Undergraduate student projects in our reserves: Alex Bentley studying the pit viper Bothrocophias micropthalmus


Every semester for many years, we have hosted undergraduate students doing independent study projects for a study-abroad program run by the School for International Training in Quito. This year we had six students, each doing independent study projects lasting about two weeks. They are assisted in the field by our reserve guards Santiago, Luis, Fausto, and Jesus Recalde, and they usually stay with families in the commmunities around our reserves. It is a nice way to integrate the reserves with the surrounding communities. Students really enjoy the experience, and their studies are often useful to us for conservation planning. Sometimes they even discover new things!

This semester’s students are finishing up their reports now. I will post excerpts from a few of the more photogenic projects here in the following weeks (and I hope to go back and write about some of the nice projects of past students as well, as time permits).

Alex Bentley is intensely interested in herpetology, so he chose to study our impressive pit viper, Bothrocophias micropthalmus. I’ve written about this snake previously here, here, and here. Alex told me that as he looked for internet data on this snake, my blog posts were almost the only field observations he could find. So clearly there was a need to do more.

Alex spent his time in the Rio Anzu area, where we have a reserve at 1100m-1200m. He stayed much of the time at a nearby Rio Anzu reserve, Sumak Causay, which has a cooperative agreement with us. He found plenty of Bothrocophias in his two weeks. I’ll wait for his paper to write more about his actual study, but for now I just want to put up some of his beautiful photos of our area’s reptiles and amphibians.

Today will be Lizard Day.

Enyalioides praestabilis near the Rio Anzu. Photo: Santiago Recalde/EcoMinga.

Enyalioides praestabilis near the Rio Anzu. Photo: Santiago Recalde/EcoMinga.


Enyalioides praestabilis again. Photo: Alex Bentley.

Enyalioides praestabilis again. Photo: Alex Bentley.


Lizard near Rio Anzu. Photo: Alex Bentley.

Lizard near Rio Anzu. Photo: Alex Bentley.


Closer view of the previous lizard. Photo: Alex Bentley.

Closer view of the previous lizard. Photo: Alex Bentley.


Sleeping lizard near Rio Anzu. Photo: Alex Bentley.

Sleeping lizard near Rio Anzu. Photo: Alex Bentley.


Potamites ecleopus near the Rio Anzu. Photo: Alex Bentley.

Potamites ecleopus near the Rio Anzu. Photo: Alex Bentley.

A brief hike in our Rio Anzu Reserve


A couple of weeks ago I made a short visit to our lowest-elevation reserve, the Rio Anzu Reserve (1100-1200m elevation) in the Amazon basin, to mark some special orchids for a visiting student to study. Lowland Amazonia is the richest habitat on earth for birds and trees, and also hosts a seemingly never-ending parade of crazy insects. A trip to this reserve is always a mind-boggling experience, even though the reserve is very small and lacks larger birds and diurnal mammals due to indigenous hunting pressure in the surrounding area. (However, black jaguars stalk this forest unseen by human eyes, but recorded in several different camera traps…)


For a minute or two I saw this Fulvous Shrike-tanager (Lanio fulvus), a core species of mixed-species insectivorous bird flocks here. Lou Jost/ EcoMinga.

For a minute or two I saw this Fulvous Shrike-tanager (Lanio fulvus), a core species of mixed-species insectivorous bird flocks here. Lou Jost/ EcoMinga.

Quite often at the trail entrance of this reserve there will be a big mixed flock of mostly-insectivorous birds scouring the branches and leaves of the forest. On this trip I met with the flock as soon as I got out of the taxi-truck that brought me there. The flock and I seemed to follow the same forest path for a long way, and I enjoyed their noisy company. A particularly sharp bird call alerted me to the “leader” of the flock, a Fulvous Shrike-tanager (Lanio fulvus) an uncommon bird which does not occur at our higher elevation reserves. This is one of the famous “liar” birds (not to be confused with Lyre-birds!) that watches for hawks, etc, and warns mixed flocks of danger, but will sometimes “freeze” the flock with a false alarm call when it sees a bird flush a particularly appetizing insect. It then grabs the insect for itself (Munn 1986). In spite of its occasional duplicity, the presence of this species allows the other flock members to find more food, since they don’t have to waste as much time looking around for danger (they rely on the Shrike-tanager to do that). So a flock will generally cluster around the local pair of Shrike-Tanagers, and they move together through the forest.

Heliconius butterfly in the Rio Anzu. Photo: Lou Jost/EcoMinga.

This Heliconius butterfly sat on the trail in the Rio Anzu reserve. Photo: Lou Jost/EcoMinga.

Throughout the day fancy butterflies filled the air. My favorite (at least on this day) are the Heliconius butterflies. These butterflies have larva that feed on poisonous passionflower (Passiflora sp.) leaves, and they themselves thus become poisonous to birds. The adults have strong warning colors and patterns, which show a very complex but interesting geographical variation. In any given area, often two different Heliconius species will share exactly the same pattern, but in a different region, the same two species can share a completely different pattern. The geographical variants are intensely studied to give clues about the process of incipient speciation, the possible locations of wet “refugia” during past hot dry epochs, etc. I saw many species that day, but only managed to photograph one.

Passionflower in the forest understory. Photo" Lou Jost/EcoMinga.

Passionflower in the forest understory. Photo:Lou Jost/EcoMinga.

Appropriately I soon found a giant passionflower plant nearby. This species is a canopy liana but has specialized short clambering flowering stems that often come out near the ground. They are pollinated by hummingbirds.

The crown of white pointy “tentacles” in the center of the flower have an important function. Flowers that attract hummingbirds generally produce a lot of nectar, and this nectar is a tempting resource for other creatures, including many that play no role in pollination. Flowers with better defenses against nectar robbery will leave more descendants than those that don’t, so very elaborate defenses have evolved in many hummingbird flowers, including this one. The white spikes protect the nectar below them. They are easily parted by a hummingbird’s needle-like beak, but a clumsy ant or bee can’t get its head close to the nectar.

The back of the flower also has a defense against nectar robbers. The bracts surrounding the base of the flower have “extrafloral nectaries”, glands that produce a bit of nectar themselves. Ants and wasps like to hang out there and drink this nectar, and these nasty bugs scare away other kinds of bugs that could chew through the back to get to the big store of nectar inside.

The day was full of grasshoppers. I photographed an especially flashy one, but many more escaped my lens. One of the grasshoppers I did manage to photograph was carrying two parasitic mites (ticks) on one leg. Mites are commonly seen on insects in the tropics, but I don’t know much about them.

Mites (one healthy, one dead) on a grasshopper's leg in the Rio Anzu. Photo: Lou Jost/EcoMinga.

Mites (one healthy, one dead) on a grasshopper’s leg in the Rio Anzu. Photo: Lou Jost/EcoMinga.

Along with the grasshoppers were many katydids. Most North American katydids eat leaves, but in the tropics things are more complicated. I found a nasty carnivorous katydid munching the severed torso of a walking stick, while the walking stick’s mate another walking stick sat and watched, motionless. The juices of the half-eaten walking stick, in turn, attracted tiny gnats which gathered under the katydid’s head waiting for a chance to steal a mouthful. It was a miniature Serengeti. The annoyed katydid repeatedly swatted the gnats with its forelegs, just like I was swatting the slightly larger gnats that were bugging me. [Edited Dec 1 to reflect my growing doubts that these two walking sticks really belong to the same species. They seem too different from each other. Any experts out there with an informed opinion?]

I found this carnivorous katydid munching on a walking stick while the walking stick's mate looks on. Photo: Lou Jost/EcoMinga.

I found this carnivorous katydid munching on a walking stick while the walking stick’s mate another walking stick looks on. Photo: Lou Jost/EcoMinga.

Victim's head. Photo: Lou Jost/EcoMinga.

Victim’s head. Photo: Lou Jost/EcoMinga.

This walking stick looked on while the katydid ate the other one.

This walking stick looked on while the katydid ate the other one Photo: Lou Jost/EcoMinga.

After a couple of hours I reached the Rio Anzu itself, an easy 15-minute walk if I had ignored the interesting bugs. This is where the ladyslipper orchid Phragmipedium pearcei grows on the wet riverside limestone. The plants are often submerged when the river rises. On this day the river was low and there were many individuals in flower.

A ladyslipper orchid, Phragmipedium pearcei, on the limestone of the Rio Anzu. Photo: Lou Jost/EcoMinga.

A ladyslipper orchid, Phragmipedium pearcei, on the limestone of the Rio Anzu. Photo: Lou Jost/EcoMinga.

The Rio Anzu. Lou Jost/EcoMinga.

The Rio Anzu. Lou Jost/EcoMinga.

At first glance the texture of this ladyslipper orchid flower is unremarkable. It looks smooth like any other flower. I had never given it a second look until that day. A microscope revealed that the flower was a complex mosaic of textures, hairs, glands and stuff I still don’t understand. The hairs were clearly guides for the insect pollinators, which must first land on the white flat rim of the orchid’s pouch or “slipper” (the pouch is called the “lip” in orchid terminology). This white rim has a row of random green spots, and another loosely organized row of larger brown spots. When magnified, the green spots turn out to be many long parallel dark green ridges, separated by greenish brown “valleys”. The effect is almost iridescent. Edit Dec 1: In response to Lisa’s question below, I did some research and found that the pollinator is a female fly that thinks these green spots are actually aphids, the prey of the fly larvae. The female lands on the flower to lay eggs among the “aphids”, and falls into the pouch. My speculations about the spots looking like fly eyes were wrong.

Top view of the "slipper" or lip of the ladyslipper orchid Phragmipedium pearcei. Lou Jost/EcoMinga.

Top view of the “slipper” or lip of the ladyslipper orchid Phragmipedium pearcei. Lou Jost/EcoMinga.

The staminode above the "slipper" or lip. At this magnification the green spots on the lip begin to show their true complexity. Lou Jost/EcoMinga.

The staminode above the “slipper” or lip. At this magnification the green spots on the lip begin to show their true complexity. Lou Jost/EcoMinga.

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Under higher magnification the green spots on the lip reveal complex textures and stiff hairs. Lou Jost/EcoMinga.

Under higher magnification the green spots on the lip reveal complex textures and stiff hairs. Lou Jost/EcoMinga.

Remarkably complex surface details of the green spots. Lou Jost/EcoMinga.

Remarkably complex surface details of the green spots. Lou Jost/EcoMinga.

Closer view of the green spots reveal they are not just smooth spots of color. Lou Jost/EcoMinga.

Closer view of the green spots reveal they are not just smooth spots of color. Lou Jost/EcoMinga.

Eventually the pollinator must fall into the pouch (perhaps drugged by the orchid). Once the pollinator enters the pouch, it finds itself trapped, with limited ways out. Most of the inner surface of the lip is only lightly hairy, but one strip is carpeted with long hairs, and this strip leads the insect up to an escape route that passes directly under the stigma and anthers. The insect thus is forced to pollinate the flower if it wants to get out of there.

A cross-section view of the "slipper". Lou Jost/EcoMinga.

A cross-section view of the “slipper”. Lou Jost/EcoMinga.

Closer cross-sectional view. Note the various kinds of hairs. Lou Jost/EcoMinga.

Closer cross-sectional view. Note the various kinds of hairs. Lou Jost/EcoMinga.

The variety of textures on this flower make me eager to look more closely at other flowers. Expect to see many more micro-photos here in the future!

A jumping spider watched me photographing the grasshoppers. Photo: Lou Jost/EcoMinga.

A jumping spider watched me photographing the grasshoppers. Photo: Lou Jost/EcoMinga.

Lou Jost
EcoMinga

References

Munn, C. A. 1986. Birds that ‘cry wolf.’ Nature 319: 143-145.

First piece of the “Forests in the Sky” is now protected!!!!!!!!!!

This week's "Forests in the Sky" purchases.  Los Llanganates National Park is outlined in green. Our purchase of the yellow property clears out a potentially-problematic inholding in the high peaks of the park and almost reaches down to the low valley of the Rio Pastaza (visible near the bottom of the photo). The blue property links the yellow property to the Rio Pastaza over a tunnel for the Banos-Puyo highway. The red outlines south of the Rio Pastaza are some of our Naturetrek and Cerro Candelaria properties.

This week’s “Forests in the Sky” purchases. Los Llanganates National Park is outlined in green. Our purchase of the yellow property clears out a potentially-problematic inholding in the high peaks of the park and almost reaches down to the low valley of the Rio Pastaza (visible near the bottom of the photo). The blue property links the yellow property to the Rio Pastaza over a tunnel for the Banos-Puyo highway. The red outlines south of the Rio Pastaza are some of our Naturetrek and Cerro Candelaria properties.

Today, thanks to so many generous donors to the World Land Trust “Big Match” campaign, our ten-year-old dream of protecting a strip of forest that would connect Parque Nacional Los Llanganates to Parque Nacional Sangay is now within reach. In just two weeks World Land Trust was able to raise 280,000 pounds for us, and with this we have finally been able to build a protected corridor all the way from some of the high peaks of P. N. Los Llanganates (altitude 3400m) down to the north bank of the Rio Pastaza (alt. 1500m). There our new protected area meets our south bank reserves, the Naturetrek and Cerro Candelaria Reserves, which extend (with a few minor gaps) all the way to the high alpine grasslands of PN Sangay at 3860m. Thus the biological corridor declared long ago by the local governments is now a real thing instead of just a piece of paper.

I know that many of my friends and readers of this blog helped to make this possible. We are very grateful to all of you!!!! (And I hope some of you will come visit this land you helped protect!)

The biological corridor between the national parks of Los Llanganates and Sangay is especially important for large endangered mammals like the Mountain Tapir (Tapirus pinchaque). Photo: Juan Pablo Reyes/EcoMinga.

The biological corridor between the national parks of Los Llanganates and Sangay is especially important for large endangered mammals like the Mountain Tapir (Tapirus pinchaque). Photo: Juan Pablo Reyes/EcoMinga.

The purchase outlined in yellow includes a significant inholding in Parque Nacional Los Llanganates. Our purchase of this inholding gives greater protection to the park, which suffers from heavy abuse (including burning, ranching, and abuse of wildlife) in other inholdings. This property is especially important to me personally, since it is where I discovered my first Teagueia orchid species. The unexpected evolutionary radiation of these Teagueia orchids on this and neighboring mountains is the most remarkable thing I’ve ever found in nature. We had protected about half the members of this radiation when we purchased Cerro Candelaria. Now we have protected the other half of the radiation (curiously, there are no Teagueia species in common between the two mountains).

Teagueia alyssana, one of the first Teagueia species I discovered. I named it after my friend Alyssa Roberts. Photo: Lou Jost.

Teagueia alyssana, one of the first Teagueia species I discovered. It grows within the yellow-outlined property in the map above. I named it after my friend Alyssa Roberts. Photo: Lou Jost.

The main Banos-Puyo highway, which limits the possibilities for a biological corridor, here goes underground through a tunnel deep below one of our purchases (the one outlined in blue in the map at the top of this post). That piece of land was absolutely critical, and we are very lucky to have gotten it.

The Banos-Puyo highway disappears into the mountain (lower left) and comes back out a kilometer away.

The Banos-Puyo highway disappears into the mountain (lower left) and comes back out a kilometer away.

The protected strip we’ve bought this week is very narrow, just 200 meters wide in places. Our original Big Match goal was to also buy the extensive block of excellent cloud forest adjoining that first strip. That financial goal hasn’t been met yet. The owners are negotiating with us now, and WLT has offered to continue the “Forests in the Sky” campaign until this land is secured. When it is secured, we will not only have created a robust corridor, we’ll also have protected a unique cloud forest with many new endemic plant species!

The dark golden-orange outlines are the properties we are still trying to buy, along with the one in aqua, which belongs to a different owner. This will protect the whole watershed of the Rio Machay.

The dark golden-orange outlines are the properties we are still trying to buy, along with the one in aqua, which belongs to a different owner. This will protect the whole watershed of the Rio Machay.

Lepanthes marshana, a new species I discovered in the cloud forest block that we are still trying to buy. Photo: Lou Jost.

Lepanthes marshana, a new species I discovered in the cloud forest block that we are still trying to buy. Photo: Lou Jost.

Lou Jost

Last photos of our young Black-and-chestnut Eagle (Spizaetus isidori)

An adult Black-and-chestnut Eagle delivers a guan to the waiting chick. Photo: Mark Wilson.

An adult Black-and-chestnut Eagle delivers the trunk and legs of a guan to the waiting chick. Photo: Mark Wilson.


Regular readers of this blog know that we (along with many other biologists) were immensely excited about our nesting Black-and-chestnut Eagles, but that the eaglet died when it left the nest. Mark Wilson, who is writing a book on the world’s giant eagles, came here to photograph the nest and eaglet, right before it died. He lost one of his memory cards, but just a few days ago he found it tucked into one of the pockets of his clothes. He sent me the new pictures. Some of them show the adult bringing in the trunk and legs of a Sickle-winged Guan (Chamaepetes goudotii). This was the main prey item brought to the nest throughout the eaglet’s life. A photo by our guards taken a month or two earlier shows exactly the same kind of trunk and legs in the nest.

Black-and-chestnut eaglet mantling prey item at the nest. It does not want to share! Photo: Mark Wilson.

The eaglet mantling prey item at the nest. It does not want to share! Photo: Mark Wilson.

Dreamed Forest- The young Black-and-chestnut Eagle watches life in the cloud forest. Photo: Mark Wilson.

Dreamed Forest- The young Black-and-chestnut Eagle watches life in the cloud forest. Photo: Mark Wilson.

In memory of the  Black-and-chestnut Eagle chick that lived a beautiful but far too short life in its cloud forest nest. Photo: Mark Wilson.

In memory of the Black-and-chestnut Eagle chick… Photo: Mark Wilson.

The last picture shows the eaglet jumping up from the nest in practice flight. This is apparently what got it into trouble, as it somehow got stuck under the canopy and could not regain the nest.

Lou Jost

Cryptic new orchid from our Rio Zunac Reserve, Neooreophilus chaoae, published this week

 Neooreophilus chaoae from our Rio Zunac Reserve. Photo: Lou Jost.

Neooreophilus chaoae from our Rio Zunac Reserve. Photo: Lou Jost.

A few days ago the journal Lankestariana published the description of a new species of orchid I discovered thirteen years ago in what is now our Rio Zunac Reserve in central Ecuador. My friend Sebastian Vieira independently discovered another population of the same species near Sibundoy in southern Colombia, and we wrote the article together. This orchid, which we named Neooreophilus chaoae, is a tiny but beautiful orchid with a very unusual growth habit. A nearly universal rule about plants is that they grow toward the light. For reasons unknown, the genus Neooreophilus defies this rule: its growing tip points downward, toward the ground, so that the newest leaves are lower than the older leaves. The chains of tiny roundish leaves usually hang from a branch or trunk. The chain of leaves looks a lot like some of the small pendant fern fronds that grow in the same habitat. The fern rhizome grows towards the light, like most self-respecting plants, so that each new frond is higher than the last, but each individual frond unfolds downward on a weak stem, just like the orchid.

Left, a climbing fern. Right, a Neooreophilus orchid. Both plants were on the same tree. The resemblance is striking and the orchid may be mimicking the much more common fern.

Left, a climbing fern. Right, a Neooreophilus orchid. Both plants were on the same tree. The resemblance is striking and the orchid may be mimicking the much more common fern.

The resemblance between the orchid and the fern may not be an accident. If there are herbivorous insects that eat orchids and that use their eyes to spot potential hosts, they may have a hard time finding these orchids amid the much more common ferns. The presence of such herbivores would drive the evolution of orchids that mimic other more common plants. In our area there are other orchids that do this, such as certain Elleanthus species that mimic bamboo. (I think plant mimicry of other plants is more common than people realize, and I’ll devote a later post to this topic, complete with local examples.)

Maybe the resemblance to these common ferns also fools field botanists. Members of this very rich genus are rarely noticed or collected by non-specialist botanists, so they are poorly represented in the world’s herbaria. It doesn’t help that they are often partly covered by moss. They are also very fussy about where they grow. They need wet but bright and airy cloud forests, and even within such forests their distributions are patchy, occupying only a small percentage of the seemingly-suitable habitat. The patch size is often just a few meters, but curiously there are often many different species of Neooreophilus in a given patch. Many orchid biologists suspect that this genus-level patchiness is probably due to the presence of a particular fungus in the patch. Orchid seeds need to form relationships with specific fungi in order to germinate, and if the right fungus for Neooreophilus is present in an area, seeds of all the species in the region will eventually fall there and germinate.

Sebastian Vieira's beautiful illustration of our new species. From our paper: Vieira, S., and Jost, L. (2015) A colorful new species of Neooreophilus (Orchidaceae: Pleurothallidinae) from the eastern Andes of Colombia and Ecuador. Lankesteriana 15(3): 213–217.

Sebastian Vieira’s beautiful illustration of our new species. From our paper: Vieira, S., and Jost, L. (2015) A colorful new species of Neooreophilus (Orchidaceae: Pleurothallidinae) from the eastern Andes of Colombia and Ecuador. Lankesteriana 15(3): 213–217.

Sebastian’s discovery of a second population of this species in Colombia, more than 250 km to the north of the population I found, was very surprising. He was on a conservation trip with directors of the Orchid Conservation Alliance (US); the trip led the Alliance to fund the establishment of a new reserve in that area. Sebastian’s discovery was especially interesting because there is another rare and local orchid species shared between Sibundoy and our Rio Zunac Reserve, Dracula exasperata. Neither of these two species are known from any other places in between these two reserves. In the case of N. chaoae, the forms appear to be absolutely identical in the two locations, suggesting that there may be connecting populations somewhere in the poorly-known mountains between the reserves. The Dracula populations, on the other hand, show quite significant differences between the two reserves, and several of my Dracula specialist friends think that the Zunac population should be described as a new species. I’ll write more about this later.

The story of N. chaoae’s twin discoveries highlights our poor level of knowledge about east Andean orchid distributions. Orchid specialists had visited both the Sibundoy and Zunac populations multiple times over the past thirty years, as indicated by the early discoveries of Dracula exasperata in Sibundoy, and Dracula fuligifera, Lepanthes zunagensis, and many others in the Rio Zunac area. Yet none of these specialists noticed N. chaoe. After the first population of N. chaoae was finally found, it took another ten years to find a second population. It is one of many cryptic, often-undetected species in these mountains. A similar story surrounds the discovery of another member of the genus, Neooreophilus (ex Lepanthes) exiguus. Stig Dalstrom and I discovered this species at a remote site in extreme southern Ecuador. Many years later my student Stella Copeland and I found a second population 150 km away in our Rio Zunac Reserve (the same area where I had found N. chaoae). The existence of so many species known from only a few widely separated sites suggests that many new species remain to be discovered in the eastern Andes of Ecuador and Colombia.

Neoorephilus exiguus, another new species I discovered in this genus. Photo: Lou Jost.

Neooreophilus exiguus. Photo: Lou Jost.

My colleague Anne Chao is famous for showing that this relation between the number of rare species and the number of still-undetected species can actually be made precise. She proved that when individuals are randomly sampled, the number of undetected species in the population is expected to be greater than
\frac {f_1^2} {2f_2}  ,
where f_1 is the number of species represented by only a single individual in the sample, and f_2 is the number of species represented by exactly two individuals in the sample. Her formula is widely used in biology and is known as the Chao estimator. Thus it is particularly appropriate that Sebastian and I named our cryptic, often-undetected new species in honor of Anne.

The genus Neooreophilus is very diverse and deserves more attention from botanists. Since I’ve started paying attention to them I’ve been able to discover several new species besides N. chaoae: N. exigua, N. viebrockiana, N. ariasii, and several others that I have not yet published. I am sure there are many more waiting to be discovered!

Neooreophilus viebrockianus, which I discovered about eighteen years ago in what is now our Rio Zunac Reserve. Photo: Lou Jost.

Neooreophilus viebrockianus, which I discovered about eighteen years ago in what is now our Rio Zunac Reserve. Photo: Lou Jost.

Incidentally, my earlier Neooreophilus discoveries were published under the genus Lepanthes, subgenus Brachycladium, where they had originally been placed (Reichenbach 1856, Luer 1996). The flower structure and the characteristics of the sheaths around the leaves and stems suggested this placement. However, recent DNA analyses by Mark Wilson, Alec Pridgeon, and coauthors have clearly shown that these plants are not particularly closely related to Lepanthes, and belong in a completely different branch of the orchid family tree. [Note added Nov 17 2015: The flowers look like Lepanthes flowers because they use the same pollination method: pseudocopulation. The distinctive sheaths apparently evolved independently in the two groups as good solutions to some unknown environmental challenge in wet cloud forests.] Scientists recently proposed the replacement names Brachycladium and Oreophilus for this group, but both these names violate the careful rules of botanical nomenclature, and are hence illegitimate; they were replaced by the new name Neooreophilus which follows the rules and is legitimate. In a few days Mark, Alec, Sebastian, Frank Graham, and I will submit an extensive article about the correct phylogenetic placement of these orchids and their relatives. I’ll post a link here after acceptance.

Lou Jost