Science 26 November 2004:
Vol. 306 no. 5701 pp. 1488-1489
DOI: 10.1126/science.1106582

• Perspective

ECOLOGY

Oh the Locusts Sang, Then They Dropped Dead

1. Richard S. Ostfeld and
2. Felicia Keesing[HN14]

+ Author Affiliations

1. The authors are in the Institute of Ecosystem Studies, Millbrook, NY 12545, USA. E-mail: rostfeld@ecostudies.org
2. F. Keesing is also in the Biology Department, Bard College, Annandale-on-Hudson, NY 12504, USA.

Summary

Seventeen-year cicadas of the species Magicicada spend most of their lives underground feeding on xylem from the roots of trees. But every 17 years they emerge in vast numbers, mate, lay eggs, and die. In their Perspective, Ostfeld and Keesing discuss a new study (Yang) demonstrating that the cicada carcasses provide a rich source of nitrogen, which is released by soil microbes and results in an increase in the nitrogen content and seed size of American bellflowers, an understory plant.

==

The Bob Dylan song “Day of the Locusts” refers to the cacophony from the 1970 emergence of 17-year cicadas (Magicicada spp.), which happened to coincide with his acceptance of an honorary degree from Princeton University. These cicadas, which dutifully reappeared aboveground in 1987 and then again this year, are a quintessential case of a resource pulse—a transient, multiannual episode of resource superabundance. On page 1565 of this issue, Yang (3) [HN3] describes the ramifying impacts that massive pulses of cicada...

Related uses of the word "locust"

Egyptian grasshopper Anacridium aegyptum

The word "locust" is derived from the Vulgar Latin locusta, which was originally used to refer to various types of crustaceans and insects; English "lobster" is derived from Anglo-Saxon loppestre, which may come from Latin locusta.^[10]Spanish has mostly preserved the original Latin usage, since the cognate term langosta can be used to refer both to a variety of lobster-like crustaceans and to the swarming grasshopper, while semantic confusion is avoided by employing qualifiers such as de la tierra (of the land) when referring to grasshoppers, del mar and del rio (of the sea/of the river) when referring to lobsters and crayfish respectively.^[11]French presents an inverse case; during the 16th century, the word sauterelle (literally "little hopper") could mean either grasshopper or lobster (sauterelle de mer).^[12] In contemporary French usage, langouste is used almost exclusively to refer to the crustacean (two insect exceptions being the langouste de désert and the langouste de Provence).^[13][14] In certain regional varieties of English, "locust" can refer to the large swarming grasshopper, the cicada (which may also swarm), and rarely to the praying mantis ("praying locust").^[15]

The use of "locust" in English as a synonym for "lobster" has no grounding in anglophone tradition, and most modern instances of its use are usually calques of foreign expressions (e.g. "sea locust" as mistranslation of langouste de mer).^[16] There are, however, various species of crustaceans whose regional names include the word "locust." Thenus orientalis, for example, is sometimes referred to as the flathead locust lobster (its French name, Cigale raquette, literally "raquet cicada," is yet another instance of the locust-cicada-lobster nomenclatural connection). Similarly, certain types of amphibians and birds are sometimes called "false locusts" in imitation of the Greek pseud(o)acris, a scientific name sometimes given to a species because of its perceived cricket-like chirping.^[17] Often, the linguistic nondifferentiation of animals not only regarded by science as different species, but that also often exist in radically different environments, is the result of culturally perceived similarities between organisms, as well as of abstract associations formed within a particular group's mythology and folklore (see Cicada mythology). On a linguistic level, these cases also exemplify an extensively documented tendency, in many languages, towards conservatism and economy in neologization, with some languages historically only allowing for the expansion of meaning within already existing word-forms.^[18] Also of note is the fact that all three so-called locusts (the grasshopper, the cicada, and the lobster) have been a traditional source of food for various peoples around the world (see entomophagy).

The word "locust" has, at times, been employed controversially in English translations of Ancient Greek and Latin natural histories, as well as of Hebrew and Greek Bibles; such ambiguous renderings prompted the 17th-century polymath Thomas Browne to include in the Fifth Book of his Pseudodoxia Epidemica an essay entitled Of the Picture of a Grasshopper, it begins:

“

There is also among us a common description and picture of a Grasshopper, as may be observed in the pictures of Emblematists, in the coats of several families, and as the word Cicada is usually translated in Dictionaries. Wherein to speak strictly, if by this word Grasshopper, we understand that animal which is implied by τέτιξ with the Greeks, and by Cicada with the Latines; we may with safety affirm the picture is widely mistaken, and that for ought enquiry can inform, there is no such insect in England.[19]

”

Browne revisited the controversy in his Miscellany Tracts (1684), wherein he takes pains (even citing Aristotle's Animalia) to both indicate the relationship of locusts to grasshoppers and to affirm their like disparateness from cicadas:

“

That which we commonly call a Grasshopper, and the French Saulterelle being one kind of Locust, so rendered in the plague of Ægypt, and in old Saxon named Gersthop.[20]

”

Compound words involving "locust" have also been used by anglophone translators as calques of archaic Arabic, Greek, Hebrew, or other language names for animals; the resulting formations have, just as in the case of the Brownian grasshopper/cicada controversy, been, at times, a cause of lexical ambiguity and false polysemy in English. An instance of this appears in a translation of Pliny included in J.W. McCrindle's book Ancient India as Described in Classical Literature, where an Indian gem is said by the Roman historian to have a "surface [that] is even redder than the shells of the sea-locust."^[21]

[EDLIS Notes]

Of course we know it was really the Day of the Cicadas that June at Princeton. But we are not ones to argue with the greatest songwriter of the last century about his choice of words. Perhaps two syllables worked better than three in the song. Or one could easily imagine that Dylan liked using a word that brought up images of the locust plagues in Egypt written about centuries before in the Bible. We think it's a rather good choice for that reason alone.

Second song on the album, New Morning
Released October 21, 1970
Remastered version released March 31, 2009
Please purchase the MP3 from Amazon or iTunes

Cicada song

	Cicada song The sound of a cicada in Lower Hutt, New Zealand. Recorded February 2006. Cicadas in Greece The sound of a tree full of Cicadas in Ithaca, Greece. Recorded July 2008.
Problems listening to these files? See media help.

Male cicadas have loud noisemakers called "tymbals" on the sides of the abdominal base. Their "singing" is not the stridulation (where one structure is rubbed against another) of many other familiar sound-producing insects like crickets: the tymbals are regions of the exoskeleton that are modified to form a complex membrane with thin, membranous portions and thickened "ribs". Contracting the internal tymbal muscles produces a clicking sound as the tymbals buckle inwards. As these muscles relax, the tymbals return to their original position producing another click. The interior of the male abdomen is substantially hollow to amplify the resonance of the sound. A cicada rapidly vibrates these membranes, and enlarged chambers derived from the tracheae make its body serve as a resonance chamber, greatly amplifying the sound. The cicada modulates the sound by positioning its abdomen toward or away from the substrate. Additionally, each species has its own distinctive "song".^[1]

Average temperature of the natural habitat for this species is approximately 29 °C (84 °F). During sound production, the temperature of the tymbal muscles was found to be slightly higher.^[10] Cicadas like heat and do their most spirited singing during the hotter hours of a summer day, in a roughly 24 hour cycle.

Although only males produce the cicadas' distinctive sound, both sexes have tympana, which are membranous structures used to detect sounds and thus the cicadas' equivalent of ears. Males can disable their own tympana while calling.^[11]

Some cicadas produce sounds up to 120 dB (SPL)^[11] "at close range", among the loudest of all insect-produced sounds.^[12] This is especially notable as their song is technically loud enough to cause permanent hearing loss in humans, should the cicada sing just outside the listener's ear (unlikely). Conversely, some small species have songs so high in pitch that the noise is inaudible to humans.^[13] Species have different mating songs to ensure they attract the appropriate mate. It can be difficult to determine from which direction(s) cicada song is coming, because the low pitch carries well and because it may, in fact, be coming from many directions at once, as cicadas in various trees all raise one another to make noise in unison. Although relatively loud, cicada song can be comforting and even hypnotic at times, as it is at its loudest during the hottest time of an already hot day.

In addition to the mating song, many species also have a distinct distress call, usually a somewhat broken and erratic sound emitted when an individual is seized. A number of species also have a courtship song, which is often a quieter call and is produced after a female has been drawn by the calling song.

[EDLIS Notes]

Two examples of the cicada song. Neither sound file gives one a complete idea of the din created by hundreds of them singing at the same time in an area.

Magicicada

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Periodical cicada
Magicicada sp. from Brood XIII, 2007
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hemiptera
Family:	Cicadidae
Genus:	Magicicada
Species
See text.

Magicicada is the genus of the 13- and 17-year periodical cicadas of eastern North America. They are sometimes called "17-year locusts", although cicadas belong to order Hemiptera, while locusts are Orthoptera.^[1]

Magicicada spp. spend all but a very few weeks of their long lives silent and immobile underground. Then thousands of nymphs emerge from holes in the earth and, within hours, become gregarious flying adults, whose courtship calls make them the loudest insect known. Two months later, however, the adults of that brood are all dead, leaving behind eggs that will hatch to become a new underground generation, one that will emerge again in 13 or 17 years.

Contents [hide] 1 Description 2 Life cycle 3 Predator satiation survival strategy 3.1 Impact on other populations 3.2 Broods 4 Taxonomy 5 Media 6 References 7 External links

Description

The familiar winged imago (adult) periodical cicada has red eyes and a black dorsal thorax. The wings are translucent and have orange veins. The underside of the abdomen may be black, orange, or striped with orange and black, depending on the species.^[2]

Adults are typically 2.4 to 3.3 cm (0.9 to 1.3 in), depending on species, slightly smaller than the annual cicada species found in the same regions of the United States. Mature females are slightly larger than males.^[3]

Magicicada males typically form large aggregations that sing in chorus to attract receptive females. Different species have different characteristic calling songs. The call of decim periodical cicadas is said to resemble someone calling "weeeee-whoa" or "Pharaoh."^[4] The cassini and decula periodic cicadas have songs that intersperse buzzing and ticking sounds.^[3]

Cicadas do not bite or sting. Like other Hemiptera, they have mouthparts for piercing and sucking sap from plants. The cicada's proboscis can also pierce human skin, which is painful if it occurs but in no other way harmful. They are not venomous, and there is no evidence that they transmit diseases. They pose little threat to mature vegetation, although planting new trees or shrubs is best postponed until after an expected emergence of the periodical cicadas. Mature plants rarely suffer lasting damage, although some twig die-off or flagging may result if egg-laying is heavy.^[5]

Life cycle

Transformation of the periodical cicada from the mature nymph to the adult

Nearly all cicadas spend years underground as juveniles, before emerging above ground for a short adult stage of several weeks to a few months. The seven periodical cicada species are so named because, in any one location, all of the members of the population are developmentally synchronized—they emerge as adults all at once in the same year. This periodicity is especially remarkable because their life cycles are so extremely long—13 or 17 years. Cicadas of all other species (perhaps 3000 worldwide) are not synchronized, so some adults mature each summer and emerge while the rest of the population continues to develop underground. Many people refer to these non-periodical species as annual cicadas since some are seen every summer. The life cycles of most annual species range from two to ten years, although some could be longer.

The nymphs of the periodical cicadas live underground, often at depths of 30 cm (1 ft) or more, feeding on the juices of plant roots.^[6] They stay immobile and go through five development stages before constructing an exit tunnel in the spring of their 13th or 17th year. These exit tunnels have a diameter of about 1–1.5 cm (0.4–0.6 in).

Emergence holes

Magicicada molting

Magicicada in final molting stage prior to hardening of exoskeleton.

The nymphs emerge on a Spring evening when the soil temperature at about 20 cm (8 in) depth is above 17 °C (63 °F). In most years, this works out to late April or early May in far southern states, and late May to early June in the far northern states. Emerging nymphs climb to a suitable place on the nearby vegetation to complete their transformation into an adult cicada. They molt one last time and then spend about six days in the leaves waiting for their exoskeleton to harden completely. Just after this final molt, the teneral adults are white, but darken within an hour.

Adult periodical cicadas live only for a few weeks—by mid-July, all have disappeared. Their short adult life has one purpose: reproduction. The males "sing" a species-specific mating song; like other cicadas, they produce loud sounds using their tymbals. Singing males of a single Magicicada species form aggregations (choruses) that are sexually attractive to females. Males in these choruses alternate bouts of singing with short flights from tree to tree in search of receptive females.^[7] Most matings occur in "chorus" trees.^[2]

Receptive females respond to the calls of conspecific males with timed wing-flicks, which attract the males for mating. The sounds of a "chorus"—a group of males—can be deafening and reach 100 dB. In addition to their "calling" or "congregating" song, males produce a distinctive courtship song when approaching an individual female.^[2]

Magicicada egg slits (circled in red)

Both males and females can mate multiple times, although most females seem to mate just once. After mating, the female cuts V-shaped slits in the bark of young twigs and lays approximately 20 eggs in each, for a total of 600 or more eggs. After about six to ten weeks, the eggs hatch and the newborn nymphs drop to the ground, where they burrow and begin another 13 or 17-year cycle.

Predator satiation survival strategy

The nymphs emerge in large numbers at about the same time, sometimes more than 1.5 million individuals per acre (>370/m²).^[8] Their mass-emergence is a survival trait called predator satiation: for the first week after emergence, the periodic cicadas are an easy prey for reptiles, birds, squirrels, cats, and other small and large mammals.^[9][10] Early ideas maintained that the cicadas' overall survival mechanism was simply to overwhelm predators by their sheer numbers, ensuring the survival of most of the individuals. It was hypothesized that the emergence period of large prime numbers (13 and 17 years) was a predatory avoidance strategy adopted to eliminate the possibility of potential predators receiving periodic population boosts by synchronizing their own generations to divisors of the cicada emergence period.^[11] A more parsimonious viewpoint holds that the prime numbered developmental times represent an adaptation to colder soil temperatures during Pleistocene glacial stadia, and that predator satiation is a short term maintenance strategy.^[12] This hypothesis was subsequently supported through a series of mathematical models, and stands as the most widely accepted explanation of the unusually lengthy and mathematically precise immature period of these bizarre insects.^[13] The length of the cycle appears to be controlled by a single gene locus, with the 13-year cycle dominant to the 17-year one.^[14]

Impact on other populations

Cycles in cicada populations are significant enough to affect other animal and plant populations. For example, tree growth has been observed to decline the year before the emergence of a brood because of the increased feeding on roots by nymphs.^[15]Moles, which feed on nymphs, have been observed to do well during the year before an emergence, but suffer population declines the following year because of the reduced food source.^[16]Wild turkey populations respond favorably to increased nutrition in their food supply from gorging on cicada adults on the ground at the end of their life cycle. Uneaten carcasses of periodic cicadas decompose on the ground, providing a resource pulse of nutrients to the forest community.^[15]

Cicada broods can also have a negative impact. squirrel populations have been decimated because the egg laying activity of female cicada damaged upcoming mast crops.^{[citation needed]}

Broods

Magicicadas (both teneral and fully developed). Photo by Arthur D. Guilani

Magicicada septendecim

Cicada prior to final molt

Newly molted Brood XIII

Periodical cicadas are grouped into broods based on the calendar year when they emerge.

In 1907, entomologist C. L. Marlatt assigned Roman numerals to 30 different broods of periodical cicadas: 17 distinct broods with a 17-year life cycle, to which he assigned brood numbers I through XVII (with emerging years 1893 through 1909); plus 13 broods with a 13-year cycle, to which he assigned brood numbers XVIII through XXX (1893 through 1905).^[17]

Many of these hypothetical 30 broods, however, have not been observed. Furthermore, two of the brood numbers assigned by Marlatt (Broods XI and XXI) existed at one time but have become extinct. The Marlatt numbering scheme has been retained for convenience, although today only 15 broods survive. ^[18]

Name	Nickname	Cycle (yrs)	Last Emergence	Next Emergence	Extent
01 !Brood I	blue ridge !The Blue Ridge Brood	17	1995	2012	Western VA, WV
02 !Brood II	east coast !The East Coast Brood	17	1996	2013	CT, MD, NC, NJ, NY, PA, VA
03 !Brood III	iowan !The Iowan Brood	17	1997	2014	IA
04 !Brood IV	kansan !The Kansan Brood	17	1998	2015	KS, MO, OK
05 !Brood V	zz !none	17	1999	2016	Northeast OH, MD, PA, VA, WV
06 !Brood VI	zz !none	17	2000	2017	GA, NC, SC
07 !Brood VII	onondaga !The Onondaga Brood	17	2001	2018	Upstate NY1
08 !Brood VIII	zz !none	17	2002	2019	OH, PA, WV
09 !Brood IX	zz !none	17	2003	2020	Western VA, WV, NC
10 !Brood X	great eastern !The Great Eastern Brood	17	2004	2021	From NY to NC along the East Coast, inland to IL and MI2
11 !Brood XI	zz !none	17	2005 !(2005)	5000 !Extinct	Last seen in 1954 in Ashford, CT along Fenton River
13 !Brood XIII	northern illinois !The Northern Illinois Brood	17	2007	2024	Northern IL and in parts of IA, WI, and IN3
14 !Brood XIV	zz !none	17	2008	2025	Southern OH, KY, TN, MA, MD, NC, PA, northern GA, Western VA and WV, and parts of NY and NJ3
19 !Brood XIX	great southern !The Great Southern Brood	13	2011	2024	AL, AR, GA, IN, IL, KY, LA, MD, MO, MS, NC, OK, SC, TN, and VA4
21 !Brood XXI	floridian !The Floridian Brood	13	2000 !(2000)	5000 !Extinct	Last recorded in 1870. Historical range included the FL panhandle
22 !Brood XXII	zz !none	13	2001	2014	LA, MS, OH, KY 5
23 !Brood XXIII	lower mississippi !The Lower Mississippi River Valley Brood	13	2002	2015	IA, IL, IN, WI
1. Consists only of M. septendecim. 2. Largest of all 17-year periodical broods. 3. Premature emergences, or "straggling" occurred in 2003 and 2006.[19] 4. Largest of all 13-year periodical cicada broods. 5. This 13-year brood does not include M. neotredecim.

 

Taxonomy

There are seven recognized species. Three species have a 17-year cycle:

• Magicicada septendecim (Linnaeus, 1758)
• Magicicada septencassini (also called Magicicada cassini) (Fisher 1851)
• Magicicada septendecula(Alexander and Moore, 1962)

Four more species follow a 13-year cycle:

• Magicicada neotredecim (Marshall and Cooley, 2000)
• Magicicada tredecim (Walsh and Riley, 1868)
• Magicicada tredecassini (Alexander and Moore, 1962)
• Magicicada tredecula (Alexander and Moore, 1962)

These seven species are also sometimes grouped differently into three subgroups, the so-called "-decim species," "-cassini species," and -decula species," reflecting strong similarities of each 17-year species with one or more species with a 13-year cycle.^[20]

Generally, the 17-year cicadas are distributed more in the northern states of the eastern United States, while the 13-year cicadas occur in the southern states, but some may overlap, for example, brood IV (17 year cycle) and XIX (13 year cycle) in western Missouri.^[21][22] If this distribution holds, their emergence will coincide in — 2219, 2440, 2661, etc., as it did in 1998^[23] (although distributions change slightly from year to year and distribution maps can be unreliable, especially older maps.^[22]).

Media

Wikimedia Commons has media related to: Magicicada

Hundreds of cicadas, Brood X
(View in high quality)

[EDLIS Notes]  

Brood X - The Great Eastern Brood

The largest of all 17-year periodical broods.
Most recent emergences were in 1970, 1987, and 2004.
Dylan got his Doctorate in Music from Princeton University on June 9, 1970.

Surely we can't be surprised that Dylan noticed the sound of the singing insects all around him.

Watch the above video to see examples of the 2004 emergence of Brood X.