Image from Steve Bowers and Worldtree
Tiny organisms which cut vegetative matter to feed the symbiotic "molds" they cultivate. Papercutters live in communities of hundreds to thousands of individuals. Since their discovery on the garden world of Macrystis, they and numerous variants have been spread across the Terragen Sphere to millions of worlds and orbitals.


Papercutters live communally underground or on flora, where they cultivate a symbiotic mold, which feeds on decaying vegetative matter, particularly materials high in cellulose. The papercutters feed on this mold and help it disperse. Papercutters are a diverse group, consisting of many species. After their discovery in 5393 on Macrystis, papercutters have spread to millions of habitats and worlds across the Terragen Sphere. A single papercutter can establish a new community because they can reproduce asexually. They can spread rapidly through cellulose-rich environments, whether as part of an engineered ecosystem or modified by malicious actors into pests that damage flora and paper products.

Papercutters are quads: members of the clade Quadrilateria, one of the main faunal groups on Macrystis. Quads developed four-way radial symmetry, which remains present in papercutters; they resemble four-legged starfish who walk on the ends of their legs. Papercutters also belong to the quad subgroup Parvocrucimorpha, which use a hydrostatic system for support and have lost most of their internal hard parts; most parvocrucimorphs are tiny (typically ranging from 0.1 — 1 mm) and often inhabit soil. Papercutters are large for parvocrucimorphs, being on average two or three millimeters in diameter, though they can range from just under a millimeter to nearly a centimeter in their native form.

Evolution and Ecology

Based on the behaviors of their closest relatives, papercutters evolved from highly social parvocrucimorphs that lived in burrows and mostly fed on underground tumuloforms: a large group which includes many sessile decomposers. They raised their young in burrows and fed them with semi-digested nourishment. Some parvocrucimorphs started to feed on sprinkle molds, a Macrystian group of filamentous decomposers, which then easily spread into their burrows. The sprinkle molds thrived among the parvocrucimorphs' young, where they too could benefit from the semi-digested leavings.

The development of a new biome type on Macrystis, the mangrove taiga, catalyzed papercutter evolution about 30 million years ago (based on time of genetic divergence). This was dominated by Umbraphyta, a new darkly pigmented type of vascular flora that grew into trees, rich in cellulose and with raised roots that trapped heat and moisture in their tangled understory. These were perfect conditions for parvocrucimorphs to live in, but they were unable to digest cellulose; sprinkle molds, on the other hand, thrive on cellulose. Thus, these parvocrucimorphs evolved into papercutters. They continued their foraging behaviors but now would harvest food that they couldn't digest themselves, cutting it up and ingesting it and then regurgitating it onto their sprinkle mold. The mold, in turn, provided a reliable food source.

Since then, their versatile lifestyle has allowed papercutters to spread into many other environments on Macrystis. Some have changed from underground burrows to living in nests on the surface or in the canopy. Some have even spread into much drier biomes. One strategy is to live in areas with seasonal rainfall (common on Macrystis) and to stay dormant throughout the dry season. Another is to live close to freshwater bodies or the water table and to forage away from them for periods short enough to avoid desiccation.

Papercutter species have variable levels of aggression. Some coexist peacefully and even merge their communities when they meet. Others can recognize relatives by scent and will attack members of other communities. At the extreme, papercutters will invade other communities.


Most papercutters are brown or tawny colors, sometimes shading into more orange hues to better camouflage with soil and Macrystian flora. They have poor vision, with a ring of compound eyes around their top side and a simple eye on each shoulder; all eyes lack color vision and have low resolution. However, papercutters have sensory hairs that they use to feel their environment and detect vibrations and scents, allowing them to navigate.

Papercutters have paired claws made from sclerotin at the end of each limb, useful for removing bits of flora, cutting them into bite-sized pieces, and bringing them into their mouth to later feed sprinkle molds. Papercutters have a blind gut with a single opening at the center of their underside that serves as both mouth and anus. Four radulae extend downward from the inside of the mouth, with a four-way shearing beak made of sclerotin on the outside, which the fleshy inner parts can scrape food against, allowing them to cut their food into small bits before ingestion. One papercutter subgroup has evolved additional sharp parts in the form of a spike on each shoulder, pointing upwards, for defense against predators.

They have a digestive pouch beyond the mouth in which they partially digest and store shavings of organic matter to later regurgitate onto their cultivated molds. They remain unable to digest cellulose on their own, so they rely on papercutter mold to process it into an edible form. While they primarily feed from their molds, they are still capable of digesting other cellulose-free material such as tumuloforms.

They have a closed circulatory system with blood vessels; their blood is clear, lacking a pigment, though some varieties also produce antifreeze. They have no dedicated respiratory system, instead breathing through their skin. As a result, they must live in moist environments.

The papercutter nervous system has expanded compared to most parvocrucimorphs. Like their relatives, it is a fairly distributed system, with clusters of nerves forming ganglia in the upper center of the body and toward the ends of each limb. Papercutters have a higher neuron count than their relatives, with a particularly expanded central ganglion with tens of thousands of neurons. This allows them to map and remember locations in their environment, enabling them to travel between their communities and food sources. They also create chemical trails to signal food sources to other community members. Each limb is also capable of some independent action, mainly reflexive cutting and feeding behavior. These abilities became relevant and useful during their redevelopment into biobots for maintaining artificial ecosystems within orbitals.

Life Cycle

The papercutter life cycle is typical of quads. The papercutter itself belongs to an asexual generation, known as a sporovite, or sporo for short. Sporos produce spores, which grow into sexual organisms known as gametovites or gams. Gams reproduce sexually, resulting in the birth of new sporos and completing the cycle.
In quads, the gams grow on the sporo's body; an individual papercutter grows four gams, one between each leg. It varies depending on the species, but it usually takes about four Macrystian days for a gam to grow (about fourteen Terran days). Gams are separate organisms, but their circulatory system is connected to their host's, as they have no other way to feed.

Male gams are small and resemble a nipple. They consist mainly of a testis and enough neurons to trigger the secretion of seminal fluid in response to tactile stimuli, which leaks through many tiny pores in a mass containing ciliate sperm.
Female gams are larger and are roughly thimble-shaped. They are mostly empty space, to serve as a womb, along with an ovary and neurons. They also fill with nutrient-heavy tissue to nourish their young. An opening at the front lets in sperm for fertilization when a male gam makes contact.

The sperm injects its nucleus into the egg and dies. An unfertilized egg can still be viable, however, as it goes through gene amplification, copying parts of its genome. Either way, a new papercutter grows within the gam. Usually, only one grows per gam at a time, though multiple births may be possible depending on the species.

Shortly after use, the gam's blood vessels disconnect from the sporo, and it is shed. This happens promptly with male gams. When possible, papercutters reflexively shed fertilized female gams among their mold symbiotes. The juvenile grows a while longer within the gam before cutting its way out.

By default, a gam develops into a female gam, but in the presence of a sufficient amount of pheromones from female gams, it will develop into a male gam instead. This ensures balance across the population and also means that an isolated papercutter will grow female gams and can reproduce asexually to found a new community. Because sporos have a high amount of gene copies, an individual papercutter has higher genetic diversity than most Terragen animals. This increases the chances of a viable population, though they still perform better when they have multiple founders.

Papercutter Mold

This symbiotic mold is a type of "sprinkle mold" belonging to the clade Filamentomycota. These grow as hyphae, analogous to Terragen fungi, though their cell walls are made of cellulose rather than chitin. These hyphae are long multinucleate cells.

Sprinkle molds grow in two forms, like most filamentomycetes. The mold morph consists of many long strands of hyphae, creating a mold with a striated growth pattern. They are versatile decomposers but specialize in dead matter heavy in cellulose. They come in a variety of colors, even within the same species, often bright hues from pigments that help resist toxins and environmental hazards.

Their other morph is the sprinkle morph, consisting of a smaller but still multinucleate hypha, resembling a sprinkle in its shape, at the edge of human vision. The mold grows sporangia, which produce spores, which then develop into sprinkles.

To attract fauna to spread their sprinkles, sprinkle molds produce an edible substance. In the case of papercutter mold, they produce a sticky filamentous mass, which papercutters can easily remove from the mold without damaging it. This mass serves as papercutters' primary food and is high in protein. It also has a high amount of sugars from flora that the mold had consumed. The rest of the mold is indigestible for papercutters.

When the papercutters feed on this cottony mass, sprinkles inevitably get stuck to their body. They'll transport sprinkles to other molds and to new locations. When two or more sprinkles of different mating types come into contact, they engage in conjugation, and a new mold grows from there. Having evolved from free-swimming cells, sprinkles have a limited ability to approach each other when nearby in a moist environment, whether via the sticky substance in the cotton-mass or through water.

Having evolved for easy digestion, the cottony mass is one of the safer things on Macrystis for nearbaseline humans to eat, with a rich sweet and savory flavor. Nonetheless, it contains proteins that are not used by Terragen life, so it can cause indigestion.

Outside Macrystis

Papercutters have turned out unusually adaptable and fertile on Terragen habitats and terraformed worlds beyond Macrystis. Even in their unmodified state, their reproductive system means that only one papercutter and a pair of sprinkles are necessary to start a new community. Furthermore, their molds can handle a wide variety of biochemistries, as they are decomposers, though they do require a sufficient amount of cellulose. As long as their environmental needs are met (moisture, oxygen, tolerable temperatures, etc.), they establish new communities readily. This easy upkeep makes it trivial to raise papercutter communities, which are widespread among those interested in Macrystian biology or who enjoy nurturing life or sharing a personal experience with others.

Papercutters and their molds have been subject to frequent genetic modification and are common throughout civilized space as biobots, which often comprise swarm technology. The most frequently used varieties of papercutter are grown/bred as biobots to maintain ecosystems within orbitals and make structures from various biomaterials, often cellulose or other polysaccharides. Swarms of mixed papercutter types began serving similar roles to many artificial bot swarms, as Savastian provolves began spreading a local ideology aligned with The Flow to nearby star systems among the Zoeific Biopolity. Most initial understanding and investigation of their genetics, life cycle, and habits originated from the first transapients to enter the Savastos system and survey Macrystis. Not long after the Caretakers catalogued the biosphere, and their agents released their discoveries, local modosophonts used papercutter sbmDNA (Six Base Macrystian DNA, a Macrystian DNA equivalent) to create templates for very effective biobots in local habitats and surrounding systems.

Reasons for gengineering may also be aesthetic, like creating more colorful papercutters or varieties that swarm in interesting patterns. Papercutter mold has been modified to produce proteins that can be consumed by any number of Terragen clades. Aioid bioborgs, particularly Kja Observers, sometimes include parts derived from modified papercutters; inversely, some papercutters have cybernetic parts incorporated to enhance their functionality. Another variation is to create much larger papercutters, generally drawing on genes from related quads to give them a vascularized breathing chamber and hemoglobin in their blood vessels. Some of these have become pets; one popular line takes advantage of their soft fluid-filled bodies and their sensory hairs to turn them into fluffy and responsive pillows.

Papercutters have been provolved several different ways; over time, sophont papercutters have increasingly led efforts into further provolution of additional non-sophont papercutter varieties. The earliest papercutter provolves were derived from one of the aforementioned enlarged types by giving them an expanded brain, making them sophont on an individual level. These had small populations in habs in the outer Savastos system and were initially designed by Zoeific modosophonts to incorporate their populations into The Flow.

The most influential papercutter provolution, however, happened when a provolved pterogam hive mind set out to create a similar hive mind using papercutters. They used genes from pterogams and relatives that enabled "linkage," in which gams are able to link together to connect their nervous systems, allowing them to share neural impulses. Through this, swarms of papercutters could achieve sophonce as a whole. In many cases, they also used direct neural interfaces to link the hive mind with a quicker response time, but this was not always the case.

In Zoeific habs, most populations of provolved papercutters coexisted with the same biobots they had derived from. Over time, groups developed feelings of kinship towards their automaton relatives, which fostered ideas of pan-sophontism throughout their cultures. The ruling transapients of the outer system took notice of this growing discontent and anxiety and so reached out to the local Caretakers to devise an amicable solution. Because papercutter hive minds required more living space due to their multiple bodies, the Caretakers agreed to allow colonization of the inner system but only by clades which were ultimately Macrystian in descent. Once this policy was enacted millions of papercutters and other provolved clades migrated sunward to establish colonies with the assistance of Zoeific neumanns. Over time dozens of minor polities would be established, with papercutters consistently comprising a plurality of the population.

As these provolve cultures developed, pan-sophontism lingered amongst the populace, and efforts were made to begin the provolution of papercutter mold as well. This was a great challenge, as the mold had no semblance of a nervous system, but they eventually engineered a subset of the molds' cells to have signaling capacity like neurons. They also enhanced their senses, mostly through cellular chemical detection, but they also created new cell types that extended into the air and could sense vibrations. The provolvers made it possible for papercutter colonies and symbiotic mold to connect their minds, whether through cybernetic interfacing or linkage by physical contact. In those communities, the mold could store memories for the group as a whole or provide communication networks, acting as biological DNIs and exocortices.

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Development Notes
Text by Banelord, Salty, TSSL, Martine, Worldtree, Schwefel Kamm, & DocViviLeandra
based on the original short article by Anders Sandberg, written 19 December 2001
Initially published on 02 February 2024.

Expanded 2020-2024