PART 2 OF A 3 PART SERIES Insect and Mite Galls: Myths and Misconceptions In Part II of this series, we learn about insect and mite galls, including the Six Laws of such galls. By Joe Boggs and Jim Chatfield n the first installment of this series Although insect and mite gall forma- sharp, piercing mouthparts (chelicerae) on plant galls (May 2015), we talk- tion is not entirely understood, research- to rupture plant cells so they can feed on ed about the difference between ers theorize there are two possible path- the contents. Only the feeding activity of gall-like structures and true galls, ways. Some gall researchers believe certain some species of eriophyid mites induc- including bacterial crown galls, types of plant gall growth are directed by es gall growth; there are no spider mite fungal galls, leaf/petiole galls, the feeding activity of the gall-maker. The gall-makers. This gall-growth pathway flower/fruit galls, bud galls and galls are produced by a combination of may explain how simple felt-like erineum stem galls. Galls galore! But there’s more. constant but subtle feeding irritation, per- patches (a.k.a. “erineum galls”) develop ILet’s dig deeper into insect and mite haps coupled with the release of chemical under the direction of a number eriophyid (arthropods) galls. Unlike bacterial crown inducers by the gall-maker. mite species. However, it does not explain galls, which are a mass of plant cells that Certain eriophyid mites provide an how highly organized plant gall structures have been modified by bacterial DNA, or example. These unusual mites are much develop. fungal galls, which are an assemblage of smaller than spider mites (you need at Some types of insect and mite galls fungal cells intertwined with plant cells, least 40x magnification to see them); are composed of complex plant struc- galls produced by insects and mites are however, both types of mites use their tures and may include functional plant constructed entirely of plant cells. Dominant arthropod gall-makers in- clude insects belonging to three orders: Hymenoptera (wasps, sawflies); Hemip- tera (aphids/adelgids, phylloxerans, psyl- lids); and Diptera (midge flies). The only mites capable of inducing galls are erio- phyids (order Trombidiformes, family Eriophyidae). There are more than 2,000 species of insect gall-makers in the U.S.; however, three quarters belong to only two families: Cynipidae (“gall wasps”) and Cecidomyii- dae (“gall midges”). Note the “cecido-” in the name of the gall midge family; Ceci- dology is the scientific study of plant galls. Of the over 800 different gall-makers on oaks, more than 700 are gall wasps. In other words, when trying to identify an insect gall, keep in mind that there is a high probability it was produced by a gall wasp or gall midge. Top of page, fuzzy, felt-like erineum patch galls appear on a birch leaf. All photos: Courtesy of Joe Boggs 22 | June 2015 | American Nurseryman Mossy rose gall is a good example of a plurilocular, unilarval gall (multiple chambers but only one larva per The large burl affecting this walnut is not a true gall. chamber), while hickory phylloxera petiole gall is a typical unilocular, multilarval gall (one chamber hosting several Below, the oak anthracnose fungus, Apiognomonia larvae). quercina, is seen infecting the plant tissue of the small oak apple gall.. organs such as nectaries. These types of cells that were originally destined to plant galls require a different gall-growth become leaf cells begin marching to a theory; one that includes the ability of the different drummer. Once the errant leaf gall-maker to turn plant genes on and off cells fall under the chemical spell of a gall- as they direct plant cells to form highly or- maker, there is no turning back—they ganized plant structures. will become gall tissue. This means that Research has revealed that some gall formation cannot occur once meri- gall-making insects and mites produce stematic leaf bud cells are committed to chemical replicas of plant hormones, or becoming leaf tissue; it’s one reason the “plant hormone analogs” meaning the leaf-gall season begins in the spring! How- molecules are nothing like plant hor- ever, once the galls start growing they will mones but the plant’s response is the same continue to grow, even after the leaves as with plant hormones. The gall-forming fully expand. galls, midge galls, and so on. While it is process is usually initiated by the female Stem galls that arise from cambial ultimately important to know the when she injects gall-inducing chemicals tissue present a different scenario. Since gall-maker, gall identification usually into the plant along with her eggs. The cambial cells remain “free agents” starts with the gall itself. eggs themselves may ooze gall-inducing throughout the growing season, galls can We’ll provide a quick recap from Part chemicals and once the eggs hatch, the be formed from these cells anytime during I: The first step is to consider where the interaction continues with the immature the growing season, although most stem gall is found on the plant. Most identifica- gall-makers continuing to exude chem- galls also start growing early in the season tion resources use the following locations: icals to direct plant growth to suit their to provide ample time for the gall-maker leaf/petiole galls; flower/fruit galls; bud needs. to complete its development. galls; stems galls; and root galls. The resulting galls provide both a Next, galls are described based on protective home and nourishment for Insect and mite gall their structure such as general appear- the next generation of gall-maker. The identification ance (for example, ball-like, hairy, etc.), a continual direction of gall growth by the Identifying insect and mite galls is number of chambers, and the number gall-maker using chemicals to turn plant challenging because of the limited num- of gall-makers housed in the chambers. genes on and off speaks to why some find ber of accurate and updated gall identifi- Recall that unilocular galls have only one insect and mite plant galls so fascinating. cation resources, both online and printed. chamber; plurilocular galls have multi- The chemicals exuded by gall-mak- This is particularly true for galls found in ple chambers. Unilarval galls only have ers can only act upon meristematic plant North America; the European gall litera- one gall-maker per chamber; multilarval cells, such as the cambial cells mentioned ture is more robust. galls have more than one gall-maker per in Part 1 of this series or the meristemat- The existing gall identification re- chamber. The aforementioned ball-like ic cells in leaf buds; the precursors to leaf sources tend to follow the same general larger oak apple gall is a unilocular, uni- cells. Under the influence of chemicals outline. Most begin by separating galls larval gall. The hairy-looking mossy rose exuded by a gall-maker, the meristematic based on the gall-maker, such as wasp Continued on page 24 The continual direction of gall growth by the gall-maker using chemicals to turn plant genes on and off speaks to why some find insect and mite plant galls so fascinating. www.amerinursery.com | June 2015 | 23 Looking like a green apple, a large oak apple gall contains a wasp larva; once the larva pupates, the mature gall turns tannish brown and appears empty. Continued from page 23 The First Gall Law: Galls are abnormal plant gall produced under the direction of the gall wasp, Diplolepis rosae; a species first growths produced under the direction of a living described by Carl Linnaeus, is a plurilocu- lar, unilarval gall with many chambers but gall-maker; they do not arise spontaneously, nor only one larva per chamber. The ball-like hickory petiole galls are they in response to plant wounding that does produced by several phylloxeran spe- cies (Phylloxera spp.) are usually uniloc- ular, multilarval galls; there is a single not involve a gall-maker. chamber housing many of these aphid relatives. The First Gall Law: Galls are abnor- away plant hormone production; they are mal plant growths produced under the not considered “true galls.” Insect and mite gall laws direction of a living gall-maker; they do The Second Gall Law: Insect and mite While the highly organized plant not arise spontaneously, nor are they in galls are abnormal plant structures that galls produced under the direction of response to plant wounding that does not are composed entirely of plant tissue; many insect and mite gall-makers repre- involve a gall-maker. This law removes they’re not part of the gall-maker. Infec- sent a wide range of forms and locations, certain tree growth such as “burls” from tions by fungal plant pathogens can illus- there are certain consistencies that can be the gall arena. It is believed these abnor- trate that insect and mite plant galls are summarized as “gall laws.” mal plant growths are the result of run- indeed plant structures. The oak anthrac- nose fungus, Apiognomonia quercina, normally infects the leaf cells of its name- sake host producing blackened, necrotic A history of galls leaf tissue. The fungus can also infect the plant tissue of the small oak apple gall Plant galls are remarkable plant structures that have been observed, studied and utilized produced under the direction of the gall since antiquity. The Greek naturalist Theophrastus (372-286 B.C.) wrote about “gall-nuts” of Syria. wasp, Cynips clivorum, because the gall is The Roman naturalist Pliny the Elder (23-79 A.D.) wrote about medical uses of gall extracts.