Landscape Ecology of Singing Insects 3: Changes Over Time

by Carl Strang

Some of the formalism developed by landscape ecologists clarifies the current status of our singing insects and their prospects for survival. From the standpoint of a given species, the landscape is viewed as patches of habitat imbedded within a matrix of non-habitat. If that matrix is hostile to the species, the habitat edge is a “hard boundary,” and unless the species is good at dispersing long distances, they will be confined to their habitat island. To different degrees that extreme may not be met if there are steppingstones or corridors that can act as acceptable temporary refuges for dispersing individuals. I suspect that such is the case with pine tree crickets, for example. These habitat specialists are common in conifer groves across the region, many of which seem too isolated for such a small, specialized insect to cross the intervening distances. Nancy Collins, a Wisconsin specialist on the subfamily, has noticed that pine tree cricket nymphs can live, at least temporarily, in non-conifer, herbaceous habitats. Edges of conifer groves thus are not hard boundaries for pine tree crickets. In other cases, river corridors and highway rights-of-way can serve as travel lanes. Thus, the little-known dispersal abilities of our various species are key to understanding their status.

Pine tree cricket

In some of the crickets and katydids which normally are short-winged and flightless, long-winged morphs occasionally appear. Roesel’s katydid, mentioned in the previous post, essentially is never long-winged in its native Europe, but commonly is so here, where the species is expanding its range. There is a tradeoff in play for such species. Long-winged variants are good dispersers, but their fecundity is reduced.

Roesel’s katydid, long-winged variant

Dispersal also can be facilitated by humans. I have seen examples, with Japanese burrowing crickets and jumping bush crickets, of individuals and small groups showing up far ahead of the front of their range expansion, in places where landscape materials are stored or sold.

Mate finding motivates some movement by singing insects. The females in general must travel to meet up with singing males. The males themselves also may need to change their position. In DuPage County, where swamp cicadas are relatively few and scattered, I have observed males frequently changing position between songs by tens of meters at a time. An Iowa study (Shaw, Bitzer and North 1982) found that sword-bearing coneheads shifted position an average of 6.2m between nights, but otherwise remained associated with their group of other males.

Swamp cicada

The singing insects whose habitat needs are met by landscape alterations for agriculture and residential areas are the abundant, widespread ones. They are easily picked out by the large number of sites marked on their maps in my singing insects guide. Dispersal is relatively easy for them as there are large habitat blocks well connected by amenable corridors. Even they can suffer local extinction as land is cleared of vegetation for buildings or roadways. As new vegetation grows into such places, the weedy species are quick to re-establish themselves, but this underlines the dynamism of the habitat patch mosaic.

Of greater interest, and greater concern, are those species whose needs are not met by human-created habitats. Here the habitat patches are only a small percentage of the landscape, and elements of patch size, patch isolation, and insect dispersal ability become critical to understanding. Theory suggests that when a habitat falls below 10-20% of the landscape, dispersal ability is expected to drop dramatically in the absence of viable corridors or steppingstones; such is certainly the case for many of our habitat specialists. A few species anecdotally are good dispersers. I have reports, or have seen myself, instances of slender meadow katydids and long-tailed meadow katydids showing up in locations remote from their respective habitats. These individuals were unusual among the small meadow katydids in being long-winged morphs; most cannot fly.

Slender meadow katydid

Patch isolation in some cases is such that successful dispersal is impossible. Distances among the few surviving sphagnum bogs, for example, are too great to be crossed by sphagnum ground crickets. Prairie cicadas can fly, but apparently are disinclined to do so far enough to matter. At one of their sites, the West Chicago Prairie in DuPage County, they remain confined to one area of mixed grasses and forbs. They have not crossed the 350 meters of grasses, small shrubs and wetlands that separate them from a similar area within the same preserve. So far, their populations have persisted in remnant prairies ranging from 1 to a few acres. Can such small populations survive long term?

Prairie cicada

That question raises another concept from landscape ecology: extinction debt. This is the idea that a species’ habitat may have been so reduced that it still may be present but is doomed to fade away to extinction because its numbers are inadequate to maintain reproductive viability. Such may or may not be the case for prairie cicadas. I suspect this is what happened to northern wood crickets (NWC) in the region. NWC were known from two forested areas in northern Indiana at the beginning of the 20th Century (Blatchley 1903). One of these was in Marshall County. Years ago I used the original survey notes and county soil map to reconstruct the presettlement landscape of the township in question.

Presettlement map of Union Township, Marshall County, Indiana. Mesic forest was the large green area east of Lake Maxinkuckee.

Blatchley found NWC in the area of mesic forest which, in 1834, was very large. By the end of the 19th century agricultural clearing would have been well under way, and today only a tiny portion of that forest remains, the rest having been converted to pasture and crops. The same is true of forested areas in Lake County, Indiana, the other area where NWC lived in the late 1800’s. Though fragments of the forests where Blatchley found NWC remain, the crickets are gone, and I have checked all the other relatively large forests in the region without finding them. This raises the disturbing question: how many other species presently in the Chicago region are in a state of extinction debt? Patch size needed to maintain a species is dependent upon the characteristics of the species, its population dynamics, and patch quality. These are unknowns for all the uncommon species.

Our two species are Linnaeus’s 17-year cicada, on the left, and Cassin’s 17-year cicada, on the right

I will close this section with a case study on the periodical cicadas in DuPage County. In pre-settlement times the county was, from the cicadas’ perspective, a matrix of prairies and wetlands with 18 forested areas of various sizes scattered throughout. Historical maps and aerial photos allowed me to trace changes in those forests from the early 19th Century to the present day. I measured the forest sizes, noting their smallest (bottleneck) extents and how those were reflected in the presence of periodical cicada choruses in the 2007 emergence. There was a clear threshold of local extinction: forests which had remained above 61ha (hectares) still had cicadas, those which had fallen below 52ha did not, even when they had grown larger subsequently. Old newspaper accounts placed cicadas in at least some of these. Three forests which fell between those sizes appeared to have been affected by isolation, a remote one lacking cicadas, and two near persisting populations having them. The human history peculiar to the county is important here. DuPage County is immediately west of Chicago. The western half of the county quickly became agricultural, with forests cut back to make room for fields and pastures. Forests lacking cicadas in 2007 were mainly in the west. The eastern half developed residential commuter communities, with forests being protected and expanded as people planted trees around their homes. The largest area with cicada choruses in 2007 was lobular in shape, the various lobes following the routes of commuter-serving railways and the towns that expanded along them, connecting several of the pre-settlement forest locations. When the cicada choruses reached their peak in the first half of June, numbers of the insects suddenly appeared in flight, crossing over highways and other hostile environments. Subsequently, small groups showed up in places remote from the concentration areas. Whether these will result in significant expansion of the species in the county remains to be seen, but this observation supports the notion that competition and population pressure produce responses by the cicadas. They have the advantages of stronger flight capabilities and better vision than other singing insect groups.

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

Landscape Ecology of Singing Insects 1: Glacial Influences

by Carl Strang

One of my winter projects has been to write new sections for my singing insects guide which go into ecological topics. This was inspired by my reading a newly published textbook on landscape ecology (With, Kimberly A. 2019. Essentials of landscape ecology. Oxford University Press, Oxford, U.K. 641 pp). In the next few posts I will share parts of the added sections. Today’s focus is the impact of the last continental glacier on the landscape and selected insect species:

Though the focus in most of this guide is on the individual species of singing insects, the field of landscape ecology provides a framework of broader patterns and questions for which the ecologically diverse singing insects provide a suitable lens.

Landscape ecology considers geographical patterns and dynamics of their change across time. Any complete biological understanding of the Chicago region must include not only what is here now but also how it got that way. Fortunately, we don’t have to go back too far geologically speaking, as our landscape is less than 20,000 years old. Three major lobes of the most recent continental glacier flowed down from the North and scoured our region’s bedrock, then began their final retreat around 18,000 years ago. As the glacier melted, with occasional periods of stalling when the push from the North was balanced by melting at the edge, it left behind a variably deep layer of various kinds of deposits. The topography was more elevated in the morainal arcs where the melt was stalled for a time, lower and flatter when the melt-back was more uniform and rapid. Occasional pocks formed where blocks of glacial ice were buried and later melted, resulting in small lakes, bogs and other wetlands. Our rivers had their start as glacial meltwater drainage streams. The Lake Michigan Lobe of the glacier picked up and crushed the softer shale from the bottom of what was to become that lake, so it left behind deposits heavy in clay west and southwest of the lake. When glacial crunching and meltwater eroded harder igneous and metamorphic rocks the ice had transported from Canada, gravel and sand resulted. Quartz sand, the most erosion-resistant component of such rocks, accumulated especially around the edge of Lake Michigan and the tributaries of the Kankakee River, which started as a glacial meltwater drainage stream originating in the part of the region covered by the glacier’s Saginaw Lobe.

This glacial history impacted our singing insect fauna in various ways. Some species require, or at least are only abundant, on sandy soils. These include the green-winged and northern dusk-singing cicadas, whose nymphs live on buried plant roots, and some of the grasshoppers, for which the sandy substrate for egg-laying and/or a poor-soil plant community is an important habitat component. A couple of species, the gray ground cricket and the seaside grasshopper, are limited to the beaches and dunes around the Lake Michigan edge.

The beaches at different points on the Lake Michigan shore have different compositions, resulting in selection for different colors in the seaside grasshopper. Here, at Illinois Beach State Park, there is a greater mix of different colors of ground igneous and metamorphic rocks.

The beaches of the Indiana Dunes are a more uniform quartz sand. Here, two seaside grasshoppers (same species as in the previous picture) would be nearly invisible if they were not flashing their inner femur colors at one another.

Kames are gravel hills formed by waterfalls within the melting glacier, and they provide a well-drained substrate. Isolated populations of sulfur-winged grasshoppers and tinkling ground crickets live on a kame in the Lulu Lake Nature Preserve in northern Walworth County, Wisconsin.

Exposed gravel on part of the kame at Lulu Lake.

All species were pushed south by the glacier, surviving in what is now the southern U.S. and being influenced by selective forces then and during the advance north as the climate moderated and plant communities spread back over the barren glacial deposits. Most of our familiar prairie and forest plant species were restricted to rather small refuges in the South during the glacial maximum, though oaks and hickories occupied a large part of the southern U.S.

Black oak savanna, Kankakee County

As the glacier melted back, open sedge tundra with some black spruce trees invaded first, then white spruces filled in to form a recognizable northern coniferous forest until around 15,000 years ago. By around 12,000 years ago most of our landscape was a mix of deciduous species, including woodlands with lots of oaks. Beginning around 10,000 years ago there was a drying period, which led to the spread of prairie through our area. The prairie then retreated as the climate became wetter, so that by 6200 years ago the western part of our region was a prairie with islands of woodlands and wetlands, grading to forest in the eastern part. This reflects a gradient of increasing moisture from west to east, mediated by the flow of wet air circulating north from the Gulf of Mexico. The drier prairies were maintained by fires which frequently knocked back woody plants that otherwise would have converted even the western part of the region into woodlands. The wooded islands within the prairie were not randomly located, but survived where rivers, other wetlands, and topographic breaks shielded certain spots from prairie fires pushed by the prevailing westerly winds. The upshot for our singing insects is a diverse landscape that to this day contains species specializing in prairie, woodland and various wetland habitats, as well as some that thrive around the edges between habitat types. A few species may be relicts of earlier changes in this history. For instance, the delicate meadow katydid, now apparently extinct in the region, is abundant in prairies to the west and probably accompanied the prairie advance. By the early 20th Century it was known in a very few scattered locations. I have not been able to find it anywhere in the present day.

Delicate meadow katydid females have longer ovipositors than their close relatives.

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