Tiger Beetles Agree—It’s Hot in Florida!

Florida is known for its rich assemblage of tiger beetles—27 species in all, including four endemics (Choate 2003).  However, late summer is generally considered not the best time of year for seeing this diversity, since adult populations of most species begin to wane as the intensity of the summer heat reaches its peak.  I knew the timing of my family vacation in early August might be a bit off; however, considering I had never looked for tiger beetles in Florida before, I remained optimistic that I still might encounter some interesting species.  My optimism was quickly rewarded—in one afternoon of exploring the small coastal preserve just outside the back door of my sister-in-law’s condo, I found Ellipsoptera marginata (Margined Tiger Beetle), its sibling species E. hamata lacerata (Gulf Beach Tiger Beetle), and several 3rd-instar larvae in their burrows that proved to be the Florida endemic Tetracha floridana (Florida Metallic Tiger Beetle).  Good fortune would continue when I made a one-day trip to the interior highlands in a successful bid to find Florida’s rarest endemic, Cicindela highlandensis (Highlands Tiger Beetle), finding also as a bonus the splendidly camouflaged and also endemic Ellipsoptera hirtilabris (Moustached Tiger Beetle).  Five species, including three endemics, in just over a day of searching!  I had one more day to sneak off and do what I love most, and I wanted to make the most of it. 

Pine sandhill habitat, Withlachoochee State Forest—Citrus Tract

Among the suggestions given to me by my colleagues, the most promising-sounding was the “end of the road,” a Gulf Coast salt marsh near Steinhatchee in Dixie County where I was told as many as 6-10 species of tiger beetles could be seen at once.  I didn’t know it at the time, but this particular location has achieved legendary status among tiger beetle enthusiasts (Doug Taron recently wrote about his experience, calling it the Road to Nowhere).  A 200+ mile drive from my base near St. Petersburg, it would take the better part of 5 hours to drive there, and not wanting to put all of my eggs in one basket, I looked for potential stops along the way.  About midway along the drive was Withlacoochee State Forest, where one of my colleagues had told me I might still find the fairly widespread Cicindela abdominalis (Eastern Pinebarrens Tiger Beetle) and its close relative, C. scabrosa (Scabrous Tiger Beetle)—the fourth Florida endemic.  My plan was to leave early in the morning and spend a few hours at Withlacoochee before driving the rest of the way to finish out the day at Steinhatchee. 

"Stilting" by Cicindela abdominalis (Eastern Pinebarrens Tiger Beetle)

It took some time to find my bearings upon arriving, but after some discussion with the decidedly forestry-oriented staff at the headquarters, it seemed that the Citrus Tract was where I wanted to be.  I was looking for the sand barren and pine sandhill habitats that these species require, and the staff’s description of the northern edge of the tract as having lots of sand and “not very good for growing trees” suggested this might be the place.  Pine sandhill (also called “high pine”) is a pyrophytic (fire-dependent) plant community characterized by sandy, well-drained soils, a widely-spaced longleaf pine (Pinus palustris) and turkey oak (Quercus laevis) canopy, and an herbaceous layer dominated by wiregrass (Aristida stricta).  I quickly found such habitat in the area suggested, and it wasn’t long before I found the first of the two species—C. abdominalis—rather commonly along a sandy 2-track leading through the area.  For those of you who see a distinct resemblance of this species to the rare C. highlandensis that I highlighted from my trip to the central highlands, this is no coincidence.  Cicindela abdominalis is very closely related to that species, the latter distinquished by an absence of flattened, white setae on the sides of the prothorax and the abdomen and by the highly reduced or absent elytral maculations (Choate 1984).  Dense white setae and distinct apical elytral maculations are clearly visible in the individuals shown in these photographs. 

Stilting is often accompanied by "sun-facing" for additional thermoregulation

It was a blistering hot day (just as every other day on the trip had been so far), and it wasn’t only me who felt that way.  Tiger beetles, of course, are ectothermic and rely upon their environment for their body temperature.  Despite this, they are able to regulate body temperatures to some degree by using a range of behavioral adaptations intended to mitigate the effects of high surface temperatures and intense sunlight.  The photos above show one of these behaviors, known as stilting.  In this behavior, the adult stands tall on its long legs to elevate its body above the thin layer of hotter air right next to the soil surface and as far off the sand as possible (Pearson et al. 2006).  As the heat of the day intensifies and the zone of hot air at the soil surface broadens, stilting alone may be insufficient to prevent overheating. When this happens, the beetles combine stilting with sun-facing, a behavior in which the front part of the body is elevated with the head oriented towards the sun. This position exposes only the front of the head to the sun’s direct rays, thus minimizing the body surface area exposed to incident radiation.

Stilting and sun-facing by Ellipsoptera hirtilabris (Moustached Tiger Beetle)

I was also fortunate to have another chance at photographing the beautiful and marvelously-camouflaged Ellipsoptera hirtilabris (Moustached Tiger Beetle), which, in similar fashion to C. highlandensis, I found co-occurring with C. abdominalis in rather low numbers. As before, they were extremely wary and difficult to approach, especially in the extreme heat of the day, and all of my best efforts to get a good shot of the species in its “classic” pose were frustrated. The photo above was about as close as I could get to any of these beetles when they were out in the open before they would flee; however, it nicely demonstrates the use of stilting combined with sun-facing during the hottest part of the day.

"Shade seeking" is another behavioral response to intense heat.

Another behavioral response to extreme heat is shade-seeking—adults may either remain active, shuttling in and out of shaded areas, or avoid exposed areas altogether and become inactive.  One thermoregulatory behavior for extreme heat that I did not observe was daytime-burrowing, in which adults construct temporary shallow burrows during the hottest hours of the day. Although I did not observe this behavior by either species at Withlacoochee, I have seen it commonly among several species in sandy habitats here in Missouri and in the Sandhills of Nebraska (e.g., Cicindela formosa, Cicindela limbata, Cicindela repanda, Cicindela scutellaris, Cicindela tranquebarica, Ellipsoptera lepida).

There was one disappointment on the day—I did not see C. scabrosa.  However, I still had the “end of the road” to explore, so I remained happy with the now six species I had encountered and optimistic about finding additional species later in the day… 

Photo Details: Canon EOS 50D, ISO 100.
Habitat: Canon 17-85mm zoom lens (landscape, 17mm), 1/100 sec, f/10, natural light.
Insects: Canon 100mm macro lens (manual), 1/250 sec, f/16–18 (C. abdominalis) or f/20–22 (E. hirtilabris), MT-24EX flash w/ Sto-Fen diffusers.

REFERENCES: 

Choate, P. M., Jr.  1984.  A new species of Cicindela Linnaeus (Coleoptera: Cicindelidae) from Florida, and elevation of C. abdominalis scabrosa Shaupp to species level.  Entomological News 95:73–82.

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.

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 2009

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Florida Scrub Lizard

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The Florida scrub lizard (Sceloporus woodi) is restricted to isolated sand scrub habitats in peninsular Florida.

Tiger beetles were not the only rare endemic species that I encountered during my visit to the Lake Wales Ridge in central Florida last August.  I didn’t know what this small lizard was as I watched it bolt from the trail and scamper for cover during my approach; however, having already found two endemic tiger beetles, I had a feeling that this lizard might also be a good one.  The photo shown here is admittedly not one of my best, but it was the only one I managed to get before the lizard ducked into the brush for good.  Horribly overexposed, I did what I could with it in Photoshop to make it halfway presentable, but there is no question that its subject represents a Florida scrub lizard, Sceloporus woodi¹.  This small, diurnal, ground-dwelling lizard belongs to the family Phrynosomatidae (same family as the Texas horned lizard that I featured in this post) and is restricted to Florida’s rare sand scrub and sandhill habitats.  Like the recently featured Highlands Tiger Beetle, this species is threatened by the isolated, disjunct nature of its required habitat—a threat made worse by the ever increasing pressures of agricultural conversion and urban development.

¹ Sceloporus is derived from the Greek word scelos meaning “leg” and the Latin word porus meaning “hole”, referring to the pronounced femoral pores found in this genus of lizards. The species epithet honors Nelson R. Wood, a taxidermist at the U.S. National Museum who collected the type specimen in 1912.

Distribution of the Florida scrub lizard (from Branch et al. 2003).

The Florida scrub lizard is related to and closely resembles the much more common and widely distributed southern fence lizard (Sceloporus undatus), which co-occurs with the scrub lizard in northern Florida.  Fence lizards, however, lack the dark brown lateral stripe that is clearly visible in the above photo, a feature seen in juveniles and adults of both sexes of the scrub lizard.  Juvenile and adult female scrub lizards also exhibit a dorsal zigzag pattern; however, this fades in males as they reach adulthood and develop the characteristic bright blue belly patches that are seen in both this species and in the fence lizard (Branch and Hokit 2000).  Since light blue patches are just visible on the belly and throat of the individual in the photograph, I haven’t been able to determine whether it represents a mature female or a still-juvenile male—any help from a knowledgeable reader would be greatly appreciated.  Unlike the fence lizard, the scrub lizard displays a high degree of habitat specificity, occurring as disjunct populations in strict association with the major sand scrub ridges of Florida.  The healthiest populations are found on the Mt. Dora Ridge in northern peninsular Florida, on which significant remnants of scrub habitat are preserved in the Ocala National Forest.  Populations also occur on the Lake Wales Ridge of central Florida and the Atlantic Coastal Ridge, but the status of these populations is less secure.  Populations also once occurred along the southwestern coast on the Gulf Coast Ridge, but these populations are now believed extirpated as a result of urban development (Jackson 1973, Enge et al. 1986).  While the Florida scrub lizard is not listed as a threatened or endangered species at the state or federal level, its high specificity to an increasingly isolated and fragmented habitat and its apparently low dispersal capabilities are clear causes for concern over its long-term prospects. As remnant habitats continue to shrink and become more isolated, the threat of localized extinction becomes an increasing concern for the lizard populations that they support.

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Scrub lizard habitat is threatened by development, fragmentation and increased vegetation.

The precarious status of scrub lizards and their occurrence in several disjunct, isolated populations makes them interesting subjects for genetic studies. Mitochondrial DNA analyses suggest that scrub lizard populations exhibit a high degree of phylogeographical structure, with populations diverging significantly not only between major scrub ridges, but also within them (Branch et al. 2003).  The findings support the notion of long-term isolation of scrub lizard populations on the major scrub ridges and confirm their low dispersal rates among adjacent scrub habitats within ridges (as little as a few hundred yards of “hostile” habitat may be sufficient to prevent movement to adjacent habitats).  More significantly, the results support the concept of two distinct morphotypes on the Mt. Dora and Lake Wales Ridges and also raise the possibility that Atlantic Coastal Ridge populations represent a distinct evolutionary entity as well.  These findings are consistent with the hypothesis that scrub lizards evolved in central Florida, where they were isolated when surrounding lands were inundated by rising sea levels during the late Pliocene and subsequent interglacial periods during the Pleistocene.  During periods of low sea level they dispersed to the younger Atlantic and Gulf Coastal Ridges, where they were isolated from parent populations when more mesic conditions returned during the Holocene (12 kya to present).  The genetic distinctiveness of these different ridge populations may justify qualifying each of them for protection as “significant evolutionary units” under the U.S. Endangered Species Act, since it raises concerns about the use of translocations, a common strategy for establishing new populations in restored habitat or augmenting existing populations, as a conservation strategy for the species as a whole.  Since lizards located on different ridges are more divergent than lizards from populations located on the same ridge, movement of lizards between ridges could compromise the integrity of the genetic differences that have accumulated over millions of years and result in loss of genetic diversity.  As a result, augmenting populations on the Lake Wales and Atlantic Coast Ridges with lizards from robust populations on the Mt. Dora Ridge may not be desirable.  Instead, it may be necessary to protect individual scrub lizard populations on each of the major scrub ridges in order to preserve as much of their genetic diversity as possible.

REFERENCES:

Branch, L. C. and D. G. Hokit. 2000. Florida scrub lizard (Sceloporus woodi). University of Florida, IFAS Extension Service Publication #WEC 139, 3 pp.

Branch, L. C., A.-M. Clark, P. E. Moler and B. W. Bowen.  2003. Fragmented landscapes, habitat specificity, and conservation genetics of three lizards in Florida scrub.  Conservation Genetics 4:199

Enge, K. M., M. M. Bentzien, and H. F. Percival. 1986. Florida scrub lizard status survey. Technical Report No. 26, U.S. Fish and Wildlife Service, Jacksonville, Florida, U.S.A.

Jackson, J. F. 1973. Distribution and population phenetics of the Florida scrub lizard, Sceloporus woodi. Copeia 1973:746–761.

Copyright © Ted C. MacRae 2009

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Moustached Tiger Beetle

Lake Wales Ridge

Dry sand scrubland on Lakes Wales Ridge in central Florida—home to Cicindela highlandensis and Ellipsoptera hirtilabris

In my previous post, I featured the rare Cicindela highlandensis (Highlands Tiger Beetle), restricted entirely to sand scrubland and pine woodland habitats along the Lake Wales Ridge in central Florida (Choate 2003).  However, that would not be the only Florida endemic tiger beetle that I would encounter during my early August visit.  Another of the several tiger beetle species that I’d hoped to see would also be found that day, although in much lower numbers.  Ellipsoptera hirtilabris (Moustached Tiger Beetle) is so named¹ because of the dense covering of prostrate hairs on its labrum that distinguish it from the closely related E. gratiosa (Whitish Tiger Beetle). Both of these species exhibit striking white maculations that cover almost the entire elytral surface and dense white pubescence covering the head, thorax, underside and legs.  They are the only species of the genus occurring in Florida, but their ranges do not overlap (Pearson et al. 2006)—E. gratiosa occurs in the coastal pine barrens of Virginia, the Carolinas, southern Georgia and the Florida panhandle, while E. hirtilabris is restricted to peninsular Florida in pine woodlands, sand hills and other habitats with open white sand.  Although the latter is considered a Florida endemic, it has been found just outside of Florida in extreme southeastern Georgia on St. Simon’s Island (Choate 2003)In addition to the pubescence of the labrum and their allopatric distributions, the two species may further be distinguished by the slightly less expanded markings and more diffuse edges where they contact the central bronze area in E. hirtilabris and the slightly larger size of E. gratiosa.    Like C. highlandensis and C. abdominalis, it seems likely that E. hirtilabris and E. gratiosa evolved from a common ancestor, diverging in isolation from each other during the pre-Pleistocene separation of peninsular Florida from the North American mainland.   

¹ The species epithet is derived from the Latin words hirtum meaning “hairy” and labrum meaning “lip”.

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Ellipsoptera hirtilabris in the alert position

I found E. hirtilabris to be exceedingly difficult to see and photograph.  Unlike C. highlandensis, which resemble bits of debris laying on the surface of the white sands where it lives, the largely white E. hirtilabris blend into the white sand itself and are almost impossible to see until they move.  The small bronze-colored patches along the elytral suture augment their cryptic capabilities by resembling small bits of debris, which is especially evident in the photo below.  Both Pearson et al. (2006) and Erwin and Pearson (2008) state that adults of this species freeze in position when approached, which may be the reason why I saw so few individuals.  Once I did see them, they were extremely wary and difficult to photograph no matter how cautiously I approached.  The photos shown here represent the only two individuals that I succeeded in photographing, and in neither case did I succeed in getting a frontal perspective to show the pubescent labrum (stifling heat and oppressive humidity during the photo session did not help matters, either).

Ellipsoptera_hirtilabris_IMG_1131_1200x800_enh

The white coloration with small brown markings helps adults blend in perfectly in their white sand habitats

Photo details:
Photo 1: Canon 100mm macro lens on Canon 50D (landscape mode) ISO-100, 1/250 sec, f/16, natural light.
Photos 2 & 3: Manual mode, f/25, MT-24EX flash w/ Sto-Fen diffusers @ 1/8 ratio.

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.

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 2009

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Highlands Tiger Beetle

When my wife and I made plans to spend a week at her sister’s condominium in Florida this past summer, I began making a list of the tiger beetle species that I wanted to see.  I would be happy to see anything, since I had never before tiger beetled in Florida, but early August was looking to be on the late side for many things.  In addition, since this was a family vacation, I would only have a couple days at most to sneak off on my own and immerse myself in bug hunting.  All this meant that I would have to be very judicious about where I went and what I looked for.  I sought advice from a few other cicindelophiles on species and localities, and by the time we made the 16-hour drive from St. Louis to Seminole (near St. Petersburg) I had settled on two destinations—the Lake Wales Ridge of central Florida to look for Cicindela highlandensis (Highlands Tiger Beetle), and the so-called “Road to Nowhere” near Steinhatchee where as many as 10 species of tiger beetles can be seen when the season is right.  Things started out well when, before even looking for any of these species, I stumbled upon Ellipsoptera marginata (Margined Tiger Beetle), its sibling species E. hamata lacerata (Gulf Beach Tiger Beetle), and some 3rd-instar larvae in their burrows that proved to be the Florida endemic Tetracha floridana (Florida Metallic Tiger Beetle) in the small coastal preserve just outside the back door of my sister-in-law’s condo.

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Dry sand scrubland on Lakes Wales Ridge in central Florida

The big target of the trip, however, was not so straightforward.  Cicindela highlandensis is one of Florida’s rarest endemic tiger beetles, being restricted entirely to remnant sand scrubland and pine woodland habitats along the Lake Wales Ridge of Polk and Highlands Counties in central Florida (Choate 2003).  The Lake Wales Ridge represents former shorelines deposited when the rest of peninsular Florida was covered by seas.  The quick draining sands have created desert-like open habitats dominated by oaks, pines, and other drought-tolerant species.  Cicindela highlandensis is one of many plants and animals endemic to the Lake Wales Ridge, which has the highest concentration of endangered plants in the continental U.S.  Unfortunately, the natural communities found on the Lake Wales Ridge have suffered severe reductions from their historical occurrence.  An estimated 85% of the scrub and sandhills has been converted to citrus groves and urban developments, and the few remaining tracts face not only continued development pressure, but also the threat of degradation from reductions in the frequency and extent of the wildfires that are essential for their maintenance (Turner et al. 2006).  NatureServe (2009) estimates that C. highlandensis populations have declined by as much as 90%, and only a few of the sites where it is known to occur are large enough to sustain viable populations.  While the species has a global status of G1 (critically imperiled) due to its limited range, restricted habitat, and very small population size, and is a candidate for listing as an endangered species by the U.S. Fish & Wildlife Service (Pearson et al. 2006), it remains—bafflingly—unlisted even as threatened by the State of Florida.  As a result, there is no formal conservation management plan for this species to ensure its survival.  Fortunately, the largest populations of C. highlandensis occur on an assemblage of public and private lands that are under partial to full conservation ownership, and preservation/management activities are taking place at most of these.

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Cicindela highlandensis, Highlands Tiger Beetle, in alert position

I had debated whether to look for Cicindela highlandensis at all—not because I wasn’t anxious to see it, but because I lacked confidence that I would be able find it.  A late season search for a rare species had all the hallmarks of a potential wild goose chase.  Nevertheless, I like a good challenge, and I had succeeded in obtaining information about specific locations for the species (a matter of public record; however, I prefer to maintain some discretion in this venue).  Although I began my search with tempered optimism, it didn’t take long for me to acheive my goal.  Similar to my experience with Cylindera celeripes in Oklahoma, I had barely walked ten yards into a gorgeous sand scrub habitat at the first site I had planned to search before I saw an individual.  The dark metallic blue coloration of the species would seem to make it easily seen in its white sand environs; however, in reality it is almost impossible to see until it moves.  Some have suggested that its coloration functions to make the beetle resemble the many small pieces of debris that litter the sand surface—perhaps the bits of charred wood that are common in open, fire-mediated environments.  Its dependence upon natural disturbance factors such as fire was made apparent to me by the distinct preference I noted for adults to congregate along trails kept open by human disturbance, and to a lesser degree in the larger, naturally open scrub areas.  The adults made very short escape flights and were easy to follow but difficult to approach closely enough for photographs due to extreme wariness—their long legs giving some indication of their highly cursorial capabilities.

Cicindela highlandensis

Cicindela highlandensis - note absence of setae on thorax and abdomen

Cicindela highlandensis is closely related to two other species of tiger beetles in Florida—C. abdominalis (Eastern Pinebarrens Tiger Beetle), widely distributed throughout the Atlantic and Gulf Coastal Plain, and C. scabrosa (Scabrous Tiger Beetle), confined to the Florida Peninsula and adjacent southeastern Georgia.  Both of these species are absent from the Lake Wales Ridge and, thus, do not co-occur with C. highlandensis. It is likely that C. highlandensis evolved from isolated populations of the widespread C. abdominalis that diverged during pre-Pleistocene separation of the Lake Wales Ridge from the mainland (Choate 1984).  Despite its resemblance to both C. abdominalis and C. scabrosa, C. highlandensis can be distinguished from both of those species by the complete absence of flattened, white setae on the sides of the prothorax and the abdomen and by the highly reduced or absent elytral maculations (note the very small apical markings on the individuals in these photographs).  All three of these species belong to the subgenus Cicindelidia (American Tiger Beetles) and possess red adominal coloration that is prominent during flight.

Photo details:
Photo 1: Canon 17-85mm zoom lens on Canon 50D (landscape mode), ISO-100, 1/160 sec, f/13, natural light.
Photos 2–3: Canon 100mm macro lens on Canon 50D (manual mode), ISO-100, 1/250 sec, f/14 (photo 2) or f/20 (photo 3), MT-24EX flash w/ Sto-Fen diffusers.

REFERENCES:

Choate, P. M., Jr.  1984.  A new species of Cicindela Linnaeus (Coleoptera: Cicindelidae) from Florida, and elevation of C. abdominalis scabrosa Shaupp to species level.  Entomological News 95:73–82.

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.

NatureServe.  2009.  NatureServe Explorer: An online encyclopedia of life [web application].  Version 7.1.  NatureServe, Arlington, Virginia.  Available at: http://www.natureserve.org/explorer (accessed: November 12, 2009).

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.

Turner, W. R., D. S. Wilcove and H. M. Swain.  2006.  State of the scrub: conservation progress, management responsibilities, and land acquisition priorities for imperiled species of Florida’s Lake Wales Ridge.  Archbold Biological Station, Lake Placid, Florida, iii + 44 pp.

Copyright © Ted C. MacRae 2009

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Monday Moth: White-tipped Black Moth

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Photo details: Canon 100mm macro lens on Canon 50D (manual mode), ISO-100, 1/250 sec, f/22, MT-24EX flash w/ Sto-Fen diffusers.

When is a ctenuchid moth not a ctenuchid moth?  When it’s a White-tipped Black Moth (Melanchroia chephise) in the family Geometridae!

I may be a beetle guy, but I also consider myself a competent general entomologist.  What is a competent general entomologist?  Someone who can identify any insect to order at first glance and a majority of them to family – regardless of one’s own taxa of expertise.  Thus, when I encountered this mating pair of moths on the outside wall of my sister-in-law’s condominium in Seminole, Florida, I “recognized” them as something in what I learned as the family Ctenuchidae (later subsumed within the Arctiidae, first as a subfamily and now as several disparate tribes).  They had all the hallmarks of ctenuchids—black and red coloration, narrowish wings with light colored patches, and about the size of the wasps that they presumably mimic.  Upon my return to St. Louis, I sat down to identify the moths—confident that their distinctive appearance would lead to the quick ID that never materialized after scanning through all of the ctenuchine pages at BugGuide.  Frustrated, I resorted to posting the photo on the site’s ID Request, never questioning my ctenuchine placement.  Precisely 4 minutes later, the moths were identified by John Maxwell as Melanchroia chephise and moved to their proper place—among the 50 other adult photographs of this species that can be found on the site!  I might as well have failed to identify a monarch butterfly!

Melanchroia chephise is apparently common in the American tropics, reaching its northern distributional limit along the coastal plains of Florida and Texas but straying further north in certain years.  Larvae feed on several plants in the family Euphorbiaceae, primarily Breynia and Phyllanthus species.  The adult coloration strikes me as obviously aposematic (warning coloration), but I could find no specific references to this.  However, considering that euphorbiaceous plants are famous for their diverse arsenal of latex and irritant toxins (e.g., diterpene esters, alkaloids, glycosides, ricin-type protein toxins, etc.), it seems reasonable to presume that Melanchroia larvae have evolved mechanisms for sequestering one or more of these compounds.  NABA South Texas states that adults of this species are probably mimics of the Red-bordered Pixie (Melanis pixe), an aposematic metalmark butterfly also of Neotropical distribution that reaches south Texas (but not Florida).  Personally, I don’t really see the resemblance (but then, nor am I an avian predator).  I suppose it’s possible that a species such as this can employ different defense strategies in different parts of its range, relying on Batesian mimicry in areas where suitable models occur and aposematism in areas where they don’t, but I have to admit that I’m now straying well outside the coleopteran-centric bounds of my expertise.

Copyright © Ted C. MacRae 2009

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Florida Metallic Tiger Beetle

Tetracha floridana

Tetracha floridana (Florida Metallic Tiger Beetle), dorsal view

In my previous post, I showed some photographs of the larva of an undetermined species of tiger beetle that I collected from its burrow in dry ground adjacent to a coastal salt marsh near St. Petersburg, Florida. I had assumed the larva belonged to the genus Cicindela or one of its former subgenera and was suprised to learn that this assumption was incorrect when the adult emerged 2 months later. Looking back at the photos, however, I realized that the photos and the information I gave regarding its location and habitat contained all of the necessary information to identify this larva. Five points to Mike Baker, who correctly deduced the genus (Tetracha), and in fact the larva represents Tetracha floridana (Florida metallic tiger beetle).

Hump of 5th abdominal segment, showing simple, thorn-like inner and outer hooks

The hump of the 5th abdominal segment bears simple, thornlike hooks.

The larva can be placed in the genus Tetracha by virtue of its simple, thorn-like hooks (in other eastern U.S. tiger beetle genera, the outer hooks are distinctly curved).  Two other genera of Nearctic tiger beetles that do not occur in Florida also bear simple hooks—Omus (Night-stalking Tiger Beetles, occurring along the Pacific Coast) and Amblychelia (Giant Tiger Beetles, occurring in the central and southwestern U.S.); however, the former bears three rather than two pairs of hooks, and the latter has the inner and outer hooks distinctly separated from each other.

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Tetracha floridana (Florida Metallic Tiger Beetle), lateral view.

Four species of Tetracha occur in the U.S., three of which occur in Florida—T. carolina (Pan-American Big-headed Tiger Beetle), T. floridana (Florida Metallic Tiger Beetle), and T. virginica (Virginia Big-headed Tiger Beetle).  However, T. carolina is restricted in the state to the panhandle and interior of the peninsula along rivers and in disturbed sparsely vegetated areas (Choate 2006).  Of the two remaining species, T. virginica is widespread throughout the southern two-thirds of the eastern U.S. and occurs in a variety of habitats (Pearson et al. 2006), while T. floridana is restricted to salt marsh and mud flat habitats along the Gulf coast of Florida from Dixie County south to the Keys (Choate 2006).  While the widespread occurrence and generalist tendencies of T. virginica might suggest that it is the more likely choice, the locality and habitat match precisely with T. floridana.

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The anterior lobes of the apical lunules are divergent.

Tetracha floridana is very similar to T. carolina and was long considered a subspecies of that more widely distributed species until Naviaux (2007) elevated it to species rank in his revision of this large genus.  Tetracha floridana is distinguished from T. carolina by the divergent anterior lobes of the apical lunules (photo above) and the uniformly black to dark green elytra that lack any violet or coppery reflections in the anteriolateral regions (photo below) (Choate 2003).

The anteriolateral areas of the elytra lack violet or copper reflections

The anteriolateral areas of the elytra lack violet or copper reflections.

I was happy as heck when I saw the first newly emerged adult in the rearing container, as this is a true Florida endemic.  I have encountered the two other eastern U.S. species commonly under street lamps and at building lights here in Missouri—T. virginica throughout the state and T. carolina in the southeastern lowlands, where it appears to reach its northern limit of distribution.  A fourth U.S. species in the genus, T. impressa (Upland Metallic Tiger Beetle) (T. affinis” in earlier works), occurs in northern Mexico and the Lower Rio Grande Valley (LRGV) of south Texas (Erwin and Pearson 2008).  Although I have not yet encountered it on any of my many trips to the LRGV (all of which pre-date my current cicindelophily), I understand it is regularly attracted to building and street lights in Brownsville (Pearson et al. 2006).  I believe I will have to go down there again and verify this for myself someday.

Feasting on a corn rootworm larva.

Feasting on a corn rootworm larva.

The last photo in this series illustrates the unique feeding behavior of these beetles, which despite their terrifyingly toothy mandibles are strictly fluid feeders.  The long, sharp mandibular teeth function primarily in prey subdual and in slicing and shredding their tissues, while the maxillae (second pair of feeding appendages behind the mandibles) and labium (fused third pair of appendages) comprise an “oral mill” that masticates the prey and and rolls it into a bolus.  Two brush-like structures can be seen behind the mandibles in the photo above—these are part of the maxillary laciniae and apparently function in containing and shaping the bolus as it is being masticated.  While this occurs, proteolytic enzymes are extruded from the midgut and mixed with the bolus to liquify its digestible components, which are then sucked into the beetles tiny mouth by the action of a pharyngeal pump.  Like the larva, the adult beetle thus “chews” but does not swallow its prey—a manner of feeding that is not too unlike that of spiders and other arachnids (sans the venom).

Photo details:
All photos: Canon EOS 50D, manual mode, ISO-100, 1/250 sec, MT-24EX flash w/ diffuser caps.
Photo 1: Canon 100mm macro lens w/ 68mm extension tube, f/25, 1/2 power flash.
Photo 2: Canon MP-E 65mm 1–5X macro lens, f/16, 1/8 power flash.
Photos 3–5: Canon 100mm macro lens w/ 36mm extension, f/18–f/22, 1/4 power flash.
Photo 6: Canon 100mm macro lens w/ 68mm extension, f/20, 1/2 power flash.

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.

Choate, P. M., Jr.  2006.  Tiger Beetles of Florida, Cicindela spp., Megacephala spp. (Insecta: Coleoptera: Cicindelidae).  University of Florida, IFAS Extension Service Circular EENY-005, 5 pp.

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.

Naviaux R. 2007. Tetracha (Coleoptera, Cicindelidae, Megacephalina): Revision du genre et descriptions de nouveaus taxons. Mémoires de la Société entomologique de France 7:1-197.

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 2009

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Anatomy of a Tiger Beetle Larva

My first experience looking for tiger beetles in Florida had gone well.  Despite its small size and urban surroundings, the narrow strip of coastal scrub and saltwater marsh along the intracoastal waterway behind my sister-in-law’s condominium boasted a robust population of what I took to be a single tiger beetle species.  The specimens I collected and photographs I took would later reveal that two co-occurring and closely related species were present: Ellipsoptera marginata (Margined Tiger Beetle) and E. hamata lacerata (Gulf Beach Tiger Beetle).  I had spent close to two hours under the August sun observing and photographing the beetles before I decided that I had given the preserve a thorough enough look.

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As I was heading back, I noticed a little bit of high ground alongside a red mangrove thicket and went over to give it a look.  As I approached I saw something I hadn’t yet seen that day – tiger beetle larval burrows.  Larval burrows, especially larger ones such as these were, are unmistakeable – almost perfectly circular (slightly cut out on one edge) and smoothly beveled around the perimeter.  There were a number of burrows clustered on the small bit of high, dry ground, and my first thought was that their inhabitants represented the same (what I thought was a single) species that I had encountered so commonly that day as adults.  I then reasoned, however, that more likely they represented another species whose adults are active later in the season – perhaps one of the so-called “spring/fall” species whose larvae typically reach maturity during the heat of summer.  The size of the burrows (~5mm dia) suggested they were inhabited by 3rd instar larvae (the final instar before pupation), in which case it may be possible to rear a few to adulthood – if I could get at them. I tried fishing (Pearson and Vogler 2001) a few holes with a grass blade but didn’t get any bites, so I decided to watch for awhile and see if any of the larvae, believing the danger of my approach had passed, would reappear at the tops of their burrows.  Waiting for tiger beetle larvae to appear is a crap shoot – maybe they’re active, and maybe they’re not, and crouching in the stifling summer air of a coastal marsh in Florida is not an easy thing to do for very long.  Fortunately my wait was short, as within a few minutes I saw one re-appear at the top its burrow.  I slowly got out my knife and moved to place the tip on the soil about 1″ from the burrow at a 45° angle for an attempted tunnel block (Pearson and Vogler 2001), but it spooked and dropped back down into its burrow before I could get then knife in place.  No matter, I knew it was in there now and that it would likely reappear if I could muster the patience.  I positioned the knife and waited – crouched under the baking Florida sun, until when it did re-appear I plunged the knife into the soil with authority.  It was a good jab – I had blocked its retreat without injuring it, and a quick flip of the knife popped out the soil plug and exposed the startled larva, flipping vigorously in a vain attempt to escape before settling down amidst its unfamiliar, exposed surroundings.

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For those of you who have never seen a tiger beetle larva, they are among the most other-wordly creatures one can imagine.  The large, heavily sclerotized head bears two long, sickle-shaped, upward-pointing mandibles and up to three pair of highly-acute eyes whose arrangement on each side conveys the image of a “face” with congenital birth defects.  The top of the head is flattened to lie flush with the surrounding soil as the larva sits at the top of its burrow, and huge, powerful mandibular muscles fill the cranial cavity.  The remainder of the body – long, narrow, and cylindrical – hangs from the head at a 90° angle down into the burrow and is unremarkably grub-like, save for a curious hump on the dorsal side of the 5th abdominal segment.  Close examination of the hump reveals an intricate pattern of forward-facing hooks and spines that function in anchoring the larva against the side of its burrow to prevent struggling prey from dislodging it.  The life of a tiger beetle larva is a life of waiting – unlike the adults who run down their prey, the larvae sit in their burrows and wait for prey to come to within lunge’s reach.  While the eyes of most grub-like insects detect little more than light and dark, those of tiger beetle larvae are densely packed with photoreceptors that permit detailed focusing and depth perception for detecting whether potential prey has ventured close enough to their burrow (Pearson et al. 2006).  When that happens, they strike with lightning speed, plunge their mandibles into their prey, and drag it down into the depths of their burrow where it is summarily dispatched with a few bites of their powerful mandibles.  Larvae consume they prey in a manner similar to that of adults in that they chew but don’t swallow their prey. Rather, they secrete digestive secretions containing proteolytic enzymes that begin digesting the prey extra-orally as they chew.  The resulting bolus is masticated and its liquid components sucked out until nothing but a dry wad of indigestable chitin remains, which is spat out of the burrow (Pearson and Vogler 2001).

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Looking at this strange insect, it occurred to me that I had not yet attempted macrophotographs of a tiger beetle larva out of its burrow, and this would be a good opportunity to get more practice with my Canon MP-E 65 mm macro lens – a lens with incredible magnification capabilities, but one that is also a bit of a temperamental beast to use hand-held in the field.  The subject was unusually cooperative, perhaps too stunned by its sudden predicament to know what to do, and as I took the photographs I focused in particular on characters of the head and dorsal hump (often useful in identifying tiger beetle larvae, at least to genus).  Time was growing short once I finished taking photographs, so I placed the larva in a vial and returned the following day to extract a chunk of native soil to place in a rearing container, managing to collect two more larvae as well (unfortunately, one became instant “prey” for the other.  Note to self: when placing multiple tiger beetle larvae in a container of soil, seal the artificial burrows into which you place each one!).  I paid little further attention to the photographs, other than to transfer them onto my computer and add metadata upon my return to St. Louis.  I didn’t know what species the larvae represented, but I assumed they were something in the genus Cicindela or one of its several former subgenera.  However, had I studied the photos and considered the locality and habitat, I would have realized that my assumption was incorrect¹.  That realization would come in surprise fashion two months later when the two adults emerged within a few days of each other…

¹ Ten points to whoever can use this information to arrive at an identification before my next post 🙂

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Photo details:
All photos: Canon EOS 50D, manual mode, ISO-100, 1/250 sec, MT-24EX flash w/ diffuser caps.
Photo 1: Canon 100mm macro lens, f/22, 1/4 power flash (photo slightly cropped).
Photos 2-4: Canon MP-E 65mm 1-5X macro lens f/16, 1/8 power flash.

REFERENCES:

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.

Pearson, D. L. and A. P. Vogler.  2001. Tiger Beetles: The Evolution, Ecology, and Diversity of the Cicindelids.  Cornell University Press, Ithaca, New York, 333 pp.

Copyright © Ted C. MacRae 2009.

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(My) Introduction to Florida Tiger Beetles

On the last day of July, I hopped into my stuffed-to-the-brim SUV and began the +1000-mile drive to St. Petersburg, Florida.  I had with me my camera and my collecting gear, but also my bike, some beach wear… and my family!  No, this was not a collecting trip, but a family vacation.  We would be staying at my sister-in-law’s condo with my niece and enjoying typical Florida vacation fare – beaches, Disney World, sightseeing.  I even brought my road bike along to enjoy some pancake-flat terrain (a rarity around St. Louis) for a nice change of pace.  Still, I can be rather single-minded when it comes to opportunities for bug collecting, and summer in Florida presents opportunities galore!  Virtually everything there is different, and while I have been to Florida a number of times, my visits have all been during spring and prior to my more recent interest in tiger beetles.  It would be a family vacation, but still I would find a way to sneak off a couple times and satisfy my compulsions, and while the girls talked about Disney World and the beach, visions of Cicindela abdominalis, C. highlandensis, C. scabrosa, Ellipsoptera gratiosa, E. hamata lacerata, E. hirtilabris, E. marginata, Habroscelimorpha dorsalis, H. severa, and H. striga danced through my head!

I wouldn’t have to go far to find my first Florida tiger beetles.  My sister-in-law’s condo is in Seminole (west side of St. Petersburg peninsula), and as my niece and her husband showed us around after our arrival, they pointed out the “wild area” off their back patio next to the intertidal waterway and suggested, innocently, that I could go bug collecting back there.  In such a developed urban/tourist area, I figured it must be highly disturbed, dominated by exotics, and offering little in the way of quality habitat for the serious bug collector.  However, that night, as a cacophony of nasal queenks from the area made it clear that a healthy population of what I presume to be green treefrogs were thriving in the area, I reconsidered my skepticism and decided to take a quick look a day later.  What I found was a small but high-quality strip of coastal scrub and saltwater marsh bordering the intracoastal waterway, with thickets of red mangrove (Rhizophora mangle) along the water’s edge, black mangrove (Avicennia germinans) “woodlands” in the high scrub areas, and moist, briney, barren ground in between (Photo 1).  The whole area couldn’t have been more than a hundred yards wide but extended along the length of the waterway, and a wooden sign as I entered the area indicated it was a bona fide, albeit private, nature preserve established as part of the condominium development.

Intertidal salt marsh, Tara Cay Sound Nature Preserve, Seminole, Florida

Intertidal salt marsh, Tara Cay Sound Nature Preserve, Seminole, Florida

Within minutes after entering the preserve I saw the first tiger beetle.  I didn’t know what it was, and my first stalking efforts were woefully inadequate.  I saw another one a few minutes later and got within net handle distance but muffed the swing.  A few feet further along the path and onto an exposure by the water revealed several individuals, one of which I was finally able to capture after several clumsy misses.  I hadn’t yet memorized key characters for all of the species I had the potential to encounter, but I could see in the hand that this individual almost certainly belonged to the genus Ellipsoptera (Ellipsed-winged Tiger Beetles) because of its resemblance to E. macra (Sandy Stream Tiger Beetle) and E. nevadica knausii (Knaus’ Tiger Beetle), which I had photographed earlier this year in Missouri and Oklahoma, respectively.  With one now in the hand as a studio backup, I began my efforts to obtain field photographs.  The beetles were extremely wary, with fast running and strong flight capabilities that made them very difficult to approach.  It was only their abundance and my dogged persistance that allowed me to finally get close enough to one (Photo 2) to fire off a sequence of frames.  I then spent some time collecting a voucher series – finally getting a beat on their behavior and able to capture them with a little more efficiency despite their ultra-wariness before concentrating on getting more photographs of different individuals (including those shown in Photos 3 and 4).

Ellipsoptera marginata - Margined Tiger Beetle

Ellipsoptera marginata - note "tooth" under right mandible of this male

Ellipsoptera marginata - Margined Tiger Beetle

Ellipsoptera marginata - another male, with a distinctly bronzed pronotum

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Ellipsoptera hamata lacerata - Gulf Beach Tiger Beetle (mate guarding)

I returned to the condo after a couple of hours – completely drenched (remember, it was August in Florida!). When I had the chance to consult my “bible” (Pearson et al. 2006), I learned there there were two possibilities: Ellipsoptera marginata (Margined Tiger Beetle), occurring along the Atlantic seaboard from Maine to the Gulf coast of Florida; and Ellipsoptera hamata lacerata (Gulf Beach Tiger Beetle), resembling and closely related to E. marginata and occurring along the Gulf Coast from Florida to Texas.  Both species inhabit coastal beaches, mud flats and salt marshes, and their distributions overlap along Florida’s Gulf Coast. While they are easily distinguished from other species in the genus by the distinctively diffuse middle band of the elytra, distinguishing between the two requires examination of the male right mandible (marginata bears a distinct tooth on the underside, hamata does not) or female elytral apices (marginata curiously bent down at a 90º angle, hamata not).  At this point, I didn’t know if I had one species or two – and if I did have two, did I have photographs of both (and would I be able to identify them)?

Fortunately, closer examination of the voucher series I collected revealed both species present, and even more fortunately I had managed to get photographs of both.  The individual in Photo 2 is a male, and the angle of the photograph clearly reveals a distinct tooth on the underside of the right mandible, identifying it as E. marginata.  Photo 3 is another male, and although the angle doesn’t afford a view of the mandibles, what can be seen is a bronze cast to the pronotum – in my voucher series, all of the E. marginata specimens have a bronze cast to the pronotum, while the E. hamata lacerata specimens exhibit an olive cast. This suggests that this individual also represents E. marginata. In Photo 4, no tooth can be seen on the male mandible, but the angle of the photo doesn’t necessarily make it visible were it to exist.  The male does, however, exhibit an olive cast on the pronotum, and the female elytral apices show no indication of being bent down as would be expected for E. marginata (see Photo 5 below of E. marginata photographed a few days later at another locality).  As a result, the individuals in Photo 4 can be identified as E. hamata lacerata.

Ellipsoptera marginata - female elytral apices showing curiously "bent tips"

Ellipsoptera marginata - female elytral apices showing curiously "bent tips"

Of the 19 vouchers that I collected, 15 represent E. marginata and 4 represent E. hamata lacerata.  This exemplifies the challenges of field identification and photography of insect species in unfamiliar places.  Had I been satisfied with photographing only one or two individuals of what I thought were all the same species, I would have missed one of the species.  The experience further exemplifies the importance of adequate voucher series, as it was only a result of close examination of that series that I realized two species were present.  Based on the numbers of individuals that I collected, as well as the photographs that I took, it appears that E. hamata lacerata was much less common at that location than E. marginata. This difference in population density between two similar, co-occurring species emphasizes the importance of sampling an adequate number of individuals at a given location before concluding what species are – and are not – present.

REFERENCES:

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 2009

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