Landscape Ecology of Singing Insects 2: Human Influences

by Carl Strang

The previous post illustrated that the Chicago region has been a dynamically changing landscape through the recent millennia, but that now is overshadowed by the alterations our own species has made. Burgeoning human numbers have overwhelmed the planet’s ecosystems, and the native habitats described earlier mostly have been replaced by agriculture and urban growth in the Chicago region. One of the more dramatic changes is the loss of the Kankakee wetland, once described as the Everglades of the North. That vast wetland was drained for agriculture, and only a few pockets of it survive in preserves. Much of the Kankakee River in Indiana is now a straight channel with constructed high levee banks. Other smaller wetlands received similar treatment, with drainage ditches spreading across the agricultural portion of the region. This is not universally devastating to wetland species. Northern mole crickets, for instance, occasionally can be found along drainage ditches.

Drainage ditch, upper reaches of the Kankakee River, St. Joseph County, Indiana. Note farm fields on both sides.

Prairie mostly has been replaced by agricultural fields, and fire suppression has led to its invasion by woody plants. Specialists such as the prairie meadow katydid, prairie cicada and short-winged toothpick grasshopper are hard to find.

Prairie meadow katydid

My singing insects research has required a lot of driving to reach the relatively tiny surviving preserves and parks to which many of the species are now restricted. Much management effort is required in these little islands to maintain their habitats. There are exceptions, of course. Many other species have thrived under our influence. These are mainly weedy ones such as the striped ground cricket, short-winged meadow katydid and Carolina grasshopper, which do well in disturbed habitats, along with woodland edge species such as the greater angle-wing, snowy tree cricket, and jumping bush cricket, which can meet their needs in residential neighborhoods dominated by lawns and scattered trees and shrubs.

Jumping bush cricket

Habitat destruction is not the only human influence. Climate change is the probable cause of northward range expansions by several singing insect species, and it likely will lead to the extinction of the sphagnum ground cricket from the region as the sphagnum bogs dry up. Say’s cicada and some northern grasshoppers already appear to be pushed out.

Sphagnum ground cricket

Climate change isn’t simply a matter of rising temperatures, as the term “global warming” may seem to imply. Global warming is an accurate enough term, as the simplest way to measure climate change is to track the global average temperature. But the point is that our changes to the Earth’s thin skin of atmosphere are increasing its held solar energy. That energy alters patterns of atmospheric flow and the behavior of storms. Droughts, more frequent flood-causing rains, and seasonal increases or decreases in temperature that seem abnormal are examples of results we observe locally. The singing insects are forced to adjust as best they can. Droughts force sphagnum ground crickets into the wettest parts of their bogs. The severe drought of 2012 concentrated wetland meadow katydids and marsh coneheads into the small portions of the Great Marsh in the Indiana Dunes National Park that remained wet. Oblong-winged meadow katydids may be pre-adapted to such year-to-year variability. Blatchley (1920) observed that their eggs, laid in moist soil, can take 2-3 years to hatch. In my travels through the region I failed to hear a single individual in the years 2010 and 2019, but in other years they have been abundant and widespread. Some of the cicadas and other species may have similar flexibilities.

Oblong-winged katydid

People also have introduced plant species from other parts of the world which, released from the consumers and competitors which hold them in check in their native lands, have become invasive plants here. Their unfair competitive advantage has led to their displacing the region’s native vegetation in an increasing number of places. This is most evident in our wetlands. Wetland meadow katydids and other singing insects are limited to places where native wetland grasses have not been supplanted by reed canary grass, common reed, purple loosestrife, and hybrid cattails. These invasive plants are proving difficult to control, and the outlook is not good for species such as the dusky-faced meadow katydid and marsh conehead. To my knowledge the once relatively widespread stripe-faced meadow katydid now is confined to a single site, and the slender conehead, never known from many locations, apparently is gone from the region.

Stripe-faced meadow katydid

Introductions have not been limited to plants. Several species of singing insects also have been imported. Roesel’s katydid is the most common of these in our region. A European predaceous katydid, Roesel’s was introduced to the Montreal, Quebec, area several decades ago and expanded from there. They occur in open habitats with tall herbaceous vegetation throughout the Chicago region. Japanese burrowing crickets are thought to have arrived at the port of Mobile and spread out from there. They are abundant as far north as Indianapolis, and common in Rensselaer in the southern part of our region. With scattered new appearances each year occurring as far north as DuPage County in Illinois, so far, I expect them to become widespread and abundant here. The tropical house cricket represents the possibility of other, short-term introductions that are unlikely to persist in our climate.

Roesel’s katydid

Making a Case 2: Prairie Meadow Katydid

by Carl Strang

After more than ten years of study, I still have not found several species of singing insects that historically were known in the Chicago region. One of these was the prairie meadow katydid, one of the smaller members of its group. On August 16, Wendy Partridge and I were checking out the bike trail near the nature center at Illinois Beach State Park. Through the SongFinder, which allows me to hear high pitches, I heard faint rapid ticking sounds which, when heard from a closer position, resolved into brief buzzes. They were mechanical and katydid-like rather than cricket-like in quality, and didn’t match anything I had heard before. Wendy, with her unusually good hearing, barely could pick them up. Later, checking reference recordings, I decided they fit the song of the prairie meadow katydid. They were not in the habitat I would expect, however, being in dense vegetation where grasses were mixed with woody plants and forbs, generally shaded. That katydid species has been known to occur in that park, however.

Then on September 27 I caught and photographed a small female meadow katydid in a relatively dry portion of the Gensburg Prairie in Cook County.

Posterior portion of the Gensburg Prairie female.

Posterior portion of the Gensburg Prairie female.

Her color didn’t seem quite right for a short-winged meadow katydid, which is a common species at that site especially in the wetter portions. Studying the photos, and looking back at references, I have decided that she was a prairie meadow katydid. Along the way I looked back at all my photos of short-winged meadow katydids, and found two other individuals that met at least some of the criteria for prairie meadow katydid, one in 2011 at Mayslake Forest Preserve, and one in 2014 in the savanna woodland at Illinois Beach State Park.

The Mayslake female

The Mayslake female

The Illinois Beach State Park female

The Illinois Beach State Park female

All three have curved ovipositors that are slenderer in proportion than in a typical short-winged meadow katydid.

Typical female short-winged meadow katydid

Typical female short-winged meadow katydid

All three also have femoral patterns in which there is a lengthwise pair of lines, as in the short-winged, but have ladders of narrow crossbars rather than being clear between them as in typical short-winged. There is enough overlap in body length, according to references, that it is not a consideration in comparing those two species. The ovipositors are too short, and the femur patterns wrong, for straight-lanced meadow katydid to be a consideration. The Mayslake female is different from the other two individuals in three possibly significant ways: the femoral ground color is green rather than brown or tan, the wings are much longer, and the front of the head does not appear to rise so much (this last being a difference mentioned by W.S. Blatchley in his classic Orthoptera of Northeastern North America, which gives unusually detailed descriptions of species).

Anterior portion of the Gensburg Prairie female. Note the abrupt rise of the tip of the head when viewed from the side.

Anterior portion of the Gensburg Prairie female. Note the abrupt rise of the tip of the head when viewed from the side.

Dorsal view of the Gensburg Prairie female. Blatchley describes the wings of prairie females as covering only one-fourth of the abdomen, two-thirds in short-winged females. Furthermore, the tip of the head, when viewed from above, is relatively wide after an inward bending of its sides, expected from Blatchley’s description.

Dorsal view of the Gensburg Prairie female. Blatchley describes the wings of prairie females as covering only one-fourth of the abdomen, two-thirds in short-winged females. Furthermore, the tip of the head, when viewed from above, is relatively wide after an inward bending of its sides, expected from Blatchley’s description.

Compare this common meadow katydid, with its relatively narrow and straight-sided tip of the head, to the previous.

Compare this short-winged meadow katydid, with its relatively narrow and straight-sided tip of the head, to the previous.

 

Side view of the same male as in the previous photo. Again, note the lack of a rise in the top of the head profile.

Side view of the same male as in the previous photo. Again, note the lack of a rise in the top of the head profile.

Blatchley also believed that prairie meadow katydids occur only in “raw prairie,” a description which applies to Gensburg and Illinois Beach but not to Mayslake. I am inclined to regard the Mayslake individual as an anomalous short-winged meadow katydid, but pending study of museum specimens, am naming the other two prairie meadow katydids.

Recent Indiana Excursions

by Carl Strang

In recent weeks I have visited a few spots in Lake and Newton Counties, Indiana, for the first time. One site in Gary is a state nature preserve with several interdune swales.

Though there are some patches of invasive wetland plants, more than 95% of the area is in native vegetation.

Though there are some patches of invasive wetland plants, more than 95% of the area is in native vegetation.

I had high hopes for this site, which I thought might have stripe-faced meadow katydids and slender coneheads. On an evening excursion and an afternoon one I built a rather mundane species list. In this rainy year it is possible that the target species are present but widely scattered. I want to get in there again in a year when drier conditions might concentrate the species of interest, and also make a larger portion of the site easily navigable.

Willow Slough Wildlife Area in Newton County is a large and diverse area that I barely have begun to explore for singing insects. One target for this year was a roadside ditch lined with native sedges and grasses.

The ditch proved to have only common singing insects, but there was a remarkable concentration of clipped-wing grasshoppers, a non-singing species I have seen in only one other location. This is a nymph; most were adults at this late point in the season.

The ditch proved to have only common singing insects, but there was a remarkable concentration of clipped-wing grasshoppers, a non-singing species I have seen in only one other location. This is a nymph; most were adults at this late point in the season.

I also checked out some narrow drainage swales along an access road closely bordered by forest.

One non-singing species there was the graceful grasshopper.

One non-singing species there was the graceful grasshopper.

Short-winged meadow katydids were abundant, but the population was unusual in that nearly half the individuals were the long-winged variant.

Short-winged meadow katydids were abundant, but the population was unusual in that nearly half the individuals were the long-winged variant.

I wonder if the narrow, constrained habitat has something to do with the oddity of that population.

 

McHenry County Visit

by Carl Strang

On September 3 I drove up to McHenry County, Illinois, to continue my regional survey of singing insects. I spent most of that hot afternoon at the Pleasant Valley Conservation Area.

This county park has some very good woodlands and savannas.

This county park has some very good woodlands and savannas.

The day produced 7 county records.

My first four-spotted tree cricket in McHenry was at Pleasant Valley.

My first four-spotted tree cricket in McHenry was at Pleasant Valley.

A shift to the Hickory Grove Conservation Area produced additional observations, some of them remarkable.

The most unexpected find was a small group of gladiator meadow katydids, still singing weeks after they normally are done.

The most unexpected find was a small group of gladiator meadow katydids, still singing weeks after they normally are done.

The photo shows the characteristic pronotum profile and cerci. The marsh habitat and the distinctive song pattern, with the ticks finishing, rather than preceding, the buzz portion of the song all were consistent with gladiator meadow katydid. The black spots on the abdomen may be signs of a parasite load; could that have delayed the completion of development?

The same site produced this marsh meadow grasshopper.

The same site produced this marsh meadow grasshopper.

The Lyons Prairie and Marsh, administered as part of Hickory Grove by the McHenry County Conservation District, actually is in Lake County. I followed the trail into a portion of the marsh dominated by reed canary grass. In addition to abundant slender and short-winged meadow katydids, I got an intriguing glimpse at a female Orchelimum that might have been a dusky-faced meadow katydid, which I have yet to find in Illinois. I was unsuccessful in getting a better look in that late afternoon, but at some point I need to get back there for a thorough search.

On the way back to the car I spotted this tiny grasshopper. Mature at around 3/8 inch long, it is a non-singing species, the black-sided pygmy grasshopper.

On the way back to the car I spotted this tiny grasshopper. Mature at around 3/8 inch long, it is a non-singing species, the black-sided pygmy grasshopper.

Wetland Concerns

by Carl Strang

A continuing theme in my regional survey of singing insects is the paucity of wetland species. The only one that is present in good numbers in many wetlands is the black-legged meadow katydid. Other species common in wetlands are habitat generalists such as the Carolina ground cricket and short-winged meadow katydid, which don’t truly count as wetland insects. One clear cause of this problem is the loss of high quality habitat to four invasive plant species (purple loosestrife, reed canary grass, common reed, and cattails). All four are capable of completely taking over a wetland, and examples of this can be found for all four. Small numbers of the singing insects mentioned above can be found in such places, but not the other wetland insect species.

This female long-tailed meadow katydid was a rare wetland species holdout in a small pocket of surviving diverse wetland plants surrounded by cattails, reed canary grass and common reed at Waterfall Glen Forest Preserve in DuPage County, Illinois.

This female long-tailed meadow katydid was a rare wetland species holdout in a small pocket of surviving diverse wetland plants surrounded by cattails, reed canary grass and common reed at Waterfall Glen Forest Preserve in DuPage County, Illinois.

On the other hand, there are some good wetlands out there. Many are small, and this along with their isolation may limit them.

This beautiful little fen at Richard Young Forest Preserve in Kendall County is completely surrounded by woodland.

This beautiful little fen at Richard Young Forest Preserve in Kendall County is completely surrounded by woodland.

Many of the interdune swales at Miller Woods in the Indiana Dunes National Lakeshore have been taken over by the common reed. This one was an exception, but I found only black-legs there.

Many of the interdune swales at Miller Woods in the Indiana Dunes National Lakeshore have been taken over by the common reed. This one was an exception, but I found only black-legs there.

This fen-like wetland at Bluff Creek in Wisconsin’s Kettle Moraine State Forest looks very good. I will want to check it earlier in the season next year.

This fen-like wetland at Bluff Creek in Wisconsin’s Kettle Moraine State Forest looks very good. I will want to check it earlier in the season next year.

Still, I have not given up hope. I found a third good population of mole crickets this year, in a swale at Miller Woods.

Mole cricket tunnels look like miniature mole tunnels. They occur around wetland edges, and occasionally have holes to let out the males’ deep chirping calls, and to provide access for attracted females.

Mole cricket tunnels look like miniature mole tunnels. They occur around wetland edges, and occasionally have holes to let out the males’ deep chirping calls, and to provide access for attracted females.

I also found melodious ground crickets at two new sites in Berrien County, Michigan. Though I did not find dusky-faced or delicate meadow katydids at the Indiana Dunes’ Great Marsh this year, I felt curiously encouraged by this.

Both species were there last year, but with the water level higher in 2013, the vegetation was arranged differently and I did not find them in the same place.

Both species were there last year, but with the water level higher in 2013, the vegetation was arranged differently and I did not find them in the same place.

Maybe the populations of many wetland species took a hit in last year’s drought, and were thinly dispersed in the expanded wet areas of 2013. This is, after all, the first year in which I have surveyed many of these sites. If they need a couple years to recover from the drought, maybe I will find the missing species in the future. Still, how to account for the lack of nimble meadow katydids? This species I have yet to find, anywhere. In the heart of the singing insect season I took my sea kayak into an area where they historically were known.

The channels at the south end of Lake Maxinkuckee in Marshall County, Indiana, have become heavily invaded by purple loosestrife.

The channels at the south end of Lake Maxinkuckee in Marshall County, Indiana, have become heavily invaded by purple loosestrife.

Even the patches of emergent water-dwelling knotweeds, a habitat specifically mentioned in some sources as good for nimble meadow katydids, had none as far as I could tell.

Even the patches of emergent water-dwelling knotweeds, a habitat specifically mentioned in some sources as good for nimble meadow katydids, had none as far as I could tell.

So the bottom line is a disappointing season for wetland species, with a few positive points and hope in the possibility that populations are at a low point from which they will recover.

Toward Singing Insect Monitoring: Dominant Frequency

by Carl Strang

When I conduct workshops or lead field trips on singing insects, people commonly ask about monitoring protocols. We have well established monitoring programs in the Chicago area for birds, frogs, butterflies, dragonflies and damselflies, plants and probably other groups I am forgetting at the moment, so how about singing insects? My answer usually revolves around the fact that different people hear singing insects differently, and this obstacle is a challenge that has yet to be solved. An important variable here is that different people hear different ranges of sound frequencies, and commonly older folks (like me) lose the ability to hear high-pitched sounds. Recently I decided to try to get a quantitative handle on this pattern, using my experience as a gauge. I went to the Singing Insects of North America website and The Songs of Insects book by Elliott and Hershberger, and lifted out the dominant frequencies sung by the species in the Chicago region.

A male short-winged meadow katydid, one of the small meadow katydids, whose song has a dominant frequency of 13 kHz.

Most singing insects produce a range of different sound frequencies when they sing, a buzz for instance consisting of a mix of many low- to high-pitched sounds. Different sounds within the mix have different energies or volumes. The dominant frequency is the loudest one in a given species’ mix. Looking at just the dominant frequency, I see that the various local species range from 1.3 to 17 kilohertz (thousands of vibrations per second, a measure of the pitch or highness/lowness of a sound). I can hear every species with a dominant frequency below 13 kHz. In fact the only species I cannot hear at all are the small meadow katydids (the members of genus Conocephalus), which sing in the 13-17 kHz range. Children and young adults can hear these, I have found. A simple, if expensive, work-around is the SongFinder device.

The long-spurred meadow katydid is my marginal species, at 12 kHz. I can hear them from close range in the woods if there aren’t a lot of competing sounds. Interesting to me is the fact that I hear them clearly at the Brookfield Zoo, where they are fairly common. I doubt that the zoo’s long-spurreds have lower dominant frequencies. My best guess is that the relative lack of other sounds in that range, plus the amplification of the songs reflecting from sidewalks and buildings, increases my ability to hear them there.

Long-spurred meadow katydid, my marginal species

Another related variable is a person’s ability to pick up a sound from a distance. Roesel’s katydid has a dominant frequency of 15 kHz. I can still hear them, but less well with each passing year, and I have to be closer to them. Children and young adults easily pick them up earlier in the song and at a much greater distance. Probably what I hear is not that dominant frequency but the lower part of the frequency range included in Roesel’s buzz. Recently I learned that young adults can hear common meadow katydids, dominant frequency 10 kHz, at a much greater distance than I can, though I hear them clearly if I am within, say, 30 feet.

Common meadow katydid

I think that monitoring protocols are possible to develop, but clearly these are variables that will need to be taken into account. There are other obstacles as well, which I will address at another time.

Meadow Katydid Nymphs

by Carl Strang

One project in my study of singing insects is to learn to identify nymphs, the immature stages, of katydids. This could extend the survey season, as I could begin a month or two earlier than at present. At some point, rearing and taking quantitative measurements will be necessary (unless there is some reference I have not found), but for now I am taking a lot of photos and looking for possible characteristics to watch. I am beginning at Mayslake Forest Preserve, which has relatively few species. Recently the first slender meadow katydids matured and began to sing.

Mature male slender meadow katydid. Note the long wings, green cerci at the abdomen tip, and the green femurs with lines of tiny, separated black dots.

Some of the nymphs I was catching in the sweep net looked like they, too, might be males of this species.

This nymph likewise has green cerci, though not yet in adult form, and green femurs with tiny, separated black dots.

Some of the female nymphs show the same color pattern, though they have undeveloped ovipositors rather than undeveloped cerci.

Again, a bright green general color with tiny black dots on the femurs.

In contrast, many of the nymphs show color patterns that I suspect tie them to adult short-winged meadow katydids, the other common member of genus Conocephalus at Mayslake.

Note the browner ground color, the many brown dots, the yellow-brown cerci, and especially the femur with its central clear zone bounded on each side by a brown band.

Females are similar.

Again the green is paler, and the femur shows the same striping.

Some nymphs are more ambiguous, but most seem to fall into one or the other color pattern, which is encouraging.

Red-legged Grasshopper

by Carl Strang

Over the past couple months there have been large numbers of grasshoppers at Mayslake Forest Preserve, especially around trail and parking lot edges where mowed lawns come up against a variety of unmowed forbs and grasses. My focus in recent weeks has been on singing insects, but with that research checklist essentially complete for the year I decided to look into those grasshoppers. Perhaps I waited too long, as every hopper I got a good look at last week appeared to belong to the same species. I collected a couple of them and, after a session with references and the microscope, settled on an identification.

Red-legged grasshopper, Melanoplus femurrubrum

As I went through the references and keys I was surprised by the number of similar species and the small features that can separate them. Very close ones often segregate by habitat, however, and red-legged grasshoppers are one of the most abundant, weediest species. Ecologists think of weeds as organisms which reproduce in large numbers and occur in disturbed habitats. Therefore, animals as well as plants can be weedy.

While scrutinizing grasshoppers in the field I also got looks at several small (Conocephalus) meadow katydids. All clearly were short-wingeds, except for one.

The oddest feature on this female is the kink in her ovipositor.

She had an abdomen tip colored like that of a straight-lanced meadow katydid, but the shorter ovipositor and leg striping pattern pointed to short-winged meadow katydid, so I am staying with that more conservative identification.

Horlock Hill

by Carl Strang

Horlock Hill Prairie is regarded as one of the highest quality bits of dry prairie in northern Illinois. It is located right at the start of the Great Western Trail in Les Arends Forest Preserve in Kane County, and I have zipped right past it on my bike many a time without realizing its significance.

The designated natural area itself is small, at 2-3 acres, but I was impressed by the floral diversity.

Adjacent meadows and prairie restoration projects enlarge the effective area in prairie or prairie-like vegetation. In one of these I spotted a small meadow katydid which appeared at first to be a short-winged, but lacked the orange abdomen tip. He allowed me to get some photos, and when I looked at them later I was pleased to see the distinctive cerci of a straight-lanced meadow katydid.

The cerci are the little pincer-like structures at the tip of the abdomen. They are long, straight, abruptly flattened in the tips, and the small, inward-pointing tooth is near the base of each.

I found a female of this species at Mayslake Forest Preserve last year, but this is the first male I have seen. The only congeners I heard around Horlock Hill through the SongFinder were abundant short-winged meadow katydids. It was a cool, cloudy day, however, and the song of the straight-lanced is a steady uninterrupted faint buzz that may have been too faint to hear under those conditions. Certainly the short-wings were slowed quite a bit.

Otherwise the singing insects there were all of common species, except that I heard a couple probable broad-winged tree crickets. All of this points to a return to that site under warmer conditions.

Stripes

by Carl Strang

Identifying species of small meadow katydids (genus Conocephalus), especially nymphs, can be challenging. Most adults are readily sorted out, especially males if you can get a good look at the cerci or claspers at the tips of their abdomens.  Females are trickier, but their ovipositors often allow distinctions to be made. I’ve decided to take a shotgun approach in my survey at Mayslake Forest Preserve, sweep sampling on a weekly basis and taking photos of as many individuals as I can. Last week I noticed something that is potentially helpful.

This is an unambiguous male short-winged meadow katydid. Cerci are right, lots of orange color around the abdomen tip, short wings, small size.

As I compared my many photos, I found that the color pattern on the sides of the hind femurs drew my eye. Notice the band of clear green color bounded by brown stripes on either side. Here’s a female, again clearly a short-winged.

Though she may be an instar short of adulthood, this one has the ovipositor of a short-winged.

Again the color pattern on the femur appears. I did an Internet search, and except for some photos from Texas, short-winged meadow katydids show this striping pattern across eastern North America. Then I began to compare other species. I started with another familiar one, the slender meadow katydid.

This male I photographed last year has clear green femurs, without the stripes.

Again an Internet and reference book search showed consistently clear green femurs on this species. However, I was surprised to find that a female I had identified a couple years ago as a slender meadow katydid had the femur stripes.

The long wings fooled me. The ovipositor actually rules out slender meadow katydid. This was a rare individual short-winged meadow katydid with long wings.

Next I turned to two species I have been seeking, which could overlap with the short-winged’s habitat. Straight-lanced meadow katydids in references and Internet photos lack the short-winged’s stripe pattern, instead often showing a diffuse blackish zone down that face of the femur. I went to photos of females I tentatively had identified as that species last year.

This one not only has an ovipositor much longer than the femur length, and lacks orange at the abdomen tip, but shows a femur color pattern different from the short-wingeds’ and consistent with straight-lanced reference photos. In fact it appears that the diffuse black zone is the same as the upper dark stripe on the short-winged, but differing in color and not sharply bounded. The lower stripe is there as well, but just as a trace of a line. I believe this individual may indeed have been a straight-lanced meadow katydid.

Others prove, on closer inspection, in fact to have been short-wingeds.

This young nymph has an ovipositor marginally longer than the femur, but has an instar or two to go until maturity and looks awfully orange around the abdomen. The femur color pattern clearly ties it to short-wingeds, and I think that is where this insect belongs.

The other species I need to sort out is the prairie meadow katydid. Few photos of this one are out there. In some there is no stripe pattern like the short-winged’s, in some there is a hint of one. For now I will need to focus on cerci (straighter, more pointed and with distinctly longer teeth than the short-winged male’s) and ovipositors (proportionately more curved than the short-winged female’s). I am encouraged, however, to continue looking for details of color pattern that might provide short-cuts to field identification at least in regional or local populations.

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