have but a single specimen that I collected in southwestern Missouri) and Chrysobothris chrysoela. I continued to beat the oaks but increasingly turned my attention to other things, including the pricklypears. The flowers were closed at first (due either to the morning hour or cool temperatures), but peeling back the petals I encountered a mating pair of Trichiotinus rufobrunneus (Florida flower scarab)—a Florida specialty!—plus one of the smaller, more widespread Trichiotinus lunulatus.

Trichiotinus rufobrunneus (Florida flower scarab—family Scarabaeidae) on flower of Opuntia australis (Florida pricklypear cactus).

I pulled the petals back on a few other flowers but didn’t see anything, then encountered a fallen dead branch of Pinus clausa (sand pine). Carefully lifting and turning over the branch revealed a Chrysobothris cribraria on the branch underside—just the second buprestid of the day for me. I then ran into Kyle again, who mentioned he’d seen a Mecas sp. on a grass stem, so I began sweeping the more open areas where the grasses—still mostly last season’s dried growth—and found two in my very first pass. These proved to be the largely endemic Mecas femoralis! I would continue to sweep grasses as I traversed the open areas of the woodland, eventually collecting a nice series of 12–15 specimens. As I was sweeping, I noticed isolated plants of the endemic Lupinis ocalensis (Ocala lupine) and, on the foliage of one, a mating pair of lixine weevils that proved to be the endemic Scaphomorphus subcylindricus.

Scaphomorphus subcylindricus (family Curculionidae)—on foliage of Lupinis ocalensis (Ocala lupine).

I continued to the south side of the road where I’d earlier seen the pricklypears in hopes of finding the flowers now open. Such was the case, and in addition to further specimens of T. rufobrunneus I also encountered Acmaeodera ornata and Strangalia strigosa—the latter yet another Florida specialty that I’d not previously encountered.

Strangalia strigosa (family Cerambycidae).

Additional sweeping not only increased my series of M. femoralis but also turned up a mating pair of Typocerus zebra (zebra flower longhorn), while Kyle encountered a single Lycochoriolaus lateralis—apparently a mimic of lycid beetles, distasteful to predators, and yet another species I’d never encountered before. After nearly five hours, I decided to call it a day and started working my way back to the car. Along the way, I encountered a patch of Serenoa repens (saw palmetto) in flower. There was quite a bit of insect activity buzzing around the inflorescences, and a closer look revealed a variety of bees and a few Zelus longipes (milkweed assassin bugs) but few beetles of then oedemerids (false blister beetles). Then I noticed something “lycid-like” approaching the inflorescence, but something about it gave me pause. When it landed, I realized it was another L. lateralis. I was able to pick it up easily (it was not skittish as are most other longhorned beetles, perhaps because it relies on its mimicry to avoid predation), and further searching resulted in a fine series of nearly a dozen individuals, along with one more S. strigosa. Those would be the last insects collected, as beating oaks along the way back to the car still turned up nothing. I suppose I’ll need to make another trip down here if I want to collect Florida Brachys!

Typocerus zebra (zebra flower longhorn—family Cerambycidae).

On the way back south (though not exactly “on the way home”), I stopped off at the Richloam Wildlife Management Area of Withlacoochee State Forest. Unlike the dry sandhill forest I’d just visited in the Citrus unit, the habitat here is wet sand forest. I was hoping to find stands of sedges (family Cyperaceae—usually found in and along wetlands) that I could sweep in the hopes of finding species of Taphrocerus—a genus of Buprestidae whose North American fauna I’ve been slowly revising over the years. These so-called “sedgy-wedgies” mine the leaves of various species of sedges, and there are still a few species of the genus that I haven’t yet myself collected—including the endemic T. floridanus. Kyle had recommended a location where I might find such sedges, which were abundant along the roadside. I swept them thoroughly, but to no avail.

As I was sweeping, a truck stopped and the driver called out to me and said “I just wanted to let you know I’ve already seen four or five good-sized rattlers here!” I thanked the man for his warning—though in reality I very well may have seen more rattlers in my lifetime than him. The man lingered as I returned to what I was doing—seemingly disappointed or surprised that his warning didn’t result in my immediate withdrawal. I suppose he was expecting me to run back to the car screaming like a little girl who’d just seen a spider. I was actually really hoping to see a rattler afterwards and was disappointed that I did not. The forest at this point was impenetrable, so I walked about a quarter-mile down the roadside, checking the flowers of Rosa palustris (swamp rose) along the way and being surprised to not see a single beetle—or any other insect, for that matter—upon them. In fact, the only insects I saw on anything were two very large “caterpillars”—actually larvae of Cimbex americanus (elm sawfly), a primitive wasp—on Salix caroliniana (Carolina willow).

Cimbex americanus (elm sawfly—family Cimbicidae) on Salix caroliniana (Carolina willow).

I crossed the road to walk the other side and soon ran into an Ilex cassine (dahoon holly) tree in full flower that was leaning over the roadside.

Ilex cassine (dahoon holly).

I saw a mating pair of Lyconotus lateralis (family Lycidae) on the flowers—their presence alerting me to the possibility of finding their longhorned beetle mimic, Elytroleptus floridanus. Despite finding perhaps a dozen lycid beetles, I found no E. floridanus. Another holly tree further up the road also hosted lycids but no longhorned beetles.

Lyconotus lateralis (family Lycidae) on flowers of Ilex cassine (dahoon holly).

Continuing a quarter-mile the other way past the car and further sweeping of sedges along the way back to the car was fruitless, and the lateness of the hour told me my insect collecting activities for the day had finally come to an end. Neverthess, I was content with series of several nice beetle species in my bottles, including a few true Florida specialities, and made my way back home.

©️ Ted C. MacRae 2024

T.G.I.Flyday—Black horse fly (Tabanus atratus)

In my previous post, I talked about a day trip to a sand scrub remnant in the Lake Wales Ridge of central Florida to find and photograph the endemic Highlands tiger beetle (Cicindelidia highlandensis). Ironically, the dry sand scrub/pine woodland habitats along this ridge are dotted with small lakes and ponds, allowing a rich aquatic flora and fauna to co-exist alongside the xeric specialists. Field mate Chris Brown and I had found and just finished photographing the tiger beetles when we encountered this rather largish lake—bright, white sand surrounding crisp, clear water reflecting white, puffy clouds in a deep, blue sky. ‘Twas a spectacular sight, indeed!

Sand scrub lake

Sand scrub remnant, Lake Wales Ridge, central Florida

As we stood looking at the scenery, I noticed something black on the stem of one of the sedges growing along the water’s edge. Something big and black! As I moved closer I could tell quickly that it was a large horse fly, but it was not simply perched and resting on the sedge—there was something else going on. Moving closer, ever so cautiously so as not to disturb the fly, I eventually realized that it was a female in the act of oviposition. How cool—I’d never witnessed this before with any species of horse fly, so to see it with such a large species was a real treat. I recognized it instantly as Tabanus atratus—commonly called the black horse fly and recognizable as such by its large size, all-black coloration, and distinctively hooked antennae (see 3rd photo below).

Tabanus atratus ovipositing

A Tabanus atratus female oviposits on a stem overhanging the water.

Before we get to the eggs, let’s dispel some misinformation that seems to persist regarding the size of this species (as it does with almost any large insect). Black horse flies are undeniably large, and in fact they are one of the largest horse flies in North America. The more credible sources (e.g., Pechuman et al. 1983, Long 2001) cite body length as ranging from 20–25 mm (up to a full inch in length). Incredibly, the species does not take the honors as North America’s largest horse fly, which goes instead to Tabanus americanus and it’s upper limit of 30 mm (in fact, T. americanus may be the world’s largest horse fly)! There are, however, on-line sources and a few popular field guides (as cited in BugGuide) that state a maximum length of 28 mm for T. atratus. How credible this figure is I cannot say, but I guarantee that the size indications of 30, 40, and even a whopping 50 mm in length found routinely among photos of this species on BugGuide were not derived from careful measurement and almost certainly instead reflect the astonished reactions that such an abnormally large insect can generate! In fact, there are precious few insects in North America that reach lengths as grand as 50 mm (i.e., two full inches)!

Tabanus atratus ovipositing

Lateral view of oviposition.

We approached carefully, again so as not to disturb the female in the middle of her act, and we watched and photographed as she laid the individual eggs one by one, using the tip of her abdomen to carefully arrange them neatly against each other in stacked layers. From a photographic perspective, balancing flash exposure of the all-black adult with the bright-white egg mass presented a real challenge. Added to that was an additional exposure challenge (my desire for a blue-sky background), making it a truly difficult-to-photograph subject. Long (2001) states that T. atratus egg masses can contain anywhere from one hundred to a thousand eggs each, always near water’s edge or somewhere quite close to water. Females are capable of laying three or four of these egg masses, which apparently gradually turn dark as the eggs develop and approach hatch.

Tabanus atratus egg mass

Freshly laid Tabanus atratus egg mass.

Despite this being the first time I’ve ever witnessed oviposition by this species, it seems to be encountered regularly. There are several photos of ovipositing females among the many photos of this species that have been posted to BugGuide. Moreover, descriptions of the egg mass of T. atatus appeared very early in the literature, first by Hart (1895) and then in photographs by Schwardt (1936). The latter author also states “T. atatus deposits its eggs in masses which are so constant in structural plan as to make specific determination of the egg mass readily possible” (as quoted in Bailey 1948). Thus, even if this female had already finished and left her egg mass, it still could have been identified to species.

REFERENCES:

Bailey, N. S. 1948. Notes on Tabanus atratus subsp. nantuckensis Hine (Diptera). Psyche 55(3):131–138 [pdf].

Hart, C. A. 1895. On the entomology of the Illinois River and adjacent water. Illinois State Laboratory of Natural History Bulletin 4:149–273 [eBook].

Jones, C. M. & D. W. Anthony. 1964. The Tabanidae (Diptera) of Florida. U.S. Department of Agriculture, Agricultural Research Service, Technical Bulletin No. 1295, 85 pp. [pdf].

Long, W. 2001. Tabanus atratus (on-line), Animal Diversity Web. Accessed 20 March 2019 at https://animaldiversity.org/accounts/Tabanus_atratus/

Pechuman, L. L., D. W. Webb & H. J. Teskey. 1983. The Diptera, or true flies, of Illinois 1. Tabanidae. Illinois Natural History Survey Bulletin 33(1):1–121 [pdf].

Schwardt, H. I. 1936. Horseflies of Arkansas. Arkansas Agricultural Experiment Station Bulletin 332:14–15, 27–32.

© Ted C. MacRae 2019

 

Revisiting the Highlands tiger beetle (Cicindelidia highlandensis)

In September 2016, I had the opportunity to attend the Annual Meeting of the Entomological Society of America in Orlando, Florida, which was being held in conjunction with the International Congress of Entomology. My first thought when I made plans to attend these meetings was that this would be a chance for me to get another look at the Highlands tiger beetle (Cicindelidia highlandensis). One of Florida’s rarest endemic tiger beetles, this species is restricted entirely to remnant sand scrub and pine woodland habitats along the Lake Wales Ridge of Polk and Highlands Counties in central Florida (Choate 2003). I was thrilled to have found adults (in good numbers) on my first attempt back in 2009, and I was also thrilled to have successfully managed to photograph the beetle at that time. However, in the years since, I have become increasingly dissatisfied with those photographs—taken during what was my very first year of insect macrophotography. I’ve learned a lot since then about lighting, diffusion, and composition, but perhaps the biggest annoyance of those photographs is the fact that in every one the antennae and/or legs are “clipped”—a result of my being so enamored with my newfound macrophotographic capabilities that I nearly completely ignored other aspects of photographic composition.

Chris Brown photographing Cicindelidia highlandensis

Chris Brown photographs a Highlands tiger beetle.

Chris Brown—long-time field accomplice and himself a tiger beetle aficionado and insect macrophotographer—was also at the meetings, and since he had never seen the Highlands tiger beetle before we made plans to slip away one day and visit the spot where I had seen them back in 2009. I knew our chances of finding them were slim—it was very late in the season (late September), and the species is a so-called “summer species” with peak of adult activity in July and August. We figured, however, that even if we didn’t find adults we would still enjoy the day in the field, and for some time after arriving at the site that’s all it was. Finally, in an open sandy area near a small lake we saw the first adult. I let Chris take his shots, as this was his first opportunity (see photo above), while I continued to search for additional adults. Eventually I found one and began the long process of “whispering” to it to coax it into allowing me the photographs I desired.

Cicindelidia highlandensis

Highlands tiger beetle (Cicindelidia highlandensis).

As you can see in the photograph above, my compositional preferences have changed since I took those first photographs back in 2009. In contrast to the “as close as possible” style that I initially adopted, I now prefer to back off from the beetle enough to include elements of the habitat in which it occurs. While this compositional style may show less detail on the beetle itself, I believe it adds perspective and results in a more interesting and aesthetically pleasing photograph. I also now like to get down as low as I can, often placing the camera directly on the ground rather than always shooting from “elbow-height”, for a more unique perspective of the beetle, and my use of better flash diffusion results in more even lighting and minimizes the distracting specular highlights that are often the hallmark of flash macrophotography.

Cicindelidia highlandensis

This individual demonstrates the thermoregulatory “stilting” behavior of the adults.

Sadly, my flash unit failed soon after I began photographing the beetle, which is a real shame because the beetle began demonstrating the characteristic “stilting” behavior that the adults use for thermoregulation in their hot environment. The photograph above is the only one that I could “rescue” through some rather heavy-handed post-processing to make up for the failure of the flash unit to fire (it is fortunate that I have shifted to routinely using a combination of ambient light and fill-flash rather than flash only, or I would have had not even this photograph to rescue!). I suppose this means I’ll just have to revisit this species once again (now that I have not one but two new flash units!), which isn’t all bad because I would also love to see and photograph once again the moustached tiger beetle (Ellipsoptera hirtilabris), another Florida endemic (or near so) that I saw here also during my first visit but not during this one.

The Highlands tiger beetle belongs to a group of species called the abdominalis species-group, with all four of the included species (C. abdominalis, C. floridana, C. highlandensis, C. scabrosa) occurring in Florida (three of which are endemic or near-endemic to Florida). For those interested, I have seen and photographed all four of the species and presented a “mini-review” with photographs and links to posts with more detailed information about each species, along with a key to the species to allow for their identification.

REFERENCES:

Choate, P. M., Jr. 2003. A Field Guide and Identification Manual for Florida and Eastern U.S. Tiger Beetles.  University Press of Florida, Gainesville, 224 pp.

© Ted C. MacRae 2019

Mother and daughter (perhaps)

Phrynus marginenotata (Florida tailless whip scorpion)

Back in May I visited the California Department of Food and Agriculture’s Plant Pest Diagnostics Laboratory in Sacramento. While I was there to visit my friend and colleague Chuck Bellamy and see him receive Honorary Membership in The Coleopterists Society, I was also anxious for the opportunity to spend time with the lab’s other entomologists—most of whom I interact with as members of the Editorial Board of The Pan-Pacific Entomologist. Among the more ‘colorful’ of these is Martin Hauser, a dipterist (although I don’t hold that against him!) who also has a passion for maintaining live, captive arthropods. For me, there is nothing finer than visiting the lab/office of a taxonomic entomologist—one wall lined with steel cabinets full of insect specimens, another wall crammed-full of books and literature (the older the better), a workbench with microscope at the center of a jumble of specimen containers and open reprints, and shipping boxes piled everywhere. I take that back—there is nothing finer than visiting the lab/office of a taxonomic entomologist that also keeps livestock! Martin’s collection of live arthropods, however, goes well beyond the requisite 10-gallon aquarium with Madagascan hissing cockroaches. I already featured one of his more unusual tenants, Damon diadema (Tanzanian giant tailless whip scorpion), and here I feature its North American relative, Phrynus marginenotata (Florida tailless whip scorpion).

The genus is characterized by five spines on the pedipalp tibia—the 3rd shorter than the 2nd and 4th.

According to Weygoldt (2000), this is the northernmost and only U.S. representative of a mostly northern Neotropical genus of eleven species, recognized by the pedipalp tibia (the thickened segment of their “claws”) with five spines—the middle one shorter than the 2nd and 4th (refer to Photo 2). This species occurs in southern Florida and some Antillean islands, where it lives under coral stones and rocks close to the beach—a habitat that presumably subjects them to periodic flooding. While most tailless whip scorpions prefer humid/moist environments, they nevertheless studiously avoid standing water itself. This species, however, has been observed to voluntarily enter the water when placed on a stone surrounded by water and remain submerged for as many as eight hours. Remarkably, submerged individuals remain active and do not drown, apparently the result of a “plastron”—an area of the cuticle surrounding the lung openings that is packed with stiff, branched structures and, thus, capable of holding a volume of air against the body while the animal is submerged. The plastron seems to function much like a gill—oxygen continuously diffuses into the plastron from the surrounding water as it is used for respiration.

The bright orange pedipalps of 2nd-instar nymphs contrast with the somber coloration of the adults.

Like the D. diadema individuals that I also photographed, these P. marginenotata individuals had also produced viable eggs which had hatched a few weeks before my visit. Two nymphs can be seen with the adult in the top photo (although I can’t say for sure whether the adult is actually the mother), and Photos 3 and 4 show one of these nymphs up close and personal. I would have liked to have seen these nymphs when they first hatched, as the 1st-instars are a soft sea-green color and remain clustered on their mother’s abdomen until they are able to start fending for themselves (for a beautiful photo showing this, see Piotr Naskrecki’s The scariest animal that will never hurt you). The 2nd-instars that I photographed had already left their mother, and while they had lost their sea-green coloration, their pale yellow/gray bodies and bright orange pedipalps were no less striking compared to the more somber coloration of the full-sized adults.

…and, of course, the signature BitB face shot!

REFERENCE:

Weygoldt, P. 2000. Whip Spiders (Chelicerata: Amblypygi): Their Biology, Morphology and Systematics. Apollo Books, Stenstrup, Denmark, 163 pp.

Copyright © Ted C. MacRae 2012

Where siblings mingle: Ellipsoptera marginata vs. E. hamata

When Erwin & Pearson (2008) formally broke up the great genus Cicindela by elevating most of its former subgenera to full genus rank, it caused a bit of consternation amongst some North American cicindelophiles. The argument went something like, “Now we have all these new genus names to learn, and we’ll have to relabel and reorganize everything in our collections, and how do we know the names won’t change again, and we can’t even tell them apart in the field anyway, and blah blah…” Pardon me, but since when did taxonomy become more about slotting species into fixed, easy-to-learn categories and less about best reflecting dynamic knowledge of complex evolutionary relationships? In the case of Cicindela and its former subgenera, however, even these arguments don’t hold up to close scrutiny—tiger beetle enthusiasts in North America should have already been quite familiar with the former subgenera due to their inclusion in the widely accepted Pearson et al. (2006) field guide, many of which actually do present a unique suite of morphological/ecological characters that facilitate their recognition in the field, and I personally find that nomenclatural recognition of individual lineages helps my attempts to learn and understand them much more than dumping them into a large, all-encompassing genus based on superficial resemblance. As for insisting that names don’t change, well that has never been a tenet of taxonomy. Stable, yes, but fixed and immutable, no.

Ellipsoptera marginata male | Pinellas Co., Florida

Enough waxing philosophic. One of the more distinctive of the former subgenera is Ellipsoptera. Morphologically the genus is defined by details of male genitalia, but the 11 North American species are generally recognizable in the field by their relatively “bug-eyed” look and long legs (Pearson et al. 2006) and, as a group, seem ecologically tied to extreme habitats with sandy and/or saline substrates that are nearly or completely devoid of vegetation. Coastal marshes and mudflats, saline flats, sandy river banks, and deep sand ridges representing ancient coastlines are some of the habitats where species in this genus are most commonly encountered. Most of the species exhibit a fairly uniform facies but differ in the details of maculation and dorsal coloration, but two species that stand apart from the rest are E. marginata (Margined Tiger Beetle) and E. hamata (Coastal Tiger Beetle) due to the highly diffuse middle band of their elytra. These are both eastern coastal species and presumably represent sibling species that have diverged based on geographical range partitioning—E. marginata along the Atlantic Coast and E. hamata along the Gulf Coast. In the field, the two species are almost identical in appearance but nonetheless easily identifiable based on geographical occurrence. There is, however, a small stretch of coastline—the lower Gulf Coast of Florida—where the ranges of the two species overlap and geography alone isn’t sufficient for species determination.

Ellipsoptera hamata lacerata male | Dixie Co., Florida

Fortunately, despite their strong resemblance to each other, field identifications in areas where these species co-occur are still possible due to the presence of small but distinct sexual characters present in one species but absent in the other. Close examination is necessary to see the characters (or their absence), so it is best to net a few individuals and examine them in the hand or, as I have done here, look at them through the viewfinder of a camera. The photos in this post include the male and the female of both species, each showing the presence or absence of the distinguishing character.

Ellipsoptera marginata female | Dixie Co., Florida

In most tiger beetles, male individuals are distinguished by a number of secondary sex characters, but easiest to see in the field are the brush-like pads on the underside of the front tarsi (“feet”). Males of E. marginata and E. hamata are further distinguished from each other by the presence (E. marginata) or absence (E. hamata) of a distinct tooth on the underside of the right mandible. Photo 1 above shows a male E. marginata from Pinellas Co., Florida, and the tooth is easily seen in that relatively distant view. Photo 2 above shows a male E. hamata lacerata (Gulf Coast Tiger Beetle)—the Floridian subspecies, and while a small bump can be seen on the underside of the right mandible, it is not nearly as well developed into a distinct tooth as in E. marginata.

Ellipsoptera hamata lacerata female | Dixie Co., Florida

Female tiger beetles, on the other hand, lack the brush-like tarsal pads present in the males and are further distinguished by the “mesopleural coupling sulcus”—an area just behind the side of the pronotum that receives the male mandible during mating and is thus devoid of setae (compare the females in Photos 3 and 4 with the males in Photos 1 and 2). Neither E. marginata nor E. hamata females possess the mandibular tooth found in E. marginata males, but they can be distinguished from each other by their elytral apices. In E. marginata females (Photo 3), the elytra are curiously “bent” at the tips, forming a distinct indentation at the apex of the elytra where they meet, while female E. hamata (Photo 4) lack this indentation.

Are there other tiger beetle sibling species groups for which you would like to see comparative posts such as this one?

p.s. I completely neglected to mark yesterday’s 5th anniversary of Beetles in the Bush! I don’t know how I missed a milestone as big as five years—hopefully my ability to provide interesting content is faring better than my middle-aged memory!

REFERENCES:

Erwin, T. L. and D. L. Pearson. 2008. A Treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of life. Volume II (Carabidae-Nebriiformes 2-Cicindelitae). Pensoft Series Faunistica 84. Pensoft Publishers, Sofia, 400 pp.

Pearson, D. L., C. B. Knisley and C. J. Kazilek. 2006. A Field Guide to the Tiger Beetles of the United States and Canada. Oxford University Press, New York, 227 pp.

Copyright © Ted C. MacRae 2012

The Third of Florida’s Three Metallic Tiger Beetles

Tetracha virginica (Virginia Metallic Tiger Beetle) | Levy Co., Florida

After three straight posts not about tiger beetles, I’m hoping readers will forgive my return to this fascinating group. The photos in this post represent Tetracha virginica (Virginia Metallic Tiger Beetle), the most widely distributed (at least in the U.S.) of the four species occurring in North America north of Mexico. Even though this species occurs in my home state of Missouri, I’d not found an opportunity to photograph it until August last year at Florida’s “Road to Nowhere“—famous among U.S. cicindelophiles as one of the country’s true tiger beetle “hot spots.” In fact, it was on the very same night at this same place that I photographed the related Tetracha carolina (Carolina Metallic Tiger Beetle) (featured in Not all Florida tiger beetles are rare) and just one day after I photographed the endemic Tetracha floridana (Florida Metallic Tiger Beetle) (featured in Why I Roamed the Marsh at Night). That’s all three species of Tetracha occurring in Florida in just two days (and if I want to photograph the fourth and only remaining U.S. species, Tetracha impressus (Upland Metallic Tiger Beetle), I’ll have to go to Brownsville, Texas and get very lucky!).

The solid green elytra without apical markings distinguish this species from all other Tetracha spp. in the U.S.

Truthfully, I had no plans to post these photos after I took them. Like the other species they were photographed at night, and when I got a better look at the photos on the computer I was disappointed to see the subject was badly covered with large particles of sand. I don’t mind a little bit of debris on insects—it is, after all, a normal part of their appearance. However, too much debris is, for me, an aesthetics killer! “Wait a minute… these don’t look too bad”, you say? Well, thanks to the Clone Stamp Tool in Photoshop Elements, and as a followup to my recent post on this subject, I now have enough confidence to tackle not only small pieces of debris, but also more difficult “debris cases” such as this one with relatively large particles. Here is the same photo as shown above and processed in exactly the same manner, except that no cloning was used to remove the debris:

Aren’t I a dirty boy?!

Obviously, there are limits to what the Clone Stamp Tool can do, and I didn’t try to deal with the sand particles clinging to more difficult to clone body parts such as legs and antennae (although I’m sure that in the right hands even these could be cloned out). Nevertheless, even just cleaning the dorsal surface of the beetle does much to improve its appearance with a relatively minor amount of effort.

And, of course, what would a tiger beetle post be if it did not end with my signature face portrait (notwithstanding a few large sand grains that I wasn’t sure I could clone out effectively)?

Copyright © Ted C. MacRae 2012

Sexual Profiles

I recently happened upon these photographs of Habroscelimorpha severa (Saltmarsh Tiger Beetle), taken in early August last year at the terminus of Florida’s famous “Road to Nowhere“. I hadn’t thought to post them afterwards because I’d already shown a photograph of this species taken at the same spot during the previous year’s visit. I should have, as they are much better photographs than that initial attempt. Blame part of the first attempt on the fact that I was only in my third month of insect macrophotography, but the biggest reason for the improvement was because I’d gotten a little smarter and learned to use a blacklight to bring these extremely wary beetles to me at night rather than try to chase after them during the day. Still, I don’t get much enjoyment out of posting photos for no other reason than to post photos, so they’ve sat on my hard drive for the past year and a half. In looking at them again, however, I realized that the first and second, profiles of a female and a male, respectively, make for a nice comparison of the secondary sexual characters exhibited by adults of this species.

Habroscelimorpha severa (female) | Levy Co., Florida

Females exhibit fewer sexual characters than males, the main one being the presence of grooves on each side at the back of the pronotum (neck). These grooves function during mating, at which time the male grasps the female by the pronotum with his mandibles. This helps to provide a more secure grip for the male to prevent him from being dislodged during mating and subsequent mate guarding. The grooves themselves are not obvious in the photo, but the lack of setae (hairs) within them is, giving the female a less “hairy” look than the male. As with most insects, females also are more robust—their abdomens larger to make room for egg-making machinery, although in this and other tiger beetle species the difference is not that obvious.

Image

Habroscelimorpha severa (male) | Levy Co., Florida

Males are immediately recognizable by several respects. In addition to the smaller abdomen and “hairier” pronotum lacking lateral grooves, male tiger beetles in most of the “higher” genera exhibit brush-like pads on the undersides of the front tarsi (feet). The function of these pads is not completely clear, but prevailing opinion is that they somehow aid in gripping the female during mating. I’m not sure I buy into this—males do sometimes hold onto females with their front legs during mating, but how these pads improve grip escapes me. Further, it is my experience that males actually spend more time during mating and mate guarding with their front legs outstretched to each side. I’ve also noticed that males are reluctant to release females even when danger approaches (even in the form of a giant insect macrophotographer). I’ve seen males tenaciously clinging to the female as she violently tries to shake him off and flee from my approach. It makes me think that perhaps the tarsal pads serve some tactile function as a final warning of impending danger to a grasping male, allowing him to not give up his female until absolutely necessary (hey, it’s an idea—if you have an alternative idea I’d love to hear it). There is more, however—look at that big head!

Habroscelimorpha severa (male) | Levy Co., Florida

Actually,  the male’s head is no larger than the female’s, but the proportionately longer mandibles give the male a distinctly “big-headed” look.  In contrast, the labrum (upper lip) is shorter than the female’s (making the mandibles look longer still). Both of these characters are, again, related to the habit of grasping the female pronotum, with the longer mandibles allowing a more secure grip of the female pronotum and the shorter labrum adding even more functional length to the mandibles (I can also imagine that this might have some effect on choice of prey by males versus females). The male mandibles also have a greater amount of white coloration at their bases—this might simply be a function of the relatively larger size of the mandibles, but given that males of many species exhibit more white overall on both the mandibles and the labrum (the latter of which is usually smaller), it seems more logical to me that the white coloration serves as a visual cue for potential mate recognition.

“Hey baby, I like your grooves!”

“Yeah, well your white lips aren’t so bad either.”

Copyright © Ted C. MacRae 2012