Friday, April 21, 2017

Pseudoflowers—Trick or Treat?

A flower-less rockcress.

Every spring some of our rockcresses forego flowering, and instead grow terminal clusters of fragrant yellow leaves dotted with sugary goo. But why? Generally plants produce color, fragrance and nectar to lure pollinators, which carry male gametes (pollen) off to where female gametes (ovules) await fertilization. But the yellow-leaved rockcresses have no flowers, no pollen, no ovules. And yet these plants are all about sex—fungal sex that is.

Rockcresses (Boechera spp.; formerly Arabis) are members of the mustard family. Most are perennials, with a few biennials. They usually produce white to pale pink or purple flowers, but the yellow-leaved versions are common enough to frequently confuse wild-flower enthusiasts.
“Almost every spring, someone brings me a picture or a plant of a strange little flower they’ve never seen before, and can’t key out or even begin to guess the family for.” Irene Shonle
“Strange little flower” (source).
Normal rockcress, and infected rockcress with pseudoflowers (Cano et al. 2013).

The yellowed rockcresses are infected with Puccinia monoica—mustard flower rust. Rust fungi are obligate plant pathogens, and include some of the most destructive agricultural pests (e.g. wheatstem rust, coffee rust). Some have extremely complex life cycles, involving five spore types and multiple host species in a single life cycle! (details here)

The life of the mustard flower rust is simpler, requiring three spore types and one or two hosts (full story here). If a wind-blown basidospore (which has a single haploid nucleus) is lucky enough to land on a suitable host plant, it germinates. Hyphae grow into the stem, tapping into the plant’s nutrient supply. But living happily ever after on a rockcress is not part of the rust's plan. Sex is its goal. Mustard flower rust is heterothallic, meaning opposite mating types are produced by separate “individuals” (rust infections) on separate rockcress plants. Opposite mating types need to get together somehow.

Puccinia monoica solves this problem by creating pseudoflowers. Like real flowers, they attract pollinators (mostly insects) by way of fragrance and the promise of sweet reward. How impressive that a simple little fungus has evolved to to grow such features! … except that’s not what happens, at least not directly. The real story is even more amazing. The plant grows these novel features … under the direction of the rust!
In addition to siphoning off nutrients, the rust reprograms the host plant, somehow changing which genes are expressed when. As a result, the infected rockcress never makes the transition from vegetative growth to flowering. Instead it elongates, grows extra leaves, and produces yellow pigment, fragrant compounds, sugary liquid, and wax. The resulting structure looks, smells and tastes enough like a flower that foraging insects show up, partake of a bit of sugar, and hopefully carry off the spore-like spermatia to receptive hypha on other rockcresses.
Bumps are spermagonia, which contain spores waiting to be dispersed and super-sweet liquid.
Pseudoflowers may mimic other wildflowers, like this nearby sagebrush buttercup (speculation for now).
With today’s molecular analysis techniques and model organisms (Arabidopsis thaliana, the thale cress, is a close relative of rockcresses), it’s possible to delve deeply into pseudoflower biology. In 2013, Liliana Cano and her colleagues looked at developmental changes in rockcresses infected with mustard flower rust. They found that for at least 31 genes, activity was significantly altered (enhanced or reduced), affecting leaf, stem and flower development; metabolism and transport of sugars and lipids; synthesis of volatiles (fragrant compounds); and wax production.

These changes can be interpreted as beneficial to the mustard flower rust. For example, consider wax production. Cano and colleagues suggest that the waxy leaves induced by rust infection serve to reduce water stress. Water-stressed plants often have shorter stems and fewer leaves—not what the rust needs. Perhaps the waxy leaves of infected plants allow taller leafier growth.
Gravelly soil drains rapidly, making for dry habitat. Looks like waxy leaves weren't enough to compensate.

Whatever the mechanisms, by enabling fungal sex, infection clearly benefits the rust. And the rockcress clearly suffers—no flowers, no sex. But what about pollinators? Are they beneficiaries or unsuspecting dupes? Some botanists consider pseudoflowers to be tricksters, luring insects into service with little reward. However in a 1998 paper, Robert Raguso and Bitty Roy pointed out that the super sweet liquid of rockcress pseudoflowers is popular with many kinds of insects, including bees, ants, butterflies and flies. And given how many sugar-oozing spermagonia there are on each yellow leaf, infected rockcresses may actually produce more yummy calories than uninfected plants. If so, then for pollinators, pseudoflowers are not a trick but a treat.
Foraging ant (in a hurry).

Puccinia monoica on Boechera sp. is the latest addition to my iNaturalist project, Plants of the Southern Laramie Mountains (two observations—one for the rust, one for the plant). To identify the rockcress to species, I have to wait until uninfected individuals are in fruit.
I found infected rockcresses scattered through this sagebrush grassland.
It’s still early spring at Blair (8000 feet elevation)—not much flower action.


Thanks to Elio Schaechter of Small Things Considered who recently blogged about Boechera pseudoflowers, which I’ve long ignored.

Caro, LM, et al. 2013. Major transcriptome reprogramming underlies floral mimicry induced by the rust fungus Puccinia monoica in Boechera stricta. PLoS ONE 8(9): e75293. (free).

Raguso, RA, and Roy, BA. 1998. ‘Floral’ scent production by Puccinia rust fungi that mimic flowers. Molecular Ecology (1998) 7, 1127-1136.

Wednesday, April 12, 2017

Wyoming Native Plant Society helps liberate Plant Names for Creative Re-use

What's your pleasure?

Let’s say you’re writing an article about a plant, or your local flora, or a pioneering botanist. Now … close your eyes and imagine you’re in a huge library dedicated exclusively to biodiversity, with 200,000+ holdings (many rare) scattered across the globe. Next, imagine giving the name of your plant or botanist to a “librarian” who then piles all relevant books, articles, field notes, correspondence, etc., on your desk almost instantaneously! In fact, this library is not imaginary. It’s quite real, though in a virtual kind of way. It’s the Biodiversity Heritage Libraryheadquartered at the Smithsonian Institution in Washington DC, but easily accessible from your office, home, or favorite coffee house.

I discovered the BHL in 2014, while putting together a post about the history of the lanceleaf cottonwood (Populus acuminata). BHL soon became my go-to site for information about botanical exploration of the American West. What I like most is the quick easy access to lots of useful information. Documents that were difficult to access or even unavailable only a few years ago are now just a search and a click away.
Per Axel Rydberg’s Populus acuminata. From American Black Cottonwoods, 1893; BHL.
Edwin James’s Jamesia. From Curtis’s Botanical Magazine, 1875; BHL.
Fossilized palm frond (Powell palmetto perhaps?) collected near Rock Springs, Wyoming. From JS Newberry’s The later extinct floras of North America, 1898; BHL.

The BHL is a consortium of natural history and botanical libraries that are digitizing legacy biodiversity literature, making it easily accessible as part of a global “biodiversity commons.” Much of this literature has been available only in select libraries, mainly in the developed world, making limited access a major obstacle—for example in research, conservation and education. Providing it online free-of-charge is a radical and exciting change. “Free global access to digital literature repatriates information about the earth’s species to all parts of the world.”

Put another way, BHL is making biodiversity literature “freely accessible to a global audience … thereby liberating taxonomic names and bibliographic data associated with the content for creative re-use.” Among the plant names most recently liberated were those in our very own Castilleja, the newsletter of the Wyoming Native Plant Society.
In October 1994, the Wyoming Native Plant Society newsletter was given a name: Castilleja.

It all started last October when the BHL blog featured a post titled A Local Focus: The Native Plant Societies of the US. When I read that native plant society newsletters were being added to the collection, I contacted Project Investigator Susan Fraser at The New York Botanical Garden, asking if Castilleja were part of the plan. Indeed it was. “We would be thrilled to include Castilleja in the project,” she replied.

Incorporating native plant society newsletters into the BHL is part of Expanding Access to Biodiversity Literature—a two-year project designed to “preserve and provide access to small natural history and botanical collections and publications.” It’s conducted by the New York Botanical Garden in partnership with Harvard University, the Missouri Botanical Garden, and the Smithsonian Institution Libraries.
“We are grateful to the native plant societies who have generously shared their local expertise by making their newsletters available to researchers through BHL. In addition to the biodiversity information they contain, these publications are a wonderful snapshot of the small, dedicated groups of people working all over the U.S. to document and preserve our native plants.” –Patrick Randall, Community Manager, Expanding Access to Biodiversity Literature; Ernst Mayr Library, Harvard University

Before Castilleja issues could be processed, a permissions form had to be signed (the society President took care of this). Fortunately, PDFs were available for all issues; these were transmitted en masse to BHL. Then the techies worked their magic. Now, whenever someone searches BHL for Boechera pusilla or Yermo xanthocephalus, for example, relevant issues of Castilleja appear on the results list. We’ve hit the big time!

The image below shows one result from a BHL search for “yermo xanthocephalus”—the desert yellowhead, endemic to Wyoming. As I scrolled through Castilleja Volume 17 Number 4 (1998), scientific names on each page appeared in the box on the lower left. Note that contents can be printed or downloaded (either the entire work or selected pages). I’ve used the latter option many times. Usually the pages arrive well within the hour, whether from the newsletter of a neighboring native plant society, or from a rare old book in a library thousands of miles away.

How did BHL manage to find yermo among the 51,749,439 pages held in the collection? It was magic!!! No, not really … sorry. But it’s just as cool as magic. As texts are processed, scientific names are extracted from each page using Global Names Recognition and Discovery (GNRD), a taxonomic name recognition algorithm. GNRD provides an open and global-names-based infrastructure to index, organize and manage biodiversity data. Like BHL, GNRD aims for easy public access, with the goal of spurring widespread and innovative use of biodiversity data. A noble goal indeed!

So if you’re in need of biodiversity literature, especially if it’s old or rare or otherwise difficult to access, pay a visit to the BHL. Adventure and discovery start here. And if you’re looking for a good time, browse the always-interesting BHL blog (warning: you'd better have plenty of time on your hands).
Above, Coffea arabica was the first coffee species to be cultivated, and still accounts for most of world's coffee production (from Köhler's Medizinal-Pflanzen; see The Berry that Changed the World). Below, Miss C.H. Lippincott Flower Seeds catalog cover (1900), from Leading Ladies in the World of Seeds (you can view over 11,000 seed and nursery catalogs in the BHL collection!).

Friday, April 7, 2017

Following a Tree, from Fronds to Rebar

April 7 has arrived, and being a tree-follower, I’m posting the latest news of my tree at the monthly gathering kindly hosted by The Squirrelbasket. If you like trees and would like to join us, check out the links above (it’s interesting and fun, with no obligation).

This year I’m following an extinct tree—Sabalites powellii, a palm that grew in southwest Wyoming 50 million years ago. I discovered it in December in our Geology Museum, and in May I will visit its ancient habitat. In the meantime, tree-following consists of learning about Sabalites and its world. This month, I decided to look into its wood. The Green River Formation—the rock layers where fossilized fronds have been found—also contains fossilized palm wood. It’s common, beautiful, and popular with collectors.
Fossilized palm wood is the state rock of Texas (source).
However, my search for “Sabalites wood” was futile, both in Google and the scientific literature. That’s because fossil palm wood is called Palmoxylon. Is this weird? Are paleobotanists exceptionally obsessed with publishing new names?
Sabalites powellii, a fossil palm frond “species” from southwest Wyoming.
Palmoxylon includes 200+ “species” of fossil palm wood (source).
Plant fossil names proliferate because paleobotanists are faced with an unfortunate situation. Most fossilized plants are actually fossilized plant parts—leaves, branches, flowers, etc., that fell off some plant. Generally there’s no way to know if different parts came from the same species, so they’re given different names.

Palm wood usually is easy to recognize. Cutting a petrified palm trunk cross-wise reveals a characteristic pattern of scattered dots—the vascular bundles that conduct water and nutrients up and down trees.
Cross-section through petrified Palmoxylon log (click on image to view dots; source).
Length-wise view of Palmoxylon (source). Xylon means “one having (such) wood—in generic names” (Merriam-Webster) … in this case, having palm wood.
In most trees (dicots), vascular bundles are arranged near the perimeter, but in monocots, they're scattered through the stem (diagram below). Monocots include orchids, grasses, lilies and more. Most are herbaceous, but a few produce wood and grow large, e.g. yuccas, bamboos and palms.
Cross-sections through dicot and monocot stems (source, modified).

Wait a minute!!! As botany students we're taught that monocots don’t make wood. Wood is produced by secondary growth in the vascular cambium (below), and monocots have no vascular cambium. Yet palms are monocots and obviously woody … ??
In dicots, wood develops from secondary growth via the vascular cambium (source, modified).
Large monocots are said to produce strong trunks through abnormal secondary growth. I think we call it “abnormal” because it’s disorderly and hard to categorize … and poorly understood. Webpage after webpage explains that palms thicken their trunks through diffuse secondary growth, in which parenchyma cells divide and enlarge. And yet a recent study revealed that at least some palms have a vascular-cambium-like meristem! (Botánico & Angyalossy 2013; article here):
“… we analysed palm stems of four species, with the aim to understand the possible presence of such secondary growth. We found that a meristematic band occurs between the cortex and the central cylinder and gives rise to new vascular bundles and parenchyma internally, producing parenchyma and fibres externally. … In fact, a meristematic band is present and may be more common than currently believed, but uneasy to detect in certain palms for being restricted to specific regions of their stems.”

Palms’ tough rigid tissue may not be “true wood,” but it’s called wood and functions as wood. Some kinds are used in construction, including that of the coconut palm. Wood from the outer part of the trunk, where the vascular bundles are dense, is harder even than oak and Douglas fir (source).

In fact, sometimes palm wood is better than true wood. It's more flexible—palms can bend 40º without breaking! The tough fibrous vascular bundles scattered through the trunk serve as plant rebar—like the steel rods that give strength and flexibility to reinforced concrete.
Flexibility is a great adaptation in hurricane-prone environments. Courtesy US Navy.
When a tropical cyclone tore through this rainforest, the palms were left standing (source).

We'll stop here. Once again, tree-following has taken me on a winding journey, this time to fossil palm wood, woody monocots, meristems, vascular bundles and finally rebar. We’re lucky there’s so much of interest in this world!

Thanks to Mike of CSMS Geology Post, for continued paleontological guidance.

Thursday, March 30, 2017

Of Microfungi, Music, Conservation & More

Remembering Martha Christensen.

In 1982, when I started as a grad student in botany at the University of Wyoming, my familiarity with fungi was limited. I knew of only two kinds: mushrooms and yucky crud (like athlete’s foot). But that’s not why I took Mycology. Few botany courses were offered that would fulfill my grad requirements, so when someone told me Mycology was “a really good class,” I signed up. It was a great choice, one that has benefitted me for 35 years.

By the time the semester ended, we had been immersed in the mind-boggling taxonomy of fungi, and exposed to the many critical roles fungi play in plant growth and ecosystem health. We experienced economic mycology: bread, beer (beer in class didn’t raise eyebrows then), cheese, tempeh and more. Especially intriguing to me was the discovery that soil microfungi in Wyoming basins include tundra species! These are Pleistocene periglacial relics, from times not so long ago (circa 10,000 years) when today’s short-grass prairies were permafrost and tundra.

But the best part of Mycology was the instructor, Dr. Martha Christensen. On the first day of class, we were taken in by her passion for the subject and her sense of humor. It was a semester filled with new knowledge, camaraderie, and lots of fun. I went on to take Martha’s Algae and Bryophytes, and was her teaching assistant for Man, Agriculture and Civilization, a non-majors course. I don't remember exactly what was taught in that course, but I’ve thought of it many times in the 30+ years since, because of one very useful piece of advice. We were frantically preparing for lab while the clock raced towards 1:10, when Martha noted with a smile: “This is just like fixing dinner for lots of guests: get as much done as you can, then quit worrying and enjoy it!”

The next year, when my advisor announced he wouldn't be available for my defense and seminar, Martha agreed to serve as committee chair. I’m not sure why I was so lucky, probably just timing, but it felt like Fortune had kindly intervened.

Those grad student days were the start of a long enjoyable friendship. But then last week Martha shocked everyone by passing away. She was 85, but still so full of enthusiasm and energy, and so busy with life, that death was unimaginable. The days since have been a time of reminiscing and sharing stories.

Martha had a long list of academic achievements. When I read her obituary and a recent tribute by one of her colleagues, I realized how little I knew about her career. That’s because our friendship was based on other mutual interests: outdoor recreation, conservation, and music. Martha’s interests were broad, and though an academic, she found time for them all. Sometimes it seems those days are gone. Is academic pressure now too great? Is there no longer time for the arts, public service, advocacy?

Martha played the viola, as do I. I love her story—told with her characteristic hearty chuckle—about how being a violist helped her land a professorship in the Botany Department. She was sure she was hired partly because several faculty members and their wives were fans of classical music and supporters of the University symphony orchestra—which was in desperate need of a good violist. Martha found herself sitting first chair (in a section of three!) and would play with the UWSO for thirty years. Though she loved classical music, and was especially fond of opera, she was no snob. Martha also played the saw!—entertaining us at departmental Christmas parties.

There was another side to Martha that we appreciated very much … and still do. She was a major advocate for public lands conservation in Wyoming, contributing both time and financial support. This is a tough place to be a conservationist; it’s easy to get burned out. But Martha persisted. That she loved nature and the out-of-doors was always clear. In retirement, she took great pleasure in visiting natural areas in the US and other parts of the world, always exploring, always learning. I looked forward to hearing of her latest adventures in our annual Christmas correspondence, for her contagious enthusiasm never disappeared!
Martha was one of the conservationists profiled in Ahead of their Time; Wyoming Voices for Wilderness.

A memorial service for Dr. Martha Christensen will be held on Saturday afternoon, April 1, 2017, at the Heritage Congregational Church, 3102 Prairie Road in Madison, Wisconsin. Family members will greet her friends at 1:00. A service of remembrance will begin at 2:00.

In September (date to be set), Martha's many colleagues, students and friends will gather in Laramie to celebrate her life. More information will be provided on the University of Wyoming Botany Department website.

Thursday, March 23, 2017

Black Hills Montane Grassland Infographic

This picture is worth 620 words! (Click on map and those below to view details.)

When our Black Hills Montane Grasslands paper appeared in the December 2016 issue of The Prairie Naturalist, I was really happy—happy to see it in print after years of work, and happy to see the map in color. I’m so glad South Dakota Game, Fish and Parks was willing to pay the extra $500. We badly needed color to drive home our message … and a clear strong message is the whole point of using an infographic.

But is this really an infographic? … or just a map?

According to the Oxford English Dictionary, an infographic is “a visual image such as a chart or diagram used to represent information or data.” The term came into use in the 1960s, originally as an adjective. Infographics include maps as well as charts, diagrams, networks and such. When a map emphasizes certain features (e.g. with symbols or colors) to show a situation, trend or pattern, it becomes an infographic.

Such maps (also called thematic maps) have been around far longer than the term “infographic.” The first that we know of was created in 1604 by Gerardus Mercator, to show religions of the world. He added symbols to various locations for the predominant religion. There was no need to refer to a list or table, because the viewer could see the situation at a glance (for example, the New World is filled with idol worshippers).
Designatio-Orbis-Christiani; David Rumsey Map Collection.
Part of legend, showing Mercator's choice of symbols.
A more powerful thematic map is one that Abraham Lincoln is said to have spent hours studying—Map Showing the Distribution of the Slave Population of the Southern States. Based on 1860 census data, counties were shaded according to the size of the slave population (darker – larger). A viewer sees immediately which parts of the region were heavily-dependent on slave labor.
Distribution of the Slave Population of the Southern States of the United States; source.
The Slave Population map played such an important role that it was included in the 1864 painting, First Reading of the Emancipation Proclamation of President Lincoln, by FB Carpenter (below, lower right; source).

In general, images are far more powerful than words. They grab our attention, and we process them “with alarming speed.” The brain can interpret a picture on the spot, whereas text must be processed linearly (more here). So if a message is amenable to being communicated with an image, that’s the way to go. A good infographic quickly shows a situation that would require many words to explain. As I worked on the grassland map, I found myself wishing I could skip the corresponding verbal explanation entirely. It would have saved me 620 words!

In creating the grassland thematic map, I had to figure out how to quickly and clearly communicate the message hidden in the massive amount of information we collected:
Black Hills Montane Grassland is a rare and endangered vegetation type endemic (limited) to the Black Hills of western South Dakota and northeastern Wyoming. Only eight mostly-native stands remain. All occur in an area of less than 35 square miles, on public land managed for multiple use. These should be managed as conservation sites.
We had the data to prove it. We thoroughly surveyed the high Limestone Plateau for Black Hills Montane Grasslands, finding 91. We evaluated and ranked each one based on size, vegetation quality (% native species), disturbances, and condition of the surrounding landscape. We found only eight A- or B-ranked grasslands. Only 10.5% of original habitat (based on presence of relic indicator species) now supports native Black Hills Montane Grassland.

After testing various symbol schemes on my colleagues, I settled on colored circles, one in the center of each grassland. Circle size indicates grassland size (three categories). Circle color indicates grassland quality (overall rank). The color scheme is intuitive: circles for the best grasslands are green, those for the worst are brown.
Only eight green and yellow grasslands remain! Fortunately, some are large.
Early on, I tried to come up with an effective black-and-white map to save money. But it was impossible to find symbols that stood out. Colors are much easier for the viewer to distinguish than shapes, which was the other choice for quality. Finally, color grabs the reader, especially in a black-and-white setting (ours is the only color figure in the issue). Fortunately, when our funding agency saw the color map, they quickly agreed color was the way to go.

I recently learned that our grassland thematic map is specifically a proportional symbol map—like Charles Joseph Minard's 1858 map about meat (livestock) sent to Paris butcheries from across France. Minard also used circles of various sizes, in this case to represent amounts of meat. The circles themselves are little infographics—pie charts, with colors representing types of meat.
Source includes much more about Minard’s famous and powerful infographics.

We dedicated our grassland paper to our late colleague and friend, Helen McGranahan (below, in white shirt). Her concern for Black Hills Montane Grasslands, her expertise in grassland biology and management, and her wonderful sense of humor are greatly missed!
Botanists at work in the Black Hills, 2011.

Sources (in addition to links in post)

Marriott, HJ, Faber-Langendoen, D, and Ode, DJ. 2016. Finding the best remaining Black Hills Montane Grasslands, the first step in conservation. The Prairie Naturalist 48:102-105. The paper will be available soon (and free) at The Prairie Naturalist archives. In the meantime, email me (see Contact tab), or download here (paste into browser bar): file:///Users/hollismarriott/current%20work/BHMG/BHMG2016/0PNATfinalrevision/BHMGMarriott%20102-105.pdf

Cartography and Visualization. PennState Department of Geography. If I had found this site before I started working on my map, it would have saved me a lot of time!

The power of pictures. How we can use images to promote and communicate science. James Balm, BioMed Central blog. (“We process images at an alarming speed.”)

Wednesday, March 15, 2017

Books from a Land of Rocks

All photo-poetry montages by Hollis (click to read). Thank you, poets, for your words.

The Grand Valley, in western Colorado, is a land bounded by rock. To the north, the Book Cliffs stretch for almost 200 miles—sinuous bands of muted sandstones and shales. To the south stands the steep face of the Uncompahgre Plateau—red, yellow, buff and orange sandstone walls, on a foundation of dark ancient rock. And in between—in the community of Fruita, just off the roundabout near the west end of town, on the second floor of the old bank building—eponymous Lithic Press makes words into books and sends them off, dispersing poetry across the Southern Rocky Mountains and Colorado Plateau (in this land of rock, we find ourselves using physiographic provinces rather than political subdivisions).

First off the press was Whacking the Punchline, a small square spiral-bound collection of compression sketches by Jack Mueller, published in 2008. Since then, 27 more books—many in custom format—have been ushered into the world. Lithic recently joined a national trend of small independent presses opening brick-and-mortar bookstores. They hung out their shingle in 2015.
Lithic’s shingle features predatory Xiphactinus—“probably the most photographed fossil specimen in the world” (click on photo to view the smaller fish inside; source).
Lithic Bookstore & Gallery—a space for design, sales, poetry readings, contemplation, inspiration.
Danny Rosen, at the office.
Last September, I stopped in Fruita to enjoy the local literary scene, as I do most years. This time I cornered Lithic-owner Danny Rosen in his office. “I have some questions for you” I said firmly, swinging a six-pack for emphasis. We sat down for an interview.

How did the name Lithic Press come about?
The name stems from my background in geology; the word means stone-like or pertaining to stone. Also, it alludes to early methods of printing. Lithography involved spreading a greasy kind of ink onto etched limestone to print from one medium to another.

What kinds of books does LP publish—what subjects, genres?
So far, Lithic has published books of poetry—largely because that has been my interest. I would like to publish other genres in the future, like natural history. More than any particular subject matter, I want to work with writers who pay attention to language, tell intriguing stories, teach me something.

What’s your policy on manuscript submission?
We do not accept unsolicited manuscripts; we accept manuscripts largely by invitation. I had an open submission period last year for chapbook manuscripts (simple books, folded and stapled, 20-40 pages long). We received over 100 submissions. It was time-consuming to go through all of them, and unpleasant to send out so many rejections. I don't anticipate doing that regularly. But we did get a few books out of it.

Though we don’t advertise for submissions, they show up anyway … frequently. Seems everyone who ever wrote a poem wants a book. Maybe that's a reflection of “celebrity culture.” Being a publisher offers interesting viewpoints on human psychology! Recently we've been reaching out to writers we admire, asking for manuscripts. This year and next we have books coming out by writers in North Carolina, South Carolina, Oregon, New York, California, and Bombay.

What is the Lithic bookmaking process?
I facilitate the process. One way or another I find manuscripts (or they find me). I don't edit so much as closely read each manuscript to find errors and give suggestions. The books I've done for Jack Mueller required a lot of work, a labor of love. He's a close friend, and this friendship played a big part in the creation of Lithic Press. I wanted to publish his work—I love it so much and I think it has great value. Amor Fati took four years to complete.

Most of our books start with manuscripts that are fairly polished when we receive them—that’s what I want. My strength is to make books for strong writers, not to help make writers strong. The early phases can be very exciting, when the idea first turns into a concept of an artifact. We decide what the book will look like: size, shape, kind of paper and binding type, font and font size, and so on. Kyle Harvey does all the design and layout for Lithic. He is very talented and imaginative. I nudge him continually, to be open to what possibilities we may conjure: new kinds of paper, different presentations.
The making of a book presents possibilities at every juncture, on every page, like Jack Mueller’s The Gate, which we made as both an inexpensive chapbook and as a limited edition, cloth-bound, foil-stamped hardcover. For Kierstin Bridger's Demimonde, a book of poems written from the point of view of prostitutes in frontier mining towns in Colorado, we paired antique images with each poem and printed them on paper vellum pages.

I want every book to stand alone as a fine artifact—a handsome object that feels good in the hand, that makes a contribution to the human story—a little something to spread around, leave behind.
I’ve been fascinated with books all my life. As a kid, I was blown away by “All the Books in the Library.” To make something that will go into that library feels like important work. Someone said to me recently that I was like a midwife to the book. That’s a strange but accurate description. It’s awesome to hold a newborn book and usher it into the world.
Going Down Grand, Lithic’s pocket-sized book of poems for your next trip down.

Yet stones have stood for a thousand years, and pained thoughts found 
The honey of peace in old poems.
–Robinson Jeffers