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Can you help identify this species?
(Link to imgur album with all images)
It was growing next to the Yarra River (maybe 10 meters from the bank) in Melbourne. At first I thought it was a type of pine however this page says Australian pines at not actually true pines. It does look much like some of the images of Australian Pine Casuarina equisetifolia
However the Wikipedia page for Casuarina equisetifolia says there are 2 subspecies, equisetifolia and incana. Equisetifolia is only found in Southeast Asia & northern Australia so that rules that one out. Incana is found on the east coast of Australia (eastern Queensland, New South Wales) - could it also be found as far down south as Melbourne?
I've also compared it against all the pines on this site as it specifically shows Australian species.
From there I thought it looks a bit like a Port Jackson Cypress Pine Callitris rhomboidea but the needles look longer and narrower.
I think it also looks a little like a Narrow Leaved Wattle Acacia linearifolia if you ignore the yellow flowers.
I'm carving a spoon out of some of it's wood so would like to know for sure what species it actually is. Any help greatly appreciated.
I went back to the tree to get some more photos but the whole tree had fallen in the river!
I did get a branch off it however to get a close up of the nodes / internodes but I could not find any seeds or cones.
I've tried to get a close up of the apical portion (the top bit right?). My twig sample has started to dry up a bit so this is the best I could get.
ONE MORE UPDATE
Due to my last pictures being rather fuzzy I got out the DSLR and the tripod. These photos are very high res and around 8mb each so may take a while to load…
Native cherry, Cherry ballart or Cypress cherryn belongs to the sandalwood family of plants.
I did some research following the latest answer on species of Exocarpos. I believe it is Exocarpus cupressiformis as well.
You can refer to this Australian website for a key to determine the actual species of Exocarpos : http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=gn&name=Exocarpos#4
Also, as I researched, I found other Exocarpos species to mismatch with your images, except for two; Exocarpos cupressiformis and Exocarpos strictus. One mismatch of E. strictus was with the way flowers were arranged in your last few pictures.
Find below the pictures of flowers of E. cupressiformis (First) and E. strictus:
You can see that E. strcitus has the flowers blooming on stem edges and not at all on tips. In contrast, your images show flowers clustering at tips just like E.cupressiformis.
Australian Pine Tree (Casuarina equisetifolia)
Australian Pine Tree, though a tree, has an exotic bird scent. And yes, as an exotic bird, unfortunatelyit does not have any particular value, but as an ornamental or productive one it is great.
The productive aspect of the tree willbe considered in a future &ndash we say now &ndash special tribute. Instead, in the article below, the ornamental side willbe presented here and now.
Australian Pine Tree Origin
Australian Pine Tree (Casuarina equisetifolia)
How do you date a tree?
If no humans were around to record the planting or germination of a tree, how can its age be determined?
The trees themselves can help tell us their age, but not just by looking at their size.
Big trees are not necessarily old trees — they might just be very healthy or fast-growing individuals.
A much more reliable way to determine the age of a tree is through their wood and the science of dendrochronology (tree-ring dating).
Flickr: Sheila Miguez, CC BY-SA
Many trees lay down different types of cell wall material in response to seasonal patterns of light, temperature or moisture.
Where the cell walls laid down at the beginning of the growth season look different to those laid down at the end of the season, rings of annual growth can be seen in cross-sections of the tree.
This map of growth patterns can also be cross-dated or correlated with major events like multi-year droughts or volcanic eruptions that spewed material into the atmosphere to be incorporated into the wood of the tree.
But the cell walls are more than just calendars.
Wollemia nobilis is an evergreen tree reaching 25–40 m (82–131 ft) tall. The bark is very distinctive, dark brown, and knobbly, quoted as resembling a popular breakfast cereal.  The tree coppices readily, and most specimens are multiple-trunked or appear as clumps of trunks thought to derive from old coppice growth, with some consisting of up to 100 stems of differing sizes.  The branching is unusual in that nearly all the side branches never have further branching. After a few years, each branch either terminates in a cone (either male or female) or ceases growth. After this, or when the cone becomes mature, the branch dies. New branches then arise from dormant buds on the main trunk. Rarely, a side branch will turn erect and develop into a secondary trunk, which then bears a new set of side branches.
The leaves are flat linear, 3–8 cm (1.2–3.1 in) long and 2–5 mm (0.079–0.197 in) broad. They are arranged spirally on the shoot but twisted at the base to appear in two or four flattened ranks. As the leaves mature, they develop from bright lime-green to a more yellowish-green.  The seed cones are green, 6–12 cm (2.4–4.7 in) long and 5–10 cm (2.0–3.9 in) in diameter, and mature about 18–20 months after wind pollination. They disintegrate at maturity to release the seeds which are small and brown, thin and papery with a wing around the edge to aid wind-dispersal.  The male (pollen) cones are slender conic, 5–11 cm (2.0–4.3 in) long and 1–2 cm (0.39–0.79 in) broad and reddish-brown in colour and are lower on the tree than the seed cones.  Seedlings appear to be slow-growing  and mature trees are extremely long-lived some of the older individuals today are estimated to be between 500 and 1,000 years old. 
The discovery, on or about 10 September 1994, by David Noble, Michael Casteleyn and Tony Zimmerman, only occurred because the group had been systematically exploring the area looking for new canyons.  Noble had good botanical knowledge, and quickly recognised the trees as unusual because of the unique bark and worthy of further investigation. He took specimens to work for identification, expecting someone to be able to identify the plants.  His specimens were identified by Wyn Jones, a botanist with National Parks and Jan Allen from the Botanical Gardens. After the identification was made, National Parks then went under a veil of secrecy, with the discoverers not learning the full magnitude of their discovery for about six months. National Parks came close to damaging the stand when a helicopter being used to collect cones inadvertently pruned one of the pines with its rotor.  The species was subsequently named after Dave. The other members of the discovery party questioned the naming but were informed that nobilis referred to the trees being noble in structure and not to David Noble. 
The first illustrations of the Wollemi Pine were done by David Mackay, a botanical artist and scientific illustrator who was working at the Royal Botanic Gardens in Sydney when the species was discovered. 
Further study would be needed to establish its relationship to other conifers. The initial suspicion was that it had certain characteristics of the 200-million-year-old family Araucariaceae, but was not similar to any living species in the family. Comparison with living and fossilised Araucariaceae proved that it was a member of that family, and it has been placed into a new genus, beside the genera Agathis and Araucaria.
Fossils closely resembling Wollemia that are thought to be related to it are widespread in Australia, New Zealand and Antarctica from Cretaceous era sediments, but Wollemia nobilis is the sole living member of its genus. These trees remained common throughout eastern Australia until around 40 million years ago but then gradually declined in range and abundance. Before the relict population was discovered in Wollemi National Park, the most recent known fossils of the genus date from approximately 2 million years ago in Tasmania.   It is thus described as a living fossil or, alternatively, a Lazarus taxon.
Fewer than a hundred trees are known to be growing wild, in three localities not far apart. It is very difficult to count individuals, as most trees are multistemmed and may have a connected root system. Genetic testing has revealed that all the specimens are genetically indistinguishable, suggesting that the species has been through a genetic bottleneck in which its population became so low (possibly just one or two individuals) that all genetic variability was lost. 
In November 2005, wild-growing trees were found to be infected with Phytophthora cinnamomi.  New South Wales park rangers believe the virulent water mould was introduced by unauthorised visitors to the site, the location of which is still undisclosed to the public. 
The grove of Wollemia trees was endangered by fire during the 2019-20 Australian bushfire season.  They were saved by specialist firefighters from the National Parks and Wildlife Service, supported by the Rural Fire Service who installed an irrigation system as well as dropping retardant.     
A propagation programme made Wollemi pine specimens available to botanical gardens, first in Australia in 2006 and subsequently throughout the world. It may prove to be a valuable tree for ornament, either planted in open ground or for tubs and planters. In Australia, potted native Wollemi pines have been promoted as a Christmas tree.  It is also proving to be more adaptable and cold-hardy than its restricted temperate-subtropical, humid distribution would suggest, tolerating temperatures between −5 and 45 °C (23 and 113 °F), with reports, from Japan and the USA, that it can survive down to −12 °C (10 °F). A grove of Wollemi pines planted in Inverewe Garden, Scotland, believed to be the most northerly location of any successful planting, have survived temperatures of −7 °C (19 °F), recorded in January 2010.  It also handles both full sun and full shade. Like many other Australian trees, Wollemia is susceptible to the pathogenic water mould Phytophthora cinnamomi, so this may limit its potential as a timber tree.  The Royal Botanic Gardens in Sydney have published information on how to grow Wollemi Pines from seed which has been harvested from helicopters from the forest trees. The majority of seeds that fall from the cone are not viable so need to be sorted to retain the plump and dark ones. These can then be sown on top of seed raising mix and watered. Once the water has drained through the mix, the pot should be placed in a plastic bag and refrigerated for two weeks. After this, the pot should be removed from the plastic bag and placed somewhere warm but not very sunny until the seed germinates (remembering to keep them moist but not wet). This could take several months.  Examples of the species can be viewed at The Tasmanian Arboretum.
The Wollemi Pine is extremely hardy and versatile in cultivation. Despite it being an endangered species, it is easy to grow and requires relatively low maintenance. It will adapt to a diverse range of climatic zones, thriving in full sun to semi shaded outdoor positions. They can be maintained in a pot almost indefinitely, and makes a good container plants for patios, verandas, and courtyards. Because it tolerates air conditioning, it can also be used as an indoor decorative plant. These are basic need to knows for care: 
When pruning the Wollemi pine, use sterile secateurs at any time of year to retain its compact form. It can be pruned heavily with up to two thirds of the plant size removed. Pruning heavily can be done on the apical growth and the branches. The best time to prune is during the winter months.
Growth Rate Edit
The Wollemi Pine has very controlled growth, especially if it is kept in a pot. It may take up to 25 years to reach 20 feet in height.
The genus Wollemia shares morphological characteristics with the genera Araucaria and Agathis. Wollemia and Araucaria both have closely crowded sessile and amphistomatic (producing stomata on both sides of the leaf) leaves, and aristate bract scales, while Wollemia and Agathis both have fully fused bracts, ovuliferous scales, and winged seeds.  Scrutiny of the fossil record likewise does not clarify Wollemia’s relationship to Araucaria or Agathis, since the former has similarly disparate leaf characters in its adult and juvenile forms, and the latter has similar cone characters.  Further, the recent description of several extinct genera within the Araucariaceae points to complex relationships within the family and a significant loss of diversity since the Cretaceous.   An early study of the rbcL gene sequence places Wollemia in the basal position of the Araucariaceae and as the sister group to Agathis and Araucaria.  In contrast, another study of the rbcL sequence shows that Wollemia is the sister group to Agathis, and Araucaria is basal.  The different outgroup selection and genes used in previous studies are the reasons behind the discrepancy over the groupings of the three genera.  Later genetic studies corroborate Wollemia's placement in the Araucariaceae as sister to Agathis based on data from the 28s rRNA gene,  a combination of rbcL and matK genes,  and a comprehensive study encompassing nuclear ribosomal 18S and 26S rRNA, chloroplast 16S rRNA, rbcL, matK and rps4, and mitochondrial coxl and atp1 genes. 
Fossils indicate that the lineage leading to modern Agathis and Wollemia evolved from the common ancestor with Araucaria in the Early Cretaceous in southern Gondwana  within climates experiencing cool moist conditions and a strong photoperiod regime.  The most recent common ancestor of Agathis and Wollemia has been proposed to be at least 110 million years old (Early Cretaceous) deduced from the reported oldest fossils of these genera.  However, genetic evidence suggests that the divergence of Agathis and Wollemia occurred 61±15 Ma around the beginning of the Cenozoic rather than in the Early Cretaceous.  In another recent molecular study, an age of only 18 Ma was inferred for the divergence of Agathis and Wollemia.  This also accords with recent revisions of the fossil record in New Zealand that reveal no examples of Agathis or Wollemia-like remains older than the Cenozoic.  The relatively minor genetic and morphological diversity in extant species of Agathis compared to the variation in Araucaria is further evidence of the earlier divergence of Araucaria. 
Below is the phylogeny of the Araucariaceae based on the consensus from the most recent cladistic analysis of molecular data. It shows the relative positions of Wollemia, Agathis, and Araucaria within the division.
This article incorporates text from the ARKive fact-file "Wollemia" under the Creative Commons Attribution-ShareAlike 3.0 Unported License and the GFDL.
The Tree That Could Help Stop the Pandemic
The rare Chilean soapbark tree produces compounds that can boost the body’s reaction to vaccines.
In early April , Paul Hiley was kicking back in the executive suite at Desert King International LLC, gazing out the window at the San Diego sunshine and daydreaming about his golf game. California had issued its initial stay-at-home order for COVID-19, but apart from the hand sanitizer around the office, life was more or less normal. Retirement was on the horizon for Hiley. Maybe he’d sell the business. Maybe his son, Damian, would take over.
For more than 42 years, Hiley has been a leading purveyor of certain plant-based food additives such as saponins, foaming agents used in root beer and Slurpees. Most of us never think about these compounds, and Hiley has always liked it that way. “My theory of business is the only two people who need to know my name are my wife and my banker,” he told me recently.
Then, one day—April 14th, to be exact—his son told him that they had a call with Stanley Erck. Erck is the CEO of Novavax, a Maryland-based maker of vaccines. Not a seller of vaccines, mind you: The company had yet to bring one of its candidates to market. But like other companies around the world, Novavax had thrown its hat into the coronavirus-vaccine race. And its success, Erck believed, depended on that odd ingredient in Slurpees.
The inner bark of the Chilean soapbark tree, Quillaja saponaria, is the source material for some of these saponins. Pulverized and soaked in water at the Desert King factory in Chile, the bark is transformed into a brown, bitter, bubbly fluid. This precious goo does many things well, and it happens to be the raw material for one of the world’s most coveted vaccine adjuvants: QS-21. Adjuvants are compounds that boost the body’s immune reaction to a vaccine. Owing to their potential risks to human health, however, only a handful of adjuvants have been approved by the U.S. Food and Drug Administration, and QS-21 is one of the newest.
A single gram of powdered QS-21 costs more than $100,000, though only about $5 worth is needed for each shot. Nine years ago, researchers estimated that the global supply of pharmaceutical-grade Quillaja extract was sufficient for just 6 million doses of vaccine. Everyone in the business knew the story of the Pacific yew tree, whose bark was the original source of the chemotherapy drug paclitaxel, and which was threatened by large-scale harvesting in the 1980s. “If you take out all the trees in one shot and deplete the source of saponin, you are in deep shit in the future,” says Garo Armen, whose company, Agenus, helped bring QS-21 to market. Novavax has its own saponin-based adjuvant, called Matrix-M, and warned investors last year that their vaccines could be delayed if they failed to “secure sufficient supplies” of high-quality extract. And the Hileys practically had a monopoly on it.
During his call with the Hileys, Erck asked if Desert King could increase their production for Novavax a hundredfold. Paul Hiley’s jaw dropped to the table. Novavax was on the cusp of receiving $384 million in international funding to help it produce 100 million doses of its COVID-19 vaccine for the world by the end of the year, and a billion doses by the end of 2021. It would also soon be on the short list of vaccine candidates funded by the Trump administration’s Operation Warp Speed. Novavax needed guarantees of 1,500 pounds of saponin now, and up to three times as much next year.
Hiley’s immediate concern was that his Chilean operation had already missed the bark-harvesting window—typically during the trees’ spring growth, between September and December. And last year he had made the ill-timed decision to postpone expanding their pharmaceutical operations in favor of investing in Desert King’s booming animal-feed business.
In the end, Hiley knew there was only one way to answer Erck: “Of course, we can deliver it,” he said. Three months later, inside the Desert King conference room in early July, sitting across from a shelf displaying Slurpee cups and bottles of Stewart’s Root Beer, Hiley let out a chuckle through his surgical mask. “I had no idea if we could.”
F or all the talk about the cutting-edge vaccines that may just get us out of the COVID-19 mess, little has been written about adjuvants. Perhaps that shouldn’t be surprising: The late Yale professor Charles Janeway famously called adjuvants the “immunologist’s dirty little secret.”
These unheralded helpers can turn a half-baked vaccine into an effective one, or stretch a scarce vaccine supply during a pandemic. Not every vaccine requires an adjuvant, but many do: Of the more than 200 vaccines listed in the Milken Institute’s COVID-19 vaccine tracker, approximately 40 percent are protein-based vaccines, which rarely work without an adjuvant. Yet adjuvants have never attracted much funding from industry and government. “Adjuvants have been the weak link in vaccines for the last hundred years,” says Nikolai Petrovsky, a vaccine researcher at Flinders University in Adelaide, Australia.
The discovery of adjuvants is credited to a bearded veterinarian named Gaston Ramon, who worked at the Pasteur Institute in Paris in the 1920s. At the time, horses were routinely injected with toxins from tetanus and diphtheria so their bodies would produce antibodies that could be used in human therapies. Ramon, who was trying to develop the first human vaccines for these life-threatening diseases, observed that the horses’ circulating antibodies generally declined over time—even if the animals were re-injected with bacterial toxins every few weeks. Every so often, however, a new injection would cause waning antibody levels to rebound.
When he examined the horses whose antibody levels rebounded, Ramon discovered abscesses at their injection sites. Those pus-filled lumps, he thought, could be temporarily trapping the toxins, giving the horses’ immune systems more time to ramp up their responses. Ramon experimented with ways of artificially slowing the absorption of the injected toxins, mixing them with bread crumbs, powdered infant formula, and tapioca starch—which happens to contain high levels of saponins—to produce local swelling without a full-blown abscess. One horse in his study, injected with a mix of toxins and tapioca, produced five times the normal levels of antibodies.
Meanwhile, a British researcher found that aluminum salts, injected into guinea pigs, had similar but more predictable effects on antibody production. For the next 70 years, they would be the only adjuvants used in vaccines. (While aluminum adjuvants can produce swelling and pain that lasts for a few days, abscesses and other side effects are uncommon.)
The earliest viral vaccines actually did fine without adjuvants. The polio and measles vaccines were initially made from weakened or inactivated whole viruses, which were more than 90 percent effective after several doses. Both vaccine types generate an antibody response, and the live ones also activated another part of the immune system, the T-cell response, which is important for fighting more complex pathogens and can even kill the body’s own cells if they become infected.
The risk of live-virus vaccines is that they can possibly revert to their more dangerous selves or replicate uncontrollably in people with weakened immune systems. Inactivated-whole-virus vaccines have also raised alarm after episodes where they altered the immune system in unpredictable ways. During a clinical trial in the 1960s, 31 infants received a vaccine made with inactivated respiratory syncytial virus (RSV). Those who later caught the virus ended up with a more severe form of the disease two of the 23 who were infected died. By the early 1980s, the quest for ever-safer vaccines against ever-trickier viruses, such as RSV, hepatitis B, and HIV, led researchers to develop vaccines that contained just a fragment of the virus, typically a protein.
But the immune system seemed blind to these new vaccines—until researchers added just the right adjuvant. Aluminum didn’t stimulate an appropriate T-cell response, which scientists noticed could be induced with other substances, such as heat-killed tuberculosis bacteria. “Why do we need to use adjuvants?” Janeway asked in 1989. “To be quite honest, the answer is not known.”
Adjuvants posed their own dangers. During a pilot study of an adjuvanted flu vaccine in the 1990s, some subjects got triple-digit fevers and egg-shaped lumps on their arms. “That scared people,” says Tyler Martin, who once worked at the Chiron Corporation, which developed that vaccine. Adjuvants became a frequent target for the anti-vaccination community, which contributed to the FDA’s conservative approach to them. “At advisory-committee meetings, people come out to rail against adjuvants,” Peter Marks, the director of the FDA’s Center for Biologics Evaluation and Research, says. “We want to make sure they are safe.”
In any event, the side effects were proof that adjuvants weren’t simply slowing down the spread of the antigen through the body. Perhaps, as Janeway himself theorized, they were flipping on some ancient danger switch to alert our immune system of an invader. In 1997, scientists located that switch: Our dendritic cells—the tentacled sentries lurking in our tissues—have at least 10 receptors attuned to pathogens. Some adjuvants act on those receptors. Martin, now the CEO of Adjuvance Technologies in Lincoln, Nebraska, told me, “Once we understand what’s the nature of the immune response we really want to create to COVID, then we can pick the right adjuvants to sculpt that response.”
A couple of weeks after meeting the Hileys, I stood across the street from a Starbucks on the wooded edge of the University of California at Berkeley. It was 10 o’clock on a Wednesday morning in mid-July, but the place was uncannily quiet—all summer classes had moved online. During the previous week, the county had reported more than 1,000 new cases of COVID-19, the highest totals since the outbreak began.
After a few minutes, a man rolled up on his bicycle, a buff cinched around his nose and mouth. Intense dark eyes peeked out from under his fluorescent-green helmet. This was Ricardo San Martin, a scientist who had helped develop the Chilean soapbark industry. He had moved on to other projects, but he still consulted for Desert King. In April, he said, he got a WhatsApp message from Damian Hiley that said simply: “Google Novavax.”
When San Martin heard that Novavax was going to need several thousand pounds of Quillaja extract each year, he started doing the calculations in his head. Since 2000, Chile has cleared 11 percent of its native forests, and mature, accessible Quillaja saponaria trees have become rare. Under Chilean law, landowners need a special permit to cut down Quillaja trees, but they are allowed to prune up to 35 percent of their biomass every five years. Over the next few years, the industry was on course to exceed one published estimate of the maximum sustainable harvest of 27,000 tons, or about 67,500 trees. The Hileys say that number is a significant underestimate of what the forests can bear. Regardless, just one major vaccine rollout would require bark from the equivalent of 5,000 to 7,000 trees per year—or more if you’re only relying on prunings. San Martin realized that while most of the world was thinking about the pandemic’s risk to the human species, someone needed to be thinking about its antidote’s risk to Quillaja. “I feel like if I don’t do it,” he told me, “then who’s going to do it?”
Keeping a safe distance from each other, San Martin and I walked around the perimeter of a small grove of trees just within the campus boundary, mostly eucalyptus and redwoods. We came to a tree about 60 feet tall with gray, sandpapery bark and waxy, oval-shaped leaves with rippled margins. This was it: a Chilean soapbark, one of a handful planted on the Berkeley campus starting in the late 1800s. San Martin—a chemical engineer, not a botanist—doesn’t know how they all ended up here, just that he’s lucky to live near them. (The soapbark is uncommon in the United States and tends to grow well only in California, which has a climate similar to Chile’s.)
Pulling a pair of garden clippers out of a pannier on the side of his bike, he snipped off a few leafy branches to tuck inside a paper bag. Later, back in his garage laboratory a few blocks from campus, San Martin ground up the dried leaves and soaked them in warm water to produce an extract. As he shook the liquid up in a large graduated cylinder that looked like a theater prop, counting to 30, it produced a dense foam like the head of a beer. The amount of that foam, he explained, was roughly equivalent to the extract’s saponin content.
Leaves aren’t currently used by industry because they represent just 5 percent of a mature tree’s weight. In a sapling, however, they may account for 30 to 50 percent of the tree’s biomass and are the only material that can be harvested sustainably. In the face of the current public-health emergency and the potentially life-saving role of saponin adjuvants, San Martin believes we should be establishing new soapbark-tree plantations inside and outside of Chile, and preparing to harvest leaves from the young trees. He’s now testing local soapbark stock inside Berkeley’s greenhouses, hunting for the plants best suited to found plantations on American soil. “What I want now, urgently, is to provide a second source of this natural raw material,” he said.
He hopes to finish a mission he began long ago.
Flash back to the early 1990s: Picture, if you will, San Martin sitting on the toilet. He was a newly minted Ph.D., working at the Catholic University in the Chilean capital of Santiago. He was looking for a project with commercial potential, something that might contribute to the Chilean economy as the country recovered from the cruelty and corruption of the Pinochet dictatorship. He had brought a copy of a United Nations newsletter into the bathroom, and one article caught his eye. It said that a “South American tree” was being tested in a vaccine against HIV.
Only later did he realize that one of those very trees was growing in his own backyard. Chilean soapbarks were once abundant in the sun-soaked hills around the capital, climbing up the flanks of the Andes to about 6,000 feet. For hundreds, if not thousands, of years, the tree bark, ground up and mixed with water, was used as soap by the indigenous Mapuche people. In his 1782 treatise on the natural history of Chile, the Jesuit priest Juan Ignacio Molina wrote of its use by locals: “There is never to be seen on their clothes the least spot or dirt.”
Over the next century, soapbark became an international commodity. American magazines offered up recipes for hair-curling liquids and wool detergents made with the soap-like saponins. Sozodont toothpaste advertised itself as “the only dentifrice” that contained “this salubrious botanical product.” (“When rosy lips part, pearls should glitter behind them.”) In the early 1900s, makers of carbonated beverages discovered that adding soapbark extract to their drinks created a coating of bubbles on the surface that kept the carbonation from escaping. From there, the applications of saponin expanded rapidly: During World War II, it was used in surveillance efforts as both a lens cleaner and an ingredient in photographic reagents.
In 1949, a U.S. government report noted that careless bark-stripping methods were destroying “thousands of trees each year” and regulations were poorly enforced. Some harvesters simply peeled off the vital bark from around the main trunk, girdling the trees and leaving behind a slowly dying forest. Felling 25-year-old trees for about 35 pounds of bark was also wasteful: Up to 95 percent of their weight was being left to rot in the field. Landowners were paid just $30 for a tree’s worth of bark, which was then sent overseas for processing. Chile was destroying its natural heritage for a pittance, and when San Martin met with forestry experts, they asked if there was anything he could do to help.
Back in his lab, San Martin began to study the chemical makeup of the tree’s fibers. Although saponins are most abundant in, and easiest to extract from, the bark, he found he could also obtain them from the trunk and the branches. Using this method, a single tree could replace five or six destroyed in the past. He also found that through judicious pruning, one could improve the condition of the scraggly stump-sprout trees left behind from previous clearcutting.
San Martin hoped to develop a soapbark-processing industry in Chile, and he set out to find international buyers for saponin. With the help of a university loan, he started a company called Natural Response and spent several years hunting for customers—with little luck. With three kids to support, his bank account was shrinking, and interest on the loan was piling up. “I had one car from 1970-something with no brakes,” San Martin said. “I was bankrupted, honestly.”
In 1995, one of San Martin’s employees sent a fax to Paul Hiley, whose business at that time revolved around saponins from Mexican yucca. Within days, Hiley was stepping off a plane in Chile. San Martin took him to the university to show him a small barrel of highly purified, powdered white saponin. “He was proud of his little production facility,” Hiley told me.
But Hiley wasn’t interested in the high-grade stuff back then. He pointed at the cola-colored syrup that San Martin had yet to process. The crude extract was exactly what Hiley needed for his clients in the soft-drink business. “I’ll buy 10 tons,” Hiley said. He purchased that first shipment and then wired San Martin an extra $300,000 to partner with him and expand the business.
San Martin was still in a hole: He needed his extract to be a pure liquid, but it came out cloudy, filled with microscopic particles. Every attempt to refine it in the factory negated the cost-saving efficiencies he’d achieved in the field, and he wasn’t hitting the price targets he had promised Hiley. “I couldn’t say, ‘Paul, pay me 12, and you sell it at 11.’”
In the late 1990s, while visiting his children in Montpellier, France, he stopped in a bookstore. He plucked a two-volume treatise on wine making off the shelf. “Why did I pull that out? I don’t know,” San Martin said. As he started reading, he realized that the food-safe processes that vintners used to remove clumps of tannin from their fermented grape juice could be applied to soapbark extract: “I rushed to Chile with that thing.” It was his Eureka moment.
It was also his introduction to the Jevons paradox, the frustrating phenomenon by which technological increases in efficiency fuel increased demand. With San Martin’s innovations, the annual harvest of Quillaja declined from a high of 20,000 tons per year to around 5,000 tons. Then it started creeping up, exceeding 11,000 tons in 2012. The annual harvest is now approaching 20,000 tons again, according to Hiley.
To some degree, this was San Martin’s own fault. He couldn’t stop inventing new applications for the soapy substance. He discovered that it could be used as a bio-pesticide for nematodes on grapes. (Saponins likely evolved as a defense against pests.) Then he found that it reduced the toxic mist of sulphuric acid that rises out of copper-extraction tanks.
San Martin sold his remaining stake in the business to Hiley in 2005, and moved to Berkeley in 2013. Over the past 15 years, the fastest-growing part of Desert King’s business has been their saponin-based animal supplements, which can improve growth rates and reduce Salmonella infections in chickens. Desert King says its saponins, manufactured and sold by major feed companies, are now fed to more than 50 percent of antibiotic-free poultry in the U.S. The supplements are also showing promise in preventing infections of viruses and parasites in fish, including farmed salmon, a big business in Chile.
Everyone at Desert King was enough of a believer in Quillaja’s juju that they began putting a few drops of extract into their coffee or orange juice each day to ward off disease. “It tastes like soap,” Damian Hiley told me. Whether this did anything for their health was doubtful, but the profits were undeniable: By the late 2000s, the company was bringing in tens of millions of dollars per year. Then, three years ago, the Food and Drug Administration gave QS-21 the nod of approval. “Everyone was knocking on our door,” Damian said.
I t is often said that vaccines are one of the most successful public-health interventions in human history. They are also bad business propositions. Two-thirds of vaccines fail in clinical trials. Once approved, they are often less profitable than drugs for cancer or rare diseases. In 2004, just five companies were manufacturing vaccines for Americans, down from 26 in 1967.
Since then, vaccine makers have lost money trying to develop vaccines for Zika and Ebola, because the outbreaks subsided and government funding dried up. When the new coronavirus landed on U.S. shores, the major vaccine makers sat on the sidelines for weeks—a situation that Anthony Fauci, the director of the National Institute of Allergy and Infectious Diseases, characterized as “very frustrating” during a February event at the Aspen Institute.
Novavax, however, was the eager kid waving a hand in the back of the room. Founded in 1987, the company had so far failed to bring a vaccine to market, and stayed afloat through private investment, research contracts, and licensing deals. Last year, its clinical trial for an RSV vaccine was a bust. Tens of millions of dollars went down the drain, employees were laid off, and two development and manufacturing facilities were sold. Soul searching ensued. “If you get bad data, everybody thinks you’re a failure,” Gregory Glenn, the company’s president of R&D, says. “I have PTSD from that.”
They got a gold star for attendance, though. Over the past decade, Glenn’s scientists have repeatedly pursued vaccines for emerging diseases including swine flu, Ebola, and Middle East respiratory syndrome, another coronavirus. Their laboratories maintained a stock of cells originally plucked from the ovaries of caterpillars in the 1970s. These cells were little factories that could be induced to pump out just about any kind of virus protein, including the coronavirus spike.
Before Glenn joined Novavax in 2010, the company wasn’t a believer in adjuvants, arguing that an unadjuvanted flu vaccine would be faster to win FDA approval. But Glenn, a pediatrician who had worked in the laboratory of an adjuvant expert at the Walter Reed Army Institute of Research, thought it was time to embrace them.
Adjuvants had undergone a renaissance, and QS-21 was its poster child. A crude saponin extract had been used in veterinary vaccines since the 1950s, but it was too toxic for humans, causing red blood cells to burst. In the 1990s, a researcher named Charlotte Kensil separated some of the 50 or so saponins in Quillaja saponaria extract, then tested them individually in mice. QS-7 was a potent adjuvant, but there wasn’t a lot of it. QS-18 proved to be the most toxic. QS-21 was relatively mild and generated both an antibody and a T-cell response.
GlaxoSmithKline licensed QS-21 from the maker. In order to tune the immune response, it combined QS-21 with a second adjuvant, a fat-like substance derived from Salmonella bacteria. Three years ago, this potent combo came onto the market in their shingles vaccine, called Shingrix. “That vaccine hit the ball out of the park,” says Janet McElhaney, an expert on aging and immunity at Health Sciences North in Ontario.
Shingrix conferred immunity on 91 percent of people over 70 years old, more than double that of a previous shingles vaccine. Last year, the same adjuvant combo was rolled out in parts of the world in GSK’s malaria vaccine, Mosquirix, and it is also a component of a late-stage-tuberculosis vaccine candidate.
Novavax, meanwhile, obtained the rights to a different saponin-based adjuvant, now called Matrix-M, which was developed by a Swedish researcher who had worked on that HIV vaccine San Martin first read about. Recently, Novavax has tested Matrix-M as part of its NanoFlu vaccine, which not only provided a stronger antibody response than existing flu vaccines but also offered cross-protection against multiple strains of influenza.
By the time COVID-19 arrived, the company was finishing up Phase 3 clinical trials of NanoFlu, which would demonstrate Matrix-M’s safety in 2,650 human subjects. In February, it began testing its COVID-19 vaccine with Matrix-M in animals, and the results coming out in the spring were promising. “We all need to be humble in front of trying to make a billion doses,” Glenn told me a couple of months ago. “But, so far, things have gone exceptionally well for us.”
N o matter how effective a COVID-19 vaccine is, it won’t put a dent in the pandemic unless it can be produced on a massive scale. The downside of an adjuvant is that it adds one more link to the global supply chain, one more crucial connection that can be broken. And by the time Novavax was preparing for its first human tests, the Hileys were struggling to keep their doors open.
As COVID-19 started to circulate in the U.S., Desert King had to provide van transportation for its 220 Chilean workers to replace the public buses forced out of service due to pandemic restrictions. The company pulled strings with local officials for lockdown waivers, and solicited letters of support from Coca-Cola and GSK. By the middle of May, however, they no longer had enough employees coming to work to run their boilers, and they had to shut down. “Every company had the same sob story,” Damian Hiley said. “Maybe our messaging was falling on deaf ears.”
The impending closure triggered a red alert at Novavax’s headquarters, some 5,000 miles away. The company sent an official letter to Chilean President Sebastián Piñera, requesting his assistance to help them put a halt to the pandemic. A couple of days later, Desert King got the exemption it needed and was cranking out Quillaja extract around the clock. It also resolved to break ground on a new pharmaceutical manufacturing suite. In July, Novavax made headlines with a $1.6 billion commitment from Operation Warp Speed, the largest award at the time.
The Hileys knew that their harvesting practices in Chile were now under the global microscope. During my visit to Desert King’s headquarters, Damian showed a brand-new company video that included drone photos of vast stands of Quillaja, an upbeat soundtrack, and the soothing voice of a female narrator describing the company’s “sustainable objectives” and “responsible management.”
“We hadn’t had to do this before,” he said of the public-relations campaign. “We really want to make sure that people understand, especially in Chile, that we are doing this in a responsible, sustainable, renewable, kosher, dotting-every-i-and-crossing-every-t way.”
“I’m not an environmentalist by any stretch,” his father added. “As a capitalist, which I am, if I can make a buck, honestly, legally, and help people and not damage Mother Earth, well, check, check, check.”
Desert King doesn’t own much land in Chile. Instead, they make agreements with local landowners. Their harvesters prune trees at intervals ranging from seven years to 20, using San Martin’s low-impact methods. They peel off all the bark they need for vaccines and use the rest of the tree biomass they harvest for their other businesses. They have mapped and tested thousands of trees to track their saponin makeup, which varies greatly by location. “Any fool can go to Chile and harvest a few trees,” Damian said. “The problem with QS-21 is that out of 100 trees, maybe five of them have the right profile.”
As part of its long-term growth plan, Desert King gives out seedlings to Chilean landowners and encourages them to plant native Quillaja instead of exotic eucalyptus and pine. The country’s forestry managers have also distributed Quillaja seedlings around the country—139,000 last year, more than any other species. Five years ago, Desert King invested in a plantation specifically for their pharmaceutical contracts, using cloned trees high in QS-21. Those trees are now large enough for harvesting, but the company still has to ensure that the adjuvant produced from their extract will be equivalent in makeup and quality to what they were using before.
Desert King plans to establish additional plantations in Chile, and possibly elsewhere, to match the needs of Novavax’s adjuvant, but any seedlings planted now will take years to produce harvestable bark—which is why San Martin’s work on Quillaja leaves is so critical. “Who knows what’s going to happen in Chile?” Damian said. “What if they say it’s illegal to harvest Quillaja?” The current supply is vulnerable in other ways: In January 2017, the country had the worst fires in its history, which burned more than a million acres of central-south Chile—a region home to Quillaja trees.
Such concerns have provided an opening for Desert King’s competitors. Tyler Martin of Adjuvance Technologies told me that they can increase the adjuvant yield from a tree by a factor of a hundred, using its semisynthetic version of QS-21. Meanwhile, Agenus is now working with another company to grow QS-21 inside vats filled with cultured plant cells. Its partner, Phyton Biotech, used this method to wean the world off the Pacific yew, and is now the world’s largest supplier of paclitaxel.
The other drugmakers partnering on a protein-subunit vaccine in Operation Warp Speed aren’t taking any chances with the saponin supply chain. GSK and Sanofi are using GSK’s less-potent oil-in-water adjuvant. It, too, contains a natural product—an oily compound from shark livers—but it is unlikely to face a supply constraint and has already been stockpiled. “That’s the way I would have gone,” Carl Alving, a retired Army adjuvant expert, says. “It’s much less expensive and much less difficult to formulate and put together in a very rapid period of time.”
Damian Hiley brushes off the suggestion that the world shouldn’t rely on Quillaja for a COVID-19 vaccine. “That’s complete bullshit,” he said. “There’s plenty of material.” What his naysayers don’t realize, he said, is that San Martin recently revamped their process of extracting saponin from bark, allowing the company to double its efficiency. The company also believes Chile’s forests can sustain four times the current annual harvest. If supplies become tight and saponins are needed for multiple vaccines, they’d just shift production away from, say, chicken feed. “Maybe one day,” he said, “we’ll have to say to those customers, ‘Sorry, guys, we’re no longer supplying this.’”
O n August 4th, Novavax released the first data from its initial safety trials, which had tested the vaccine on 131 human subjects. The immune response was stellar, activating both antibody and T-cell production. “This is the first time I’m looking at something and saying, ‘Yeah, I’d take that,’” the Cornell virologist John Moore told The New York Times. As with other COVID-19 vaccines, some patients had experienced headaches, fatigue, and swelling at the injection site, but there were no serious side effects.
Novavax’s protein-based vaccine will likely only arrive on the U.S. market after the faster-to-develop RNA vaccines from Moderna Therapeutics and Pfizer have received emergency approval. But unlike those gene-based vaccines, which require ultra-low-temperature freezers, protein-based vaccines can be stored in refrigerators, simplifying global distribution. Novavax has entered Phase 3 trials in the United Kingdom, while interim data from their Phase 2 trial in the U.S. is expected by the end of the year. Japan, Canada, South Korea, and the United Kingdom have now secured purchase agreements with Novavax, and the company expects to be able to produce 2 billion doses of vaccine annually in 2021.
San Martin wants a safe, effective vaccine as much as anyone on Earth. While I was in Berkeley, he told me that two of his old friends from Chile had recently died from COVID-19. He and his wife had decided to temporarily increase their social-distancing measures. Six feet wasn’t far enough he wanted 10.
He is looking forward to being able to have a beer with friends, listen to live music, and talk about old times with Paul Hiley without wearing a mask. At times, the thought crosses his mind that he’s the only person who can save the trees in Chile. He brushes such anxious thoughts aside, though, because they prevent him from focusing on the science.
As San Martin and I stood next to Berkeley’s soapbark trees, a groundskeeper rolled up behind us in a maintenance vehicle, a weed whacker and trash can in its bed. “What are we doing?” she hollered.
San Martin spun around, a bouquet of leaves in his hand. “I’m taking a sample,” he said. “This tree has some compounds that are now being used in the best candidates for the COVID vaccine.”
“Wow! That’s amazing,” the woman replied. “We’ll have to plant a lot of those, huh?”
We took a moment to laugh, and to appreciate a brief social connection in dark times. The woman zoomed away. San Martin turned back to the tree with a serious look in his eyes. He craned his neck up toward the drooping branches overhead, then back at the promising leaves in his hand. “Okay,” he said. “So, here we go.”
Pinus halepensis (Aleppo pine)
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|Caption||Canopy structure, leaves and cones of P. halepensis (Montpellier, France). Needles are pale green (usually 6-12 cm long and <1 mm wide), and are borne on silvery-grey branches, usually abscising after the second year.|
|Caption||Seedlings of Pinus halepensis. Community Nursery, Uppington, Northern Cape Province, South Africa.|
|Caption||Seedlings of P. halepensis. Community Nursery, Uppington, Northern Cape Province, South Africa.|
|Caption||Bark and trunk of Aleppo pine, planted as an ornamental tree (University of California, Los Angeles, USA). The bark is greyish and smooth initially, but finally fissured and exposing the reddish-brown or orange inner bark.|
|Caption||Cones of Aleppo pine, partially open and closed (5-12 cm long). Kango Valley, Western Cape Province, South Africa.|
|Caption||Open cone of P. halepensis, growing near Montpelier, France. Cones are moderately to highly serotinous, conical (5-12 cm long), turned downwards, grey to reddish-brown, usually opening after 3-4 years.|
|Title||Vegetation dominated by P. halepensis|
|Caption||Aleppo pine-dominated vegetation, near Montpelier, France.|
|Caption||P. halepensis invading renosterveld on shale-derived soils in the Kango Valley, in the far eastern part of the Western Cape Province, South Africa. P. halepensis has spread from sites of planting in many parts of the southern hemisphere, and in Australia, New Zealand and South Africa, Aleppo pine is a serious environmental weed.|
Preferred Scientific Name
Preferred Common Name
Other Scientific Names
- Pinus abasica hort. ex Carriere
- Pinus arabica Sieber ex Spreng.
- Pinus persica Strangw.
International Common Names
- English: Jerusalem pine
- Spanish: Alepo pinua pi blanc pi bord pino blanquillo pino carrasco pino de Alepo
- French: pin blanc pin d'Alep
- Portuguese: pinheiro francos pinheiro-de-alepo
Local Common Names
- Croatia: alepski bor alepskim borom alepskog bora bili bor
- Germany: Aleppokiefer Seekiefer
- Greece: halepios pefki pefko
- Israel: oren Jerushalaim
- Italy: pino d'Aleppo
- Netherlands: Aleppo pijn
- Poland: alepskiej
- Serbia: alepski bor alepskim borom alepskog bora bili bor
- Turkey: halepcami
Summary of Invasiveness
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Gymnospermae
- Class: Pinopsida
- Family: Pinaceae
- Genus: Pinus
- Species: Pinus halepensis
Notes on Taxonomy and Nomenclature
P. halepensis (2n = 24) and P. brutia together form the subsection Halepenses in the section Pinus, subgenus Pinus (diploxylon or hard pines) of Pinus in the family Pinaceae.
P. brutia was previously considered a subspecies or variety of P. halepensis by some authors, for example, by Gaussen et al. (1993) in their revision of Pinus in Europe. However, P. brutia is now almost universally accepted as a separate species (Price et al., 1998).
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
History of Introduction and Spread
It is difficult to separate natural from adventive distribution in many parts of its range and in the western part of the Mediterranean basin, P. halepensis has a large distribution assumed to be naturalized. For example, the area covered by P. halepensis in the Languedoc region of southern France increased three-fold between 1878 and 1904, and an additional 2.6 times between 1908 and 1978 as the species invaded abandoned lands (Acherar et al., 1984 Lepart and Debussche, 1992). This trend is evident in many other parts of its range, especially where large areas of agricultural land have been abandoned following the imposition of set-aside policies in the European Union (Barbéro et al., 1998).
As P. halepensis is not a species of major commercial importance, no comprehensive data are available on the global extent of plantings for this species, although there are more details concerning Southern Africa and Australia. Where P. halepensis is planted outside its natural range it can exhibit weedy characteristics and in some areas it has become an important weed, and Richardson and Higgins (1998) review the phenomenon of pines as plant invaders in the southern hemisphere. In other areas, P. halepensis has failed to establish, for example, in the Pescadore Islands in the Taiwan Straits (Kao, 1983). It is also notable that P. halepensis is not widely naturalized or a weed problem in California, USA, despite widespread plantings throughout coastal regions of the state where it grows vigorously without irrigation in areas that receive 400-500 mm of rain per year (DM Richardson, University of Cape Town, South Africa, personal communication, 1999). However, one area of California where it is reported invasive is Catalina Island ( Knapp, 2004 ).
P. halepensis has been planted in all countries of southern Africa, where it was first introduced in the mid 1800s. Plantings and trials have been most extensive in South Africa, but the species is also grown in Angola, Botswana, Lesotho, Malawi, Mozambique, Namibia, Zambia and Zimbabwe (Poynton, 1979). P. halepensis is presumed to have been introduced in South Africa before 1830 (Shaughnessy, 1986). The first record of prolific natural regeneration and presumed spread of P. halepensis in the southern hemisphere is from the Caledon district of South Africa in 1855, some 25 years after it arrived in the country. Invasion of fynbos vegetation in South Africa often occurs after a fire (Trabaud, 1991).
P. halepensis arrived in Australia in about 1850, and the main phase of planting, mainly in South Australia, was just before 1900. It has been planted fairly widely in the southern part of the continent, mainly in predominantly winter rainfall areas that receiving 450-750 mm per year. In South and Western Australia, it has invaded disturbed eucalypt forest, especially around Adelaide and on the Eyre Peninsula. P. halepensis appears on the noxious weed list for Australian states and territories, its use being controlled in parts of South Australia, and it is presently unassigned prior to assessment in Western Australia (Anon., 1998). In New Zealand, most invaded sites are extensively managed grasslands, but various semi-natural vegetation types are also invaded. In a study of fifteen alien forestry species in Argentina, Zalba (1995) found that P. halepensis showed the greatest expansion.
Risk of Introduction
|Terrestrial||Managed||Managed grasslands (grazing systems)||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Present, no further details||Harmful (pest or invasive)|
Host Plants and Other Plants Affected
Biology and Ecology
P. halepensis is a diploid species with a chrosome number of 2n=24. Genetic variation in P. halepensis is considerable, and can be clearly observed in morphological features, such as trunk straightness, branch thickness and cone shape. Giordano (1960) studied provenances in Italy differences exist between provenances from Israel, Morocco, Greece and Italy whilst Karschon (1961) established two altitudinal ecotypes in Israel. Morphological and biochemical studies of intraspecific genetic variation within the species have confirmed the existence of two main groups within the natural range of P. halepensis: East Mediterranean and West Mediterranean. The latter is subdivided into four subgroups: Eastern European subgroup (mainly in the Balkan peninsula) West European subgroup (Italy, France and Spain) West North-African subgroup (Morocco and West Algeria) and East North-African subgroup (East Algeria and Tunisia) (Schiller et al., 1986 Schiller and Grunwald, 1987). Different provenances of P. halepensis also display different levels of resistance to pests, and most of the devastating outbreaks of insect pests in Mediterranean Basin pine forests have occurred in planted, rather than natural, pine stands, where inappropriate provenances have been planted, for example, in Morocco (Questienne, 1979). Different provenances of P. halepensis also display different levels of resistance to frost and low water potentials (Schiller and Brunori, 1992) and the importance of careful site-provenance matching is now generally accepted.
It is a not a very long-lived pine, living for up to 150-200 years, though Lepart and Debussche (1991) report that P. halepensis lives until approximately the age of 100 years, but can bear cones at six years and produce viable seed from age 12.
P. halepensis reaches reproductive maturity at a relatively young age enabling it to cope with sites prone to fire. P. halepensis produces seed prolifically (Weber, 2003) and the time interval between seeding events is approximately two years (Moran et al., 2000). According to Moran et al. (2000) the seeds are stored in the canopy and are strongly serotinous. Seeds are able to germinate and establish in the immediate post-fire environment and it would appear that disturbance through fire promotes seedling establishment and invasion (Rouget et al., 2001). There is usually prolific seedling recruitment after fires, leading to dense, even-aged stands. However, seed release is not wholly dependent on fire as dry, hot conditions may also stimulate seed release, as occurs in Israel (Nathan et al., 2000). Lepart and Debussche (1991) note that P. halepensis is not able to recruit seedlings under its own canopy and in natural successions, tends to be replaced by other species such as the oaks Quercus ilex and Q. pubescens. Seeds remain viable for up to ten years (Dean et al., 1986).
P. halepensis is one of the most drought-tolerant of all pines and Rouget et al. (2001) consider that this may have contributed to its widespread invasion of semi-arid shrubland in South Africa. It also survives nutrient-poor soils, frost (provenance-specific) and salt spray, but is unable to withstand waterlogging. In warm, humid conditions, P. halepensis is susceptible to damage from hail storms and subsequent infection by Diploidia pinea. In parts of its adventive range, crowns of P. halepensis can be broken by heavy snowfalls.
The distribution of P. halepensis in the Mediterranean basin follows the 1.5°C isotherm of mean minimum temperature for the coldest months of the year (Trabaud et al., 1985), with annual precipitation in the 300-900 mm range (Houérou, 1974). In the North African part of its range, Le Houérou (1981) argued that P. halepensis forests extended to the 200-150 mm isohyet at approximately 2500 BC, but that extensive deforestation in the 1800s resulted in the shrinkage of its range to areas with at least 300 mm annual rainfall. It grows from sea level to 1700 m (only in Morocco). Summer temperatures averaging 20-25°C accompanied by 3-month drought are typical in its current natural range. P. halepensis is generally considered to be very frost tolerant, surviving winter temperatures as low as -12°C (Calamassi et al., 1999), but this is dependent on microclimatic conditions.
At drier sites within and at the edge of this range, Quercus suber often assumes dominance (Summers, 1939), whereas other pine species (P. pinaster, P. pinea, P. nigra) replace P. halepensis in more humid areas at high elevations. The climatic conditions in the range of P. halepensis are conducive to widespread wildfires indeed the P. halepensis zone is one of the most flammable of all areas in the Mediterranean Basin. Although P. halepensis is best suited to Mediterranean areas with predominantly winter rainfall, it also grows well when planted outside its natural range in uniform- and summer-rainfall regions.
In its natural range in the Mediterranean basin, P. halepensis generally grows on marls, limestone and dolomites, avoiding wet soils and compacted clay soils. In Israel, it grows on luvisols, tolerating terra rossa soils, but preferring brown and pale rendzinas. In South Africa, P. halepensis has proved very accommodating with respect to soils, but it grows best in the presence of lime. It grows well on clay-loams overlying dolerite or shale and, on the highveld, it succeeds on acid, sandy and skeletal soils derived from sandstones. In Australia, it also grows on a wide range of soils, from moderately heavy clays to rather poor sandy podsols, but it does best on sandy loams. For amenity plantings it is generally suitable for most soil types, except for tight clays. Plantations of P. halepensis in southeast Croatia appear to stop the degradation of the limestone soils (karsting) and to improve basic soil properties, resulting in improved regeneration of native trees and shrubs (Tomasevic, 1994) and similar results have been obtained in Bulgaria (Tilev, 1977). In some locations P. halepensis can experience deficiencies in phosphate (Doumas et al., 1983, 1984) and/or potassium (Mandouri, 1981 Doumas et al., 1986).
P. halepensis is often a dominant component of coniferous forests at lower altitudes in the Mediterranean basin and also occurs in maquis, various other scrub communities and on planted coastal dunes. It typically occurs above a layer of shrubs, such as Arbutus unedo, Erica arborea and Myrtus communis or Cistus species. In Australia, P. halepensis has invaded disturbed native eucalypt forest in South Australia and Eucalyptus marginata forest in Western Australia. Aleppo pine is a widespread invader of two types of vegetation in the Cape Floristic Region of South Africa: fynbos and renosterveld.
The most common mycorrhizal fungi found to be associated with P. halepensis in different parts of its range are: Suillus, Rhizopogon and Cenococcum geophilum (Iberia, Spain Diaz et al., 1996) Amanita spissa, Hebeloma edurum, Lactarius deliciosus, Pisolithus tinctorius, Suillus luteus and S. variegatus (Spain Torres and Honrubia, 1994) Pisolithus tinctorius, Rhizopogon roseolus, and Suillus collinitus (south-east Spain Roldan and Albaladejo, 1994 ) Suillus collinitus, S. granulatus, Rhizopogon roseolus, R. luteolus, Amanita muscaria and Lactarius deliciosus (Spain Torres and Honrubia, 1991) Tuber melanosporum, T. brumale var. moschatum, T. aestivum, T. albidum and T. maculatum (Italy Pirazzi and Gregorio, 1987).
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"It also provides a lot of shade as well as [being] a food source of our insects and birds."
The trees are also used by Indigenous people to determine seasonal outlooks.
"For us they are indications of the summer," Mr Blight said.
ABC Great Southern: Ellie Honeybone
"If they flower early, that means were going to have an early summer or, vice versa, if they flower late we're going to have a late one.
"Generally they're in full blossom by Christmas every year."
There we have it! Identifying pine, spruce, and fir trees can be a tricky business, but with a handy identification key like this at your disposal, it shouldn&rsquot be too much of a challenge.
Bookmark this page and visit it whenever you have a tree to identify.
And take this open invitation to send us pictures of any conifers you have difficulty identifying, in the comments section below.
Simply click on the camera icon below the comment box and you can upload up to three photos at a time (up to 6 megabytes each).
In addition to your photo, please let us know what region you are in, whether the tree is in the wild or in your own backyard, and please give us as much detail about the tree as possible, such as how long the needles are.
I&rsquoll get back to you as soon as possible with an answer!
Photos by Matt Suwak, © Ask the Experts, LLC. ALL RIGHTS RESERVED. See our TOS for more details. Uncredited photos: Shutterstock.
About Matt Suwak
Matt Suwak was reared by the bear and the bobcat and the coyote of rural Pennsylvania. This upbringing keeps him permanently affixed to the outdoors where most of his personal time is invested in gardening, bird watching, and hiking. He presently resides in Philadelphia and works under the sun as a landscaper and gardener, and by moonlight as a writer. An incessant questioning of &ldquoWhy?&rdquo affords him countless opportunities to ponder the (in)significance of the great and the small. He considers folksy adages priceless treasures and is fueled almost entirely by beer and hot sauce.
The Jack pine usually grows to be twenty-seven meters tall and sixteen and thirty-two centimeter diameter around the trunk. As it grows it gets rounder and rounder around the crown area. The bark of the jack pine is a reddish-brown. The bark is also flat. As the tree gets older the bark gets grayer. The jack pine has needles instead of leaves. The jack pine has long and slender twigs. They are a reddish color. The jack pine has pinecones that store and produce its seeds. The seeds are usually four to five millimeters long and are dark brown. The Jack Pine gives most animals their nurturance. The tree gives its nurturance to mostly rodents.
The jack pine lives in the taiga biome, which is a semi cold forest in the northern hemisphere. It is mostly found in the colder states and countries. Some examples are, New Hampshire, Maine, and Vermont. The jack pine is plentiful in the wild.
The jack pine adapts to flat or hilly areas that have sandy soil. The jack pine does not have a special root system. It is possible for the jack pine to grow in the taiga because of its waxy pine needles and rough bark. It is good to have waxy pine needles and rough bark in the taiga because the wax on the pine needles protects the needles and so does the rough bark. These things protect the tree from the cold wind in the taiga. These charicteristcs also protect this tree from the weather in the taiga, which is usually cold. In the taiga, the brush starts forest fires and the waxy pine needles and rough bark protect the tree. The jack pine has long and slender twigs so the snow does not stick onto the twig.
The jack pine has two major uses. The jack pine is used to build houses and it is used to make pulp for paper.
Promicromonospora callitridis sp. nov., an endophytic actinobacterium isolated from the surface-sterilized root of an Australian native pine tree
A new strain of the genus Promicromonospora, CAP94T, was isolated from the surface sterilized root of Callitrispreissii (Australian native pine tree). This strain was a Gram-stain-positive, aerobic actinobacterium with hyphae breaking up into fragments which were non-motile, rod-like, coccoid elements. Phylogenetic evaluation based on 16S rRNA gene sequence analysis placed this isolate as a member of the family Promicromonospora ceae, and most closely to Promicromonospora sukumoe NBRC 14650T (99.4 %), Promicromonospora kroppenstedtii DSM 19349T (99.2 %) and Promicromonosporaaerolata V54AT (99.1 %). Chemotaxonomic data including cell-wall components, major menaquinone and major fatty acids confirmed the affiliation of strain CAP94T to the genus Promicromonospora. The results of the phylogenetic analysis, including physiological and biochemical studies in combination with DNA-DNA hybridization, allowed the genotypic and phenotypic differentiation of strain CAP94T and the closest species with validly published names. The name proposed for the new species is Promicromonospora callitridis sp. nov. The type strain is CAP94T (=DSM 103339T=TBRC 6025T).