Are we loving our prairies/glades/woodlands to death?

I had such high hopes for last weekend’s collecting trip – late May is boom time for insects across Missouri, we have had good moisture this spring, and I would be visiting some high-quality natural communities that I had not visited for a long time.  My stated goals (the jewel beetles, Agrilus impexus and A. frosti) were long shots – I knew that and would have been fine coming home without those species (which I did) had the the collecting been otherwise productive (which it was not).  Still, I’ve been doing this for a long time now, and I’ve learned to draw on my accumulated experience when things don’t go as planned to give myself the best shot at turning a bad collecting trip into a decent one when things don’t go as planned.  The itinerary with which I start is rarely the one that I actually follow, and this past weekend was a good example of such.

My first stop was Ha Ha Tonka State Park, one of Missouri’s premier parks, boasting high-quality chert, dolomite, and sandstone savanna interspersed with dolomite glades.  It is on these glades and savannas that I hoped to find Agrilus impexus, or failing that at least collect a nice diversity of other jewel beetles on the oaks and hickories of the savannas and surrounding woodlands.  However, it was with some reservation that I even came here after being told by my colleague at the Department of Natural Resources just 2 days before my trip that 75% of the park’s grasslands and woodlands had been burned within the past two years.  For an insect collector, this is never good news – in all my years of collecting insects, my experience in relatively recently-burned habitats has been consistent: collecting sucks!  I decided, however, to visit Ha Ha Tonka anyway because of the quality of the natural communities it contains, thinking perhaps I might be able to find pockets of unburned habitat supporting good insect populations.  This was not to be. I beat oak after oak in the savannas and woodlands – nothing!  I swept little bluestem and Indian grass in the glades – nothing!  The foliage was lush and green and the savanna and glade landscapes highly diverse – given the time of season the place should have been teeming with insect life, yet it almost seemed sterile. Were it not for a few Chrysobothris quadriimpressa jewel beetle adults that I found attracted to a recently wind-thrown black oak tree, I would not have seen any insects here at all.  It appeared my fears about park-wide depression of insect populations had been realized.  However, not one to waste a visit I decided to explore some of Ha Ha Tonka’s fascinating geological features.  Ha Ha Tonka contains one of Missouri’s best examples of karst geology, with complex structures formed from the collapse of a major cave system.  The Devil’s Promenade is one of the more spectacular examples of such, its horseshoe-shaped cliff representing the former interior walls of a now-collapsed cave.  As dusk approached, the day’s poor insect collecting caused me to abandon my plans to stay here and blacklight for nocturnal beetles.  Instead, I decided to break from the itinerary, drive further west and explore Lichen Glade Natural Area in the morning before heading to the Penn-Sylvania Prairie BioBlitz later that afternoon.

Devils Promenade, Ha Ha Tonka State Park

Lichen Glade Natural Area is a small area owned by The Nature Conservancy that boasts a high-quality sandstone glade surrounded by post oak/black jack oak forest.  My first visit to the area more than 20 years ago was during May, and it was one of the most productive collecting trips I’ve had with a number of Agrilus spp. (including A. frosti) beaten from post oak (Quercus stellata) along the woodland edge.  I didn’t visit again until fall of 2002, when Chris Brown, Rich Thoma and I found claybank tiger beetles (Cicindela limbalis) sunning on the exposed sandstone outcrops, and I made one more visit the following May to beat more insects off of post oak.  The Lichen Glade that I returned to this past weekend was a very different place from when I last visited – the surrounding woodlands had been extensively opened (I would guess within the past few years based on the size of the post oak resprouts), and fire had been used throughout the area.  Anticipation turned to frustration when no amount of beating of the woodland vegetation and sweeping of the glade vegetation turned up beetles in any appreciable numbers (or any insects for that matter) and two hours worth of effort yielded not a single buprestid beetle!

Sandstone glade community, Lichen Glade Natural Area

With resignation, I headed on over to Penn-Sylvania Prairie, where during the introduction to the BioBlitz I learned that nearly half of the 160-acre prairie was burned last December and all of it had been burned within the past few years.  I knew what I was going to find – nothing!  Okay, I shouldn’t say nothing, as there actually were some beetles present.  However, the numbers and diversity were low, with all of the species encountered representing common, widespread species.  Moreover, it was not just beetles – all of the invertebrate group leaders (which included experts on snails, ants, butterflies, and bees) reported low overall abundance and diversity in their groups of interest.  Only the vascular plants – the metric by which the value of prescribed burning is always assessed – showed high diversity, with 300 species of mostly native prairie plants recorded for the site.  It was a fun event, with probably ~75 attendees and a delicious pot luck dinner that evening; however, it would have been more enjoyable had there actually been a nice diversity of insects present to document for the preserve.

My comments may make it seem that I am against the use of prescribed burning.  This is not true – I understand the critical role that fire as a management technique plays in restoring and maintaining examples of Missouri’s historically fire-mediated landscape. Without fire and other processes to mimic natural disturbance factors, most of Missouri’s historical grasslands and woodlands suffer relentless encroachment by woody vegetation. However, the modern landscape is very different from the historical landscape, where fires of unpredictable scale, intensity, and frequency operated within a vastly larger scale to create a shifting mosaic of natural communities in various stages of ecological succession. Such processes cannot be recreated on today’s severely fragmented landscape, where the precious few remaining tracts of native habitat are relatively to extremely small and more often than not separated from each other by vast expanses of homogeneous and “inhospitable” habitat (e.g., agricultural, urbanized, or severely degraded lands).  It is in that context that I have great concerns about how aggressively fire has been used in recent years on our state’s natural areas and the impact this is having on insect populations – specialist and generalist alike.  Fire proponents will point to published studies that show little to no effect by the use of fire for managing small, isolated remnants on specialist insects (see review in Henderson 2010).  However, there are an equal number of studies that suggest such concerns are well-founded (see review in Panzer 2002). A consistent limitation in all of the studies that have been conducted is the lack of very large and long un-burned remnants.  Prescribed burning has been adopted so rapidly and pervasively that there just aren’t any significant un-burned remnants left to properly include as controls in such studies.  As a result, the insect fauna present at a given site at the start of such a study is already skewed towards those species that successfully recolonized the area post-burn.  At a minimum, the data to this point are inconclusive, and certainly the potential for impacts has not been given the consideration it warrants in designing fire-management plans for our own state’s prairies and glades. Furthermore, as rapidly and aggressively as fire has been adopted on our few, small, widely disjuct remnants, the opportunity for proper investigation of those potential effects may be gone.  A particularly egregious example of the lack of consideration being given to prairie invertebrates in designing fire management plans is shown in these photos of Iowa’s Sylvan Runkel State Preserve before and after a late May burn and the impact of that burn on a resident population of Nevada buck moths (Hemileuca nevadensis).

Here in Missouri, as in Iowa, it’s a problem of scale – the landscape is too fragmented and remnants too disjunct to manage based strictly on floristic response.   Populations of generalist insect species will recover, and even specialist species may be able to overcome such management practices if they are widely distributed and sufficiently mobile. But what about conservative species with low vagility, such as the swift tiger beetle (Cylindera celeripes) and our disjunct population of the frosted dromo tiger beetle (Dromochorus pruinina), flightless species restricted in Missouri to the few tiny remnants of loess hilltop prairie in northwestern Missouri and a single 2.5-mile stretch of roadside habitat in west-central Missouri?  Until directly relevant data, gathered here in Missouri, are forthcoming to suggest otherwise, I believe the most judicious use of fire possible should be practiced in restoring and maintaining our grasslands and woodlands.  In-season burns may have been a part of the historical landscape, but their use today has great potential to result in local extirpations and should be used only after the most careful consideration.  Leaving un-burned refugia within remnant habitats to accelerate recovery would also be prudent – yet many land managers disregard this practice because of its logistical difficulties. This is especially true in small parcels, yet it is precisely these remnants that have the most to gain from their use (or lose from not doing so!).  In the historical landscape, every burn was a patch burn – no matter what its size, there were always adjacent or proximal unburned habitat from which recolonization could occur.  Elk and bison, too, were integral components of the presettlement prairie landscape – their roamings caused intermittent, localized disturbances that were likely not only crucial to the tiger beetles that I study but may also have contributed to vegetational diversity through patch succession.  Techniques that mimic these natural disturbance factors include mowing, haying, and managed grazing.  They can be utilized to mimic those disturbances as well as delay woody encroachment, and their use in land management should be considered for their ecological value rather than deprioritized because of their relatively greater complexity and cost to implement. Mechanical removal and selective use of herbicides offer additional tools for addressing woody encroachment while minimizing potential impacts to insect populations. An effective management program that considers all of the flora and fauna of a remnant may not be possible unless all of these management tools are utilized, or at least properly considered. As my good friend James Trager said in a recent email (quoting Andrew Williams), habitat restoration “cannot rest on any single management practice, nor practicing it too extensively.”

REFERENCES:

Henderson, R. A.  2010. Influence of Patch Size, Isolation, and Fire History on Hopper (Homoptera: Auchenorrhyncha) Communities of Eight Wisconsin Prairie Remnants.  Wisconsin Department of Natural Resources, Research Report 189, 22 pp.

Panzer, R. 2002. Compatibility of prescribed burning with the conservation of insects in small, isolated prairie reserves. Conservation Biology, 16(5):1296-1307.

Copyright © Ted C. MacRae 2010

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The Moth and Me #11

The Moth and MeWelcome to issue #11 of The Moth and Me, the monthly carnival devoted to the “forgotten” lepidopterans. Most people – even entomologists – regard these as the lesser leps, denizens of the night, as if to hide their somber-colored drabness from the flashy brilliance of their rhopaloceran relatives. Of course, this simply isn’t true, as the contributions to this month’s issue well demonstrate. Butterflies may be among the largest insects on earth, but the largest lepidopteran in the world is a moth. They may also be as gaudily colored as the rainbow itself, but what butterfly is more colorful than the Urania day-flying moths (the genus name literally means, “The heavenly one”).   And, they may be almost universally accepted by a largely insect-indifferent public, but who among us does not think back to that first sight of a luna moth as the most stunning insect we had ever seen to that point.  Yes, moths are all that butterflies are, and for this month’s issue of TMaM, 15 contributions by nine writer’s show us why.

Family Saturniidae – Giant Silkworm & Royal Moths

Tales from the Butterfly Garden: LepcuriousLuna moths belong to the royal moths of the family Saturniidae, and as the name implies they are not the only stunningly beautiful member of the group. Kristen at Tales from the Butterfly Garden: Lepcurious writes about an encounter with the Sweetbay Silkmoth (Callosamia securifera).  Like other members of the family, larvae of this species are rather particular about the type of tree that they utilize for food, which in the case of this moth is sweetbay (Magnolia virginiana).  I’m a little too far north here in Missouri for this tree, so I have never seen this moth.  However, I have seen (and reared) some of its close relatives, the Promethia Silkmoth (Callosamia promethea) which hosts on several plant species and the Tulip-tree Silkmoth (Callosamia angulifera) which hosts on Tulip Tree (Liliodendron tulipifera).

Family Zygaenidae – Leaf Skeletonizer Moths

xenogereJason Hogle at xenogere is fond of the unusual and has a gift for finding it. In his post The Unmoth, Jason shows us a male grapeleaf skeletonizer (Harrisina americana) – not your typical moth!, The uniformly black color and bright red neck collar just screams “Don’t eat me – I’m poisonous”, and indeed species in this family are among the few insects capable of producing hydrogen cyanide!  As the name suggests, larvae skeletonize the leaves of both wild and cultivated grapes (Vitis spp.), as well as the related Virginia creeper (Parthenocissus quinquefolia).

Family Noctuidae – Noctuid Moths & Tiger Moths

Tales from the Butterfly Garden: LepcuriousRoyal moths are not the only stunningly colored moths that Kristen at Tales from the Butterfly Garden: Lepcurious has found in Florida, as she shows in this post on Oleander Moths (Syntomeida epilais) and a companion piece on its Oleander host plant.  This striking day-active moth, also called Uncle Sam Moth (for its red, white, and blue colors) and Polka-Dot Wasp Moth (for obvious reasons), may seem like an easy-to-spot target for would be predators, but its gaudiness is actually warning of the toxic chemicals it has sequestered in its body from the Oleander on which it fed as a larva.  Oleander contains the toxins oleandrin, a cardiac glycoside, and neandrin and is toxic if ingested.  Although oleander is an Old World exotic, oleander moths may also be found feeding on devil’s potato vine (Echites umbellata), which may have been their native Florida host before the introduction of oleander to the United States.

See TrailAside from the underwings (genus Catacola) and the recently incorporated tiger moths, Noctuids are typically thought of as the “basic brown moths” – relying on just the aforementioned groups to add a splash of color to the family’s otherwise drearyness.  Nothing could be further from the truth – check out the stunning Eight-spotted Forester (Alypia octomaculata) in this post by Matthew York at See Trail. Larvae of this beautiful little moth feed on ampelopsis, Virginia creeper, and other plants in the grape family (similar to the grape leaf skeletonizer above). “A great moth; brilliant color, diurnal…… and yes… Noctuid. Some moths, like people, don’t go with the trends.”

See TrailFor the most part, tiger moths shun the daytime in preference for the safety of the night. That does not mean, however, that they are any less colorful, as Matthew York at See Trail shows in his post Poor Grammia. Notarctia proxima, the Mexican Tiger Moth, and its relatives have had a bit of name shuffling over the years at the hands of taxonomists – formerly placed in the genera Grammia and Apantesis. Whatever name you call it, the striking white and black striped forewings give a clue about their common name of tiger moths, and the red, black-tipped abdomen not only add to its beauty, but belies the defensive compounds it surely contains.

Speaking of tiger moths and defensive compounds, watch the video that Chris Grinter at The Skeptical Moth included in his post Moth Perfume. In it, Chetone angulosa gives a striking display of a common defensive mechanism for the group – excreting hemolymph (sweating blood, so to speak!). So spectacularly does the moth do this that you can actually hear the hissing sound of the fluid being pumped from the body. Moreover, there seem to be at least a couple of active ingredients in the froth – one that smells like peppermint, and another that causes numbing of the tongue (as Chris can testify firsthand – he is a truly dedicated experimental naturalist!).

Karthik's JournalIn similar fashion to our North American species of underwing moths (Catocola spp.), the related Eudocima materna, one of the fruit-sucking moths of south India, uses its drab-colored forewings to hide its brilliantly colored hindwings, as Karthik at Karthik’s Journal shows us in his post Startling Displays.  This forms a double line of defense against would-be predators – the forewings blend marvelously into the color of the tree trunks upon which it rests during the day, camouflaging the insect and making it nearly invisible.  If this doesn’t work, a sudden flash of the hindwings may startle the predator just enough to allow the moth to take flight to another tree – where it instantly “disappears” as soon as it closes its wings.

Snails Eye ViewAustralia also has some very colorful fruit-piercing moths, and Bronwen Scott at Snails Eye View presents some beautiful photos of the particularly strikingly-colored Othreis iridescens. Like other members of the group, this Far North Queensland endemic feeds on fruit (Pycnarrhena novoguineensis and Hypserpa laurina, both Menispermaceae, in the case of this species), but as it is apparently the rarest of the primary fruitpiercing moth species in Australia it is not considered to be a pest (and Bronwen would cut it some slack even if it was!).

EntophileAdults are but only one of four life stages that all moths go through. If moths are the “forgotten” leps, then caterpillars are the “forgotten” moths. In many cases, the caterpillar stage cannot be recognized until it becomes a moth (and in some cases the caterpillars are completely unknown). Fortunately, Navy entomologist corycampora at Entophile recognized the caterpillar he found on his croton bush, which he features in the post Croton caterpillar, Achaea janata (Linnaeus), (Lepidoptera: Noctuidae). These “eating machines” can be just as fascinating to observe as their scaled adult counterparts, and while croton seems to be a preferred host in Hawaii, it apparently also feeds on castor beans (judging by its other common name, Castor Oil Semi-looper).

Family Notodontidae – Prominent Moths

the Marvelous in NatureOften dismissed as noctuids, the prominent moths tend to be fuzzier, more thickly-bodied moths that rest with their wings curled around their abdomen or tented over their back (rather than flat like noctuids and most other moths). TMaM organizer Seabrooke Leckie at the Marvelous in Nature has a love affair with prominents, and in her post Georgian Prominent, she features the nicely thick-bodied and fuzzy Georgian Prominent, Hyperaeschra georgica. The caterpillars of this widespread species feed on oak (Quercus spp.), thus, unless you live in the Pacific Northwest you stand a good chance of encountering this species – if you’re you’re willing to make the effort.

Family Psychidae – Bagworm Moths

xenogereMany of us are probably familiar with the evergreen bagworm moth (Thyridopteryx ephemeraeformis), whose large, cone-shaped bags almost look like fruit hanging from the evergreen bushes on which the caterpillars feed. But did you know there are other species of bagworms as well? Jason Hogle at xenogere does, and he compares and contrasts two of them in this duo of posts, Rainy day on the patio and The Other Bagworm. One huge and prominent, the other (Dahlica triquetrella) very small and oft unseen. One with all manner of plant matter stuck to its bag, the other usually mistaken for small bits of dirt or wood. Jason is so good, he can even determine the sex of the caterpillar inside the bag!

Family Sphingidae – Hawk Moths

Roundtop RumingsCarolyn at Roundtop Rumings is hoping that somebody can Name this moth, which she found on the door of her cabin in the forests of Pennsylvania. Don’t let her inability to name this moth fool you, however, for her post contains loads of information on exactly the kinds of characters one should take note of when trying to identify hawk moths. Large size and membership in a popularly studied group aren’t enough – what do the hindwings look like? Are there any spots on the abdomen? As a coleopterist, I hesitate to offer my relatively uninformed opinion on the exact genus and species for this moth, but I’m going to go out on a limb here and suggest maybe something in the genus Ceratomia, perhaps the waved sphinx (C. undulosa)?


I hope you have enjoyed this issue of The Moth and Me, and my sincere thanks go out to all of those who contributed!  The hosting slot for next month’s issue of TMaM is still open, but you can submit your contributions anyway to Seabrooke Leckie at the home site for inclusion in the June 2010 issue once a host is selected.  The submission deadline is June 13, with the issue appearing a few days later.  Perhaps you might like to host the June issue – hosting is not only fun, but also a great way to introduce readers to your site and generate a little traffic.  Contact Seabrooke at the home site if you’re interested – I’m sure she would love to hear from you.

Copyright © Ted C. MacRae 2010

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