Galls 101, https://uwm.edu/field-station/bug-of-the-week/galls-i/, provided an introduction to the biology of galls and gall-makers.  Short version: “Better living through chemistry.”  This week’s BOTW features a few oak galls and a grape gall.  Remember, of the 2,000-plus kinds of galls found on North American plants, 800 different kinds form on oaks (genus Quercus). 

Cynipid wasps, which mainly target stems and leaves, are very big players in the oak gall game.  The galls caused by some Cynipid wasps are very high in tannin/gallotannin (giving them the bitter taste that gave rise to their name, gall).  Tannic acid is produced routinely by many plants (the plant’s strategy is to make itself unpalatable to grazers), but galls contain the highest tannin concentrations on most plants.  It has been suggested that the gall-makers enjoy some “tannin-perks;” since tannins are somewhat anti-microbial, high-tannin galls may protect the larva against fungi and bacteria.

Interesting as they are to Nature Appreciators and to Scientists (they are, after all, “tumors” and their dynamic in that area is being studied), galls have a history of human commerce and use that goes back thousands of years.  They have provided food, medicine, lamp fuel (in Greece, from a gall caused by a wasp named Cynips theophrastea), chemicals for tanning hides, dyes for fabric, leather and hair, beads for necklaces, and inks for tattooing and writing. 

Aleppo galls (produced by Cynips gallae-tinctoriae on certain Turkish/Eastern Bloc oaks) have the highest concentrations of tannin among the galls, 50 – 65%.  Historically, Aleppo galls provided a strong astringent and a treatment for fevers, burns, mouth ulcers and toothache.  They continue to be important trade item, now used more for tanning and dyeing and as an ingredient in inks. 

The presence of traces of iron-gall ink in the Dead Sea Scrolls makes for a pretty impressive pedigree.  S.W. Frost, in Insect Life and Natural History, wrote in 1942 that “in some places, the law requires that permanent records be made with ink derived from gallnuts……The Aleppo gall has been specified in formulas for inks used by the US Treasury, Bank of England, German Chancellery, and the Danish Government.”  The downside of iron-gall ink is the fact that it tends to fade after, oh, 1,500 years or so, and by that time, it may have discolored your paper, too.  Google “oak gall ink” or “iron-gall ink” for the recipe; you can join the artists who explore older media – and the forgers of old documents – in reviving this ancient ink.

Oak-Apple Gall – There are about 100 kinds of these marble-to-ping-pong-sized galls; they grow on a leaf’s mid-vein or on the leaf stem (petiole).  Some kinds have thin outer shells with fibrous insides, and others are denser.  The oak apple pictured is occupied by a single larva.  In fall, when you find these on the ground among the fallen oak leaves, check to see if there is a tiny exit hole and think about the size of the full-grown wasp that made it.  Stokes, in Nature in Winter (great section on galls), tells of opening an oak apple gall that held about 200 ant eggs along with their nursery workers, and he has also found nests of mud wasps inside.

Oak Bullet Gall – Because they form from the woody tissue of the twig, these half-inch galls are very firm and can stay on a tree for several years.  There are about 50 kinds of oak bullet galls, and some secrete a sticky “honeydew” that attracts other Hymenopterans- ants, bees and wasps.  One source said that in exchange for the honeydew, the honeydew-eaters discourage parasitic wasps from laying their eggs in the galls.  But, another source mentions attacks on bullet galls by parasitic wasps that insert their ovipositers into the gall and lay their egg on the gall-maker’s larva.  Birds may peck open the gall and go after the larva.

Woolly Oak Leaf Gall – These attach to the mid vein (usually) or side veins (sometimes) and they look like cottonballs.  They grow on the underside of the leaf, and they are easier to see as the leaves fall.  Based on this one sample, it looks like they may have a “vampire-like” effect on some surrounding tissue.

Grape Phylloxera Gall – In 1850, there was only one species of grape being grown in all the vineyards of Europe.  In about 1860, the Grape Phylloxera (a wingless aphid about 1/20” long) was accidentally introduced from its native North America.  The rest, as they say, is history.  By 1880, the little critter had traveled to Australia, Algeria, South Africa, and via a different route, California.  One-third of French wine-producing grapes, about 2 ½ million acres, were wiped out (Mother Nature usually finds a way to deal with monocultures). 

While leaf galls seldom damage a plant, a plant with grape phylloxera leaf galls has root galls, too, and the root galls weaken and stunt the vine.  The French fought back, and after burying live toads under the vines to draw out the poison failed to work (True!), they imported the rootstocks of resistant American Fox grapes both to graft the French vines onto and to develop hybrids from – all the while “dissing” the quality of the American grapes.  Each of the leaf galls may house a teeny, yellow Aphid Mom and hundreds of eggs and/or nymphs.

Despite the galls and the withering, the leaf continues to function.

And by the way, it really bugs/galls the BugLady that much of the information about these generally harmless growths is found on sites that have a pest control and forestry-pest bias.  The accounts invariably end with some variation of “These are harmless and don’t measurably damage the plant, but you don’t like the looks of them, so here are some chemicals you could throw at them.”

This concludes Galls I – How They Do That, and Galls II – A Date with History.  Coming eventually, Galls III – Oddball Galls.

Go outside – look for galls!ody in the past five days.

Kate Redmond, The BugLady

Bug of the Week archives:
http://uwm.edu/field-station/category/bug-of-the-week/

2016 – When the BugLady started researching zebra mussels, it became apparent that the story of this non-native, invasive mussel was inextricably entwined with that of an equally-alien and equally-invasive mussel, the quagga mussel.  And also that we are, as a species, appallingly slow learners. 

Zebra and quagga mussels are in the Phylum Mollusca, a diverse bunch that includes snails and slugs, limpets, clams, scallops, squid, octopi, and cuttlefish.  Within the Mollusks, they’re in the order Bivalvia, and in the family Dreissenidae. 

Zebra mussels (Dreissena polymorpha) and Quagga mussels (Dreissena bugensis) have traveled far from their native haunts, the Caspian Sea drainage of western Russia, but they have settled nicely into their new homes in America, hitchhiking through the Great Lakes and inland lakes and rivers on boats, boots, bait buckets, and on the feet of waterfowl.  Zebra mussels were first identified in the US in 1988 in Lake St. Clair, just east of Detroit, and they reached Wisconsin by 1992.  In 2010, they were found in 130 Wisconsin lakes and rivers. 

The first North American quagga mussel was found in 1989, but it wasn’t positively identified until 1991.  The two mussels are now found in 29 states, and the National Wildlife Federation website estimates that the Great Lakes alone are home to 10 trillion (with a “T”) zebra and quagga mussels.  They are similarly-marked, but the underside of a Zebra mussel shell is flattened, and a Quagga mussel’s shell is more rounded and doesn’t lie flat.  Pictures of the two may be found at: https://www.invasivespeciescentre.ca/invasive-species/meet-the-species/fish-and-invertebrates/zebra-and-quagga-mussels/.  Here an animated map of the Zebra mussel’s spread between 1986 and 2024: https://nas.er.usgs.gov/queries/SpeciesAnimatedMap.aspx?speciesID=5 (the BugLady loves animated maps), and graphics showing the spread of both species in the Lake Michigan basin up until 2015 (scroll down to the Lake Michigan profiles) https://greatlakesecho.org/2019/05/30/invasive-mussels-challenge-commercial-whitefish-fishing-in-the-great-lakes/. 

Did they hoof it over here on their own?  They did not; like most of us, they came over on the boat.  They undoubtedly arrived in the Great Lakes in the ballast water of ships that ply international waters, as have a rogue’s gallery of hardy gatecrashers (more than 180 species, so far) – a number of North American organisms have been toted to Europe in the same fashion. 

Here’s the physics of it: while they’re in their home ports, European vessels take water (plus whatever is swimming in that water) into tanks built on the inside of the ship’s hull, and this “ballast water” helps keep the ship upright.  A ship carrying a small cargo needs lots of ballast, but as it loads more cargo, it discharges ballast water (plus whatever’s swimming in it) https://en.wikipedia.org/wiki/Ballast_water_discharge_and_the_environment#/media/File:Ballast_water_en.svg.  Ocean-going ships routinely enter the Great Lakes via the St. Lawrence Seaway, and equally routinely, have emptied their ballast water into the Great Lakes.  In 1993, a difficult-to-enforce law was passed that required incoming ships to replace their home-grown ballast water with ocean water before entering the Seaway,

In 2011, New York State, gatekeeper for the St Lawrence Seaway, proposed stiff, new regulations about ballast water management/treatment.  Three (short-sighted) Midwestern governors pushed back, citing concerns about job loss, even though some innovative alternative transport was suggested at the time. 

Zebra and quagga mussels turned out to be plenty adaptable – although they originated in salt/brackish water, they quickly adjusted to fresh.  They are bottom dwellers that live in clusters in great, huge, astronomical numbers on the floors of lakes (at one site in Arizona, quagga mussels number 35,000 per square meter).

When quagga mussels arrived, they out-competed the zebra mussels.  Although their life histories are similar, the two mussels prefer different habitats.  Zebra mussels like water depths of 6 to 30 feet, and quaggas can live as deep as 400 feet, so zebra mussels grow closer to shore, and quaggas thrive through the deep basins of the Great Lakes.  Quaggas necessarily have a much wider temperature tolerance.  Zebra mussels prefer hard surfaces to grow on, but Quaggas thrive on softer, siltier lake floors.  Both species eat all day, but quaggas continue feeding during the winter, when zebra mussels are dormant (you snooze, you lose).  The BugLady’s photos show them as most people experience them – as “empties,” cast up on the beach. 

Mussels are “filter feeders,” which means that they suck water in through a siphon and run it over their gills.  Food particles, zooplankton, phytoplankton, and nutrients (and pollutants) are strained out of the water by cilia in the gills and are moved to the mussel’s mouth, and the water is expelled through a second siphon.  Wastes are released as mucous-covered, organic packets called pseudofeces (vocabulary word of the day).  An inch-long zebra mussel can filter a liter of water a day.  One liter per day x Biblical numbers of mussels = large bodies of very clean water. 

At first, some people were thrilled – “Yay, the lake is clean again,” shouted the headlines.  Cities around Lake Erie had been battling pollution in the form of algal blooms due to excessive nutrients (fertilizer) in the water.  In short order, you could see the bottom of the lake again (it’s called “nutrient bioextraction,” and it can be a useful tool in controlled situations where the bivalves are removed when they’re finished eating and processed into animal food or, ironically, fertilizer). 

However, the water was crystal clear because there were so few nutrients left in it, and native species that depended on the food in those liters of water were out of luck.  It was an all-out attack on the base of the food web.  Zooplankton feed on phytoplankton and are fed upon by larger animals, including tiny fish, which, in turn, feed bigger fish and a variety of other vertebrates.  But nutrient theft is not the only problem with these mussels. 

• Quaggas eat algae, but they’re picky, and non-toxic algae are their favorites.  What’s left after they feed is higher concentrations of the more troublesome algae. 

• Light is able to penetrate deeper into that nice, clean water – UV rays are bad for the very young fish but great for plants, opening the door for more algal blooms.  Decaying algae, some carrying harmful bacteria, wash up on beaches or lurk just offshore.

• The clear water can allow thick growths of other aquatic plants, too, fertilized by nutrient-rich mussel poop.  Dense aquatic vegetation discourages swimming, fishing, and boating. 

• Our native shellfish are indicators of the health of their environment.  The invasive mussels turn lake beds hard and lumpy, with wall-to-wall shells, making it hard for native bivalves to find favorable habitat.  To add insult to injury, zebra mussels will piggyback on native mussels, hindering their feeding and ultimately smothering them.  Great picture at: http://www.startribune.com/mussel-bound-lakes-could-imperil-birds/133021828/.  Said one fisheries biologist, “When I’m diving in the Mississippi River, if I come up with a ball of zebra mussels, I know that when I break that open, I’m either going to have a snail or a mussel — a native clam — inside that ball of zebra mussels.”

• Old zebra and quagga mussel shells wash up on shore, often in sharp fragments, problematic for barefoot beach-goers.  

• Zebra mussels overgrow anything that stands still long enough, especially pilings and other underwater surfaces, and they clog utility water intake/cooling pipes, requiring costly fixes.  Researchers in a few northern lakes have observed an odd (and one-sided) association – zebra mussels growing on the backs of clubtail dragonfly naiads (immature clubtails may live underwater for several years, giving mussels plenty of time to gain a foothold).  Their exoskeletons are effectively glued shut by mussel filaments on the thorax, where the exoskeleton normally splits to release the adult dragonfly, so naiads crawl up on shore and die there, unable to emerge.

• As they feed, quagga and zebra mussels accumulate toxins, with some pollutants occurring in their tissues (and their pseudofeces) in concentrations measuring many thousands of times higher than in the surrounding water.  Those toxins (including Clostridium botulinum) get passed up the food chain in a process called biomagnification.

• A mass of pseudofeces on the lake’s floor requires oxygen in order to decompose. 

Lake Superior has mostly avoided this mess, probably due to a combination of its much colder temperatures, lower levels of nutrients in the water, and its water chemistry – very little calcium for growing strong shells. 

Mussel reproduction is external and chancy.  Males and females release their bodily fluids into the water, nature takes its course (aided by water currents and propinquity), and fertilized eggs hatch into a life stage called veligers.  An adult female can produce as many as a million eggs annually, and her life span is three to five years, but the attrition rate for eggs and veligers is huge (they’re even eaten by filter-feeding adults).  Mom and Dad may be stuck in one spot, but their offspring are, temporarily, free-swimming, and currents can carry them great distances.  Veligers swim and feed for four or five weeks before they must attach, and they mature by their first birthday. 

What slows these critters down?

• Fish, like yellow perch and redear sunfish, and waterfowl, especially diving ducks like goldeneye and scaup, have learned to love the invasive mussels (98% of a Lesser Scaup’s diet is zebra mussels).  Kudos also go to the equally alien and equally invasive quagga-eating Round goby fish.  Alien species that become invasive do so because they have left their native predators behind.  In this case, the predator caught up with the mussel, but, alas, this aggressive little fish damages native fish populations, too.

• A patented bacterium called Zequanox targets these two mussel species only and has a 90% mortality rate, but it’s far too expensive to apply to a Great Lake.  There’s a copper-based treatment, too.

• Unusually warm water – In 2001, the water temperature in parts of the Upper Mississippi reached 89 degrees F, and masses of zebra mussels died. 

Good news-Bad news: For zebra mussels in the Great Lakes, the show is over, but they’ve simply been replaced by quaggas, and scientists doubt that the Great Lakes will ever return to their pre-alien-mussel state.  At this point, Lake Michigan (the 6^th-largest freshwater lake in the world) is essentially a man-made ecosystem that’s being managed as a fishery, because the base of the food web is so messed up.   

For those people whose attitude toward alien species is “Get over it – A species is a species!  New species = more biodiversity,” the BugLady has one word. “Seriously???”

For all your invasive species needs, remember our own Southeastern Wisconsin Invasive Species Consortium (SEWISC), https://sewisc.org/, a wealth of information about invasive species already in the state and on the horizon.  

Be assured that the BugLady did not use any of the information presented in the article about “Zebra Muscles” (ain’t Spellcheck grand?). 

The BugLady

---

2025   So – what’s new?

ZEBRA MUSSELS

• They continue to spread to small, inland lakes in Wisconsin – 250 of them, at latest count https://apps.dnr.wi.gov/lakes/invasives/AISLists.aspx?species=ZM, with 14% of the 13,000 untouched lakes judged to have a favorable water chemistry to support them.  Here’s a fact sheet about them https://nas.er.usgs.gov/queries/FactSheet.aspx?speciesID=5.

• An ounce of prevention is worth a pound of cure – it can be time-consuming and expensive to try to get rid of them once they’re established (and they’ve usually been around for two or three years by the time anyone notices them).  Most boat launches have signage about hosing off the trailer, boat, live wells, bilges, and motor, but the list should also include swimsuits and wetsuits – the veligers can live for three to five days out of water.

• Researchers in Minnesota drew a direct line from large Zebra mussel infestations in inland lakes to some staggering increases in mercury levels in game fish in those lakes (up 72% in walleyes and a whopping 157% for yellow perch!).  In the waters well-filtered by invasive mussels, walleye fry fail to thrive.

• Sheephead, pumpkin seed sunfish, and carp will eat Zebra mussels, but apparently, non-native mussels are less nutritious than the native mussels, and the fish are stunted (the BugLady is blown away that someone figures these things out).

QUAGGA MUSSELS

• The belief in Lake Superior’s resistance to Quagga and zebra mussels turned out to be wishful thinking, but the populations seem localized – Apostle Islands, Isle Royale, and a few harbors.  Water that averages 40 degrees does seem to discourage them.  

• Here are some Quaggas feeding – https://www.seagrant.wisc.edu/videos/quagga-mussels-feeding-sped-up-10x/.

• The population of Whitefish has plummeted by 80% in some parts of the Great Lakes, due to Quagga mussels.  There are estimates of quadrillions of Quaggas in the lower Great Lakes.

Quaggas may have completed their conquest of Lake Michigan, but their spread into our inland lakes is just starting.  They were recently found in Geneva Lake, a deep lake whose substrate is 95% sand.  In a survey they did at Geneva Lake, the DNR found that a quarter of the boats had been used on a different water body in the past five days.

Kate Redmond, The BugLady

Bug of the Week archives:
http://uwm.edu/field-station/category/bug-of-the-week/