Shaggy Volleypom

The Shaggy Volleypom split from its ancestor the Obsidoak and quadrupled in size to dominate the canopy. Reaching up to 200 meters tall, and bases of up to 18 meters across in truly ancient individuals, these behemoths of the forest are quite a remarkable sight. Supported by the massive trunk are similarly robust branches thick with dark leaves and sporangium.

The leaves of the Shaggy Volleypom have become more complex organs, with methods of dealing with heat. The supportive tracheal veins developed by it's distant ancestor, the Obsiditree, for the purpose of increasing CO2 access to the inner leaf, has gained greater branching directly into the surrounding spongy mesophyll. The entries to the tracheal system of each leaf can be found as a single pneumathode at the end of each major tracheal vein, across the edges and tip of the leaf. Each leafs tracheal system is isolated to that single leaf. There is no palisade mesophyll in the leaves, the task of photosynthesis is performed by cells throughout the spongy mesophyll. Like in it's ancestors the majority of vascular fiber and phloem in the leaf concentrates around the main tracheal veins, as the greatest rate of material exchange occurs in those regions of the mesophyll.

The movement of water is dictated, as it is throughout it's lineage leading all the way back to the Orange Spore Stalk, by guttation through its epidermis and evaporation across its surface. In the heat of the summer sunlight, far from their water source below, the rate of evaporation can become greater than it's vascular system can compensated for, so the Shaggy Volleypom has developed a couple solutions. The first solution is triggered when light intensity becomes too great for its tissues to handle, and it starts to heat up. This solution is to begin development of long pale trichomes, or hairs, that rise above the leaf and throw a shadow across its surface. It's a simple, but very effective, solution, and is triggered easily enough during springs and summers that the dense branches will be highlighted in a spotty manner with gray and white from the hairs reflecting and breaking up the pathway of light moving across them.

The second action for tissue preservation is triggered by rapid evaporation, this can be due to excess heat from surroundings, light heating the leaf, or just more wind than usual that year dragging across the leaves and stripping them of moisture. This second solution is the excretion of a thin layer of PHB, a bioplastic, across both the upper and lower surfaces of their leaves. This substance is not evenly coated, and cracks are typically left to allow for some degree of evaporation to continue. When this action is taken the leaves with it take on a glossy appearance, and generally the plastic remains for the duration of the leaf.

The pneumathode, or entrance to the tracheal system, remains fairly simple in structure, as a pore that passes through the epidermis and tracheal epithelium to allow direct gas exchange between the inside of the leaf and the outside world. However, the epidermis and tracheal epithelium do not directly join, there is a crevice that leads to the mesophyll which will expand open when in contact with water. The entrance of the pneumathode is shaped in a manner to lead rain droplets into itself, where they will be sucked up into the mesophyll through this crevice. Shortly behind this crevice as well are new tiny packets of tissue called rain bulbs, what they do is produce terpenes which seep out of the pneumathode and create aerosols in the air. Enough of this stuff will encourage the formation of rain clouds in the area over time.

Though the Shaggy Volleypom is technically "evergreen", like other black flora it takes advantage of it's pigmentation to warm up during the winter, it still has morphological differences between seasons. The summer leaves contain well developed tissue between each supportive vein, taking full advantage of the summer sun. As the year progresses these broad leaves are shed and replaced by narrower, more filamentous leaves. These leaves comprise mainly of support veins wrapped by a thin layer of mesophyll and vascularization. Their main purpose is prevention of snow build up on the more delicate portions of the branches, as the massive sudden weight addition would snap them. The leaves achieve this by simply shedding snow due to their structure not providing surface to hold it. Warming during the day also facilitates this process, and provides a little water for the leaf to absorb.

The majority of leaves can be found on young non-woody twigs and the continued growth of leaf spurs on newly woody twigs and young branches. The growth of a leaf begins with the continuation or creation of a leaf spur by the epidermis. A leaf spur is a slow growing nub of tissue that sits as a part of the epidermis, so that as the parachyma underneath grow and expands the leaf spur simply gets pushed outward. When bark develops around it the outer end of the leaf spur remains softer, photosynthetic, epidermis. However in time the bark will overtake it as the layer expands.

Off of the end of the leaf spur grows a tough and simple microphyll. This dark structure is dense with photosynthic cells that can be considered proto-spongy mesophyll. Once the microphyll is formed and functional the spur, if new, will stimulate the formation of vascular fiber offshoots from its supporting twig to grow and infiltrate it and the microphyll. The leaf spur will contain this vascular fiber for the rest of its time on the Shaggy Volleypom. Inside the spur the vascular fiber will develop normally, however in the microphyll it undergoes a different change.

The microphyll extends significantly in length, increasing it's available surface area. The vascular fiber inside expands a single chamber to increase in diameter and form a central support vein, like in the ancestral Obsidishrub, though with the more complex functionality of increased gas absorption like in it's ancestral Obsidian Tree with the formation of its primary pneumathode. The walls of the growing tracheal vein increase in thickness, multiplying the number of cells across the wall, while the cells themselves increase the rigidity of their cell walls with modest additions of lignin and other fibers. The length of the tracheal vein, and vascular fibers, continues to grow at this point and the leaf continues to grow with it.

The vascular fibers at this point begin separating from the tracheal vein, as it becomes a distinct structure and tissue. The cells along the dorsal and ventral sides of the tracheal vein remain steady in their number and growth, but the cells in the remaining two sides begin growing rapidly, once the vascular fibers break away, to start forming secondary tracheal veins. This growth pushes the tissue of the leaf outward and stimulates increased growth of mesophyll, in winter leaves this increased growth does not happen. Eventually the secondary veins mature and the leaf fully forms.

When the life of a leaf is over, the point of connection between it and the leaf spur is severed via apoptosis. The leaf falls away to join the forest litter beneath it, returning nutrients to the ground to be recycled. The leaf spur remains alive and well, rather than forming a scar at the point of severing it sheds any remaining dead tissue and fills in with fresh epidermis, increasing in length ever so slightly by a millimeter or two. Once completely repaired it proceeds to grow a new microphyll to repeat the process of creating a leaf.

The reproductive strategy of the Shaggy Volleypom has shifted from the usual method taken by most black flora. Most black flora release sexual or asexual spores into the air in huge plumes, with spores of those descending from the Salty Sunstalk combining in the air directly before falling to the ground to develop. The Shaggy Volleypom has differentiated it's sporangium, it has begun producing megaspores in megasporangium and microspores, which physiologically are more like traditional black flora spores, in microsporangium.

The megasporangium is a round trilobed structure with a diameter about 10-15cm at maturity, these grow in clusters ranging from 30cm to 90cm across. Their inside are densely packed with oval megaspores 1-2cm long which are arranged similarly to an Earth pomegranate. The three lobes are divided by crevices, ancestrally these are the points where a sporangium opens, but in the megasporangium These remain about 3mm in breadth and act as receiving channels for microspores. The clusters of megasporangium, once their cargo inside is mature, fall to the ground. The walls of the megasporangium are tough but papery, deforming on impact and scattering the thicker walled megaspores as they bounce and roll across the forest floor.

The microsporangium is a trilobed oblong structure 3-5cm long. These grow in large tight clusters ranging in size from 10cm to 120cm across. They function similarly to the ancestral sporangiums of their lineage, pumping out large amounts of orange spores into the air as they curl open like more typical black flora sporangiums. These microspores float about until they drift into the receiving channel of a megasporangium, at which point they fall in and stick to the inner surface or top of a megaspore. Once stuck to a megaspore the two will merge become a fertile megaspore, at which point the outer layer of the spore becomes a thickened wall of cellulose and lignin as armor to keep safe during falling.

Both structures are grown throughout the year, with megasporangium taking up to two years to fully mature. The scattered megaspores require either a stratification period of about 60 days minimum to break dormancy, or to remain in dormancy for two years without stratification before growing. The hardened shells meant to protect from falls perform a secondary function by preventing ease of consumption by fauna that may try to eat them, though persistence will still break them. They are commonly stashed away by creatures that horde food for storage.

Reduced to 18% of original (2315 x 3801) user posted image Simplified illustrations of the formation of microspores (top), and megaspores(bottom), with gamete positions indicated via circles.

The core components of most sporangium are fairly basic, the wall, the supportive tapetum, and the productive "mother cell layer" or materchyma with all three arising roughly at the same time upon differentiation of the parent cell clump of parachyma. The walls of both the microsporangium and megasporangium of the Volleypom are fairly conservative to black flora. These walls are several cells thick and heavily photosynthetic, they are very water permeable and guttation occurs freely across them when immature and growing, an inheritance from the tiny ancestral stalks in which these structures were their highest point. Vascular tissue, such as vascular fibers and phloem, directly pass into this layer separate from the inner developing parts. This layer can be distinctly peeled away from the other parts if care is taken. The role of the wall is essentially to act as a barrier, separating the developing inner tissue from the outside world.

Tapetum makes up any inner tissue of the sporangium that does not directly give rise to the gamete cargo of a spore (or zygote cargo, in the asexual spores of other species). There are several different categories of tapetum, from tissue that seeps up nutrients and water from what little vascular tissue that reaches the inside of a sporangium, to tissues that surround and support the materchyma, to surface tapetum that build the spores themselves. Though these different functions occur in specific places with specific versions of tapetum cell, the different layers are not distinctly separated.

The materchyma, the "mother cell layer", is a strip of cells across the inner sporangium deeply imbedded in the tapetum. The two faces I this strip are distinct, with a flat face toward the wall comprised of young cells, and toward the center, where spores mature, the face is jagged with many thin tiny peaks of older cells reaching into the tapetum. These peaks break apart into individual cells that are pushed forward by a layer of tapetum growing alongside them.

As these individual parent cells drift away from the materchyma they begin to replicate, performing mitosis they cleave perpendicular to their path of motion, so one daughter cell is ahead of another. After this tapetum signaling induces meiotic cleavage, again perpendicular to their path of motion, first in the cell ahead and then in the one behind as it travels forward. These resultant germ cells travel ahead, single file while the tapetum feeds them and bulks them up. The tapetum surrounding the forwardmost germ cell begins to centralize a more distinct form around it, this form maintaining connection to those tapetum cells ahead and behind but separating from ones next to it. Nutrients passes into the structure from cells behind it, and through the structure to cells ahead of it.Among microspores, and most black flora, this structure continues moving forward and growing, eventually breaking free entirely and resting on the inner surface of the sporangium awaiting its release. Megaspores, on the other hand, stop moving at this point. The tapetum beneath them ceases its continual push forward, trapping the germ cells next in line from ever forming a spore. The megaspore remains attached to the tapetum underneath which feeds it. Tapetum next to it, unattached, continues to grow and fill the enlarging sporangium with cushy tissue, though it does not overtake the growth of the megaspore which would result in the spore structure becoming re-embedded. The materchyma continues to grow to no detriment to the rest of the tissues, though with no real contribution at this point.

Reduced to 17% of original (3120 x 4160) user posted image Simplified cut away diagram of vascular fiber of black flora, and close up of vascular barrel system

The progression of xylem-esque tissue in black flora had been a fairly linear one. Beginning as individual freely growing open funnel-like tubes in the Bank Balgae. Transitioning to a more terrestrial life in the Orange Spore Sprout these tubes closed off and bundled together for durability and functionality, though the continual tube remained in the short statured flora which was a limiting characteristic. These tubes were cross joined by pores manned by a single central cell that formed a mesh of fibers to prevent cross contamination between tubes. Finally in the Orange Spore Stalk these hollow tubes pinched and formed staggered short cavities for passing water up the flora bodies, structurally convergent with the tracheid xylem structures of Earth, though created in the intercellular spaces of what were once hollow tubes rather than the cells themselves. The pores even developed a living version of a torus-margo system using their central mesh cell to prevent the spread of air, which would cause complete failure of the vascular system. In much of the descended lineages from that species this formation is conserved, most changes between upright species are cell wall composition rather than structure or arrangement regarding their vascular fibers.

Reduced to 20% of original (2899 x 3450) user posted image Simplified cross section illustration of a recently barked twig. Displayed is a cross section of an infiltrating vascular fiber, two established phloem fibers, and an established lead vascular fiber with fibrous sheath visible.

How vascular tissue increases in number in a given part, such as a twig, branch, root, or even the trunk or stem, is via new infiltrating fibers. In Shaggy Volleypoms these fibers may begin their existence anywhere, but the majority arise in the lower trunk and base with their production stimulated by the increasing diameter and opportunity for expansion of the root system. Twigs and root tips extend with a lead vascular fiber powering behind them. Around this fiber parachyma will expand outward over time, and provide a medium for infiltrating fibers to grow into.

The vascular fiber begins life as a vascular disk, a ring of cells created by differentiating parachyma. Via tissue growth this disk rapidly elongates both upward and downward, or if in a horizontal branch or root it would be toward tip and base, as well as slowly grows in diameter. Shortly after a given section elongates the comprising cells adjust themselves to create long hollow tubes. As the ends of the fiber elongate they trigger parachyma ahead to begin differentiating, shifting to accommodate the upcoming structure. These differentiating cells send the signal along the pathway of growth ahead of them as well, creating a cascade of differentiating cells ahead of the fiber establishing a clear route to grow along. These routes will lead to various parts in need of increased vascular availability, such as leaves, widening twigs, and sporangium. Along this route secondary vascular disks will form, elongating and growing in a similar fashion to the primary fiber and following the already established path, ultimately these will join one another and become a part of the primary fiber themselves. In this way a route can be established in a grown Shaggy Volleypom in under 36 hours.

Phloem forms in a similar fashion, with the formation of a phloem disk consisting of a single broad flat cell. Rather than elongating, however, it sends a signal through the parachyma to create a route of growth, which then subsequently triggers the parachyma to convert to phloem cells. This results in a cord of phloem one cell thick. Phloem disks typically form in the upper new growth, such as twigs and spurs.

As the older portions of the new fibers, both primary and secondary, grow in diameter the surrounding parachyma become tougher sheath cells. This layer grows with the vascular fiber while also providing structural support and a layer of separation between its contents and the rest of the black flora body, similar in function to the casparian strip in Earth plants. Eventually once a section begins reaching a threshold diameter, typically around 0.3 - 1 millimeters, the long hollow tubes inside will start pinching to form staggered vascular barrels. This is the mature form of the vascular fiber, and at that point as the fiber continues to grow in diameter it will immediately include the additions of New vascular barrels with it. Barrels found in horizontal roots tend to be notably longer.

The formation of new twigs, which in turn give rise to branches, is triggered similarly to the formation of leaf spurs, however with a continual and comparably fast extension of the spur and parachyma rather than capping off with a microphyll. This developing twig normally recruits infiltrating vascular fibers to become its lead fibers, but in unusual cases can force side growth out of younger fibers if no infiltrating ones are available.

The branches and trunk of the Shaggy Volleypom have no distinct rings, they do not have a true cambium like many earth plants. Rather their growth is carried out by a mess of undifferentiated and secondarily undifferentiated cells throughout its width as tissue analogous to parachyma. This parachyma gives rise to vascular fibers and simple phloem dispersed throughout itself in no particular pattern other than age. The vascular fibers end up roughly numbering greatest toward the center of the trunk and older branches, as these large living structures continue to expand themselves as well as have their numbers multiplied by differentiating parachyma during the duration of the organism, while never being deconstructed. The walls of the cells of the vascular fibers are in multiple layers, some highly lignified and others not, these add structural support for the whole organism.The simple phloem, thick strands of fiber comprised of disk shaped cells stacked on one another, number greatly throughout the parachyma, sections being dismantled and replaced as damage occurs. The parachyma grows between these fibers, the tough and lignified vascular fibers acting as central points it pushes away from and the phloem gets moved about freely by this growth due to size and thinner cell walls.

As parachyma cells experience increased pressure between vascular fibers they develop greater and greater lignification in their cell walls. This process tends to start first with cells roughly toward the middle, where the pressures have existed longest and the vascular fibers are closest together, but can arise between close vascular fibers away from the center as well. Once fully enclosed by lignified parachyma a vascular fiber no longer can expand outward from its center and will stop expanding in that direction. Fibers found in the partially lignified areas tend to be largest, as they are the oldest fibers not yet enclosed in hard woody tissue. Those vascular fibers found in the soft undifferentiated parachyma are generally younger and smaller. The growing parachyma will treat the lignified parachyma similarly to vascular fiber, pushing off it, which increases pressure between it and any other unbudging structures, thus perpetuating The growth of lignified tissues. As phloem is pushed away from lignified areas by parachyma growth they gradually form clusters. Eventually these clusters will become trapped in wood tissue where they no longer can move around and over time may be crushed under pressure as the surrounding cell walls continue to thicken.

The parachyma experiences very slight disconnection between cells, allowing for extracellular spacing in which air may travel and gas exchange can occur, water in these spaces is generally soaked up by cells to keep them free from blockage and freighted toward vascular fibers for further use by the black flora. Other extracellular labyrinths, unconnected, do, however, allow water to traverse between cells. This allows cells to remain hydrated and is an artifact from various ancestors using guttation and evaporation through their stem and trunk surface to assist in the movement of water up their bodies.

Reduced to 19% of original (3120 x 3727) user posted image Simplified cross section of a woody twig. P1. Undifferentiated parachyma, P2. Partially lignified parachyma, P3. Fully lignified parachyma. B1. Undifferentiated PCC, B2. Fibrous PCC, B3. Bark layer

The edge of the parachyma is ringed with a tissue reminiscent in function to a proto-cork cambium (PCC). At the boundary of the parachyma and proto-cork cambium the latter is broken apart by the growth of the former. Cells of PCC replicate here, always pushing outward from the parachyma. As observation travels further outward along the PCC the tissue becomes more uniform, vertically fibrous, and unbroken, with excepting to strips housing access to the inner air and water labyrinths. Further out still the access to the water labyrinths is severed, at this point the cells are tighter packed, and begin excretion of lignin and suberin into their extracellular matrix, or cell wall, making them more rigid and very waterproof. As these masses of cells are pushed out further by those behind them their layers fracture into macroscopic fibrous chunks that can measure up to 30cm in length. These chunks may curl slightly at their edges due to desiccation, though the cells inside at this point should all be dead and desiccated regardless. At this point it is the surface of the branch or trunk, and what remains is a flaky and fibrous, waterproof, layer that can be called a bark (though more specifically, periderm). This bark as it continues to be grown may end up several centimeters thick in older areas, and non-existent in new tissues where parachyma is directly exposed.

The roots of of the Shaggy Volleypom can reach very deep into the soil on their search to access groundwater. This helps prevent the massive organism from drying out. These organs can be as long as the organism itself, growing outward and downward. Their massive root nets higher up spread their base support and anchor them to the ground and other rooted flora, preventing wind storms from so easily uprooting them and knocking them over.

These deep roots also provide access to a range of otherwise locked away substances, including arsenic. This element when drawn up into the Shaggy Volleypom is not shed or excreted readily like other noxious chemicals found during extraction. Rather it is used in the formation of organic arsenic acids that accumulate in the leaves and new growth of the organism, though not the sporangiums. This substance deters and can even kill foraging herbivores, especially smaller ones with voracious appetites and inability to easily source other foods. As leaves are shed to the forest floor they increase the bioavailability of arsenic in their area, which will be drawn back up into the Shaggy Volleypom, but may also spread into other organisms as contamination.

Reduced to 18% of original (3843 x 1789) user posted image simplified cross section of a growing roothair.

The structure and growth among the primary and lateral roots are fairly similar to the trunk and branches, playing a similar role of transporting water and nutrients. However a few differences are that the barrels of the vascular fibers are more elongate, the bark layer is very thin, and there is more storage specialization among their parachyma. The formation of roothair structures is similar to how vascular tissue arises from Volleypom embryonic cell mats. Epidermal cells trigger parachyma just beneath them to form vascular disks, albeit these consist of a single cell rather than a whole bead of tissue. This cell divides, then divides again to form four daughter cells which then begin growing a length of lumen tube that remains to be only four cells in perimeter. This lumen tube pushes through the epidermal layer directly into the surrounding soil where water and dissolved nutrients and salts may diffuse into the space of the tube. At the point of origin the four cells of the tube pinch the tube shut and multiply so this dead end fans open in shape. The cell walls between them fill with suberin, the same waterproof stuff in the Volleypom bark, and force any water or nutrients to actively pass through the cells rather than between. This way there is a checkpoint for anything entering the root where passage can be allowed or denied. Once through this cork water and nutrients is passed through parachyma on into the vascular fibers where it can rapidly travel further up the organism.

Reduced to 18% of original (2807 x 3973) user posted image Simplified cross section diagram of a germinating megaspore, chronologically top to bottom.

The germination process of a megaspore differs somewhat from the typical black flora spore, allowed by its size. The inside of a fully mature megaspore is made up of many lobes of tissue, as well as a single massive yolk-like cell with a loose and cushy cell wall. This singular cell is the zygote. Surrounding it is a soft armor of cushion lobes to protect it from jostling or taking damage during falling. In front of it is a single lobe with the purpose of breaking through the germ pore during germination, after which it is absorbed by the growing organism. Behind the zygote are large storage lobes that provide energy and materials to the developing organism before leaving it's shell and during it's initial growth outside. The cushion lobes share this role after completing their main function. Toward the very back is a long lobe that takes on storage as a secondary function as well, the umbilicus lobe. This developed from a string of spore cells through which the parent Volleypom fed the rest of the growing spore tissue and was directly attached to the megasporangium.

Once dormancy is broken the zygote will begin rapidly dividing into smaller and smaller filamentous cells. These cells form a fibrous mat akin to the filamentous mats and films of other black flora during their early development, however this stage is able to occur entirely within the safety of the megaspore shell. Eventually the surrounding lobes are depleted and absorbed into the mat, at which point the front most lobe, or egg tooth lobe, breaks through the germ pore and surrounding shell. The cell mat grows through this new hole, spilling out as a ball of gray-black fuzz as its photosynthic pigments start being produced. Shortly after this the surface of the fuzz ball begins firming into a true epidermis and beneath this layer parachyma and prevascular disks begin forming.

Once the epidermis is complete the remainder of fibrous cell mat outside the megaspore shell is converted to parachyma. What's left of the mat inside the shell acts as a food source and depletes itself into the rest of the more defined body. The vascular disks beneath the epidermis begin developing further into true vascular fibers at this point. The largest and fastest growing one quickly becomes the dominant fiber, completely passing through the newly formed body creating a growing stalk and root. The many other vascular fibers cease their growth, essentially taking up the role of backup if the dominant fiber takes irreparable damage. If above ground they also can increase surface area for photosynthesis, and lean across the megaspore shell as a clear surface to perform this task.

Reduced to 30% of original (2370 x 2047) user posted image Shaggy Volleypom sprouts