As part of my research on the nettle family, Urticaceae I became aware of plants growing in the entrance caverns of caves several years ago and for over a year now my collaborators at the Guangxi Institute of Botany, China lead by Professor Yigang Wei and I have been working on documenting the full diversity of this unusual flora. This lead us to think about whether these plants may have become adapted to life in caves, in particular the relatively constant climate and low light. Especially for species which grow amongst the lowest light levels at the back of caverns where they are growing in a fraction of the light they could be expected to receive in a forest. We therefore applied for a grant from the Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrain, and the Foreign Experts Bureau to undertake some preliminary work to document the climate, light and photosynthesis of the plants in the caves.
We selected the Yangtse cave as we know the diversity of plants that grow there (ten species of nettle alone), we have three data-loggers recording temperature and humidity in it and it is close to a town where we can spend the night. It is also a spectacular and beautiful place to spend several days working. The aim of our work was to collect the data necessary to test the hypothesis that the plants growing within the entrance cavern of the Yangtse cave exhibit different photosynthetic performance than the same or congeneric species growing outside of the cave. To do this we randomly selected individuals of three species of nettle in the genus Elatostema, one species of Begonia and a species of fern at four different locations in the cave, the back, midway into the entrance cavern, at the entrance and outside of the cave. We also brought two species of Elatostema from the living collection at the Guangxi Institute of Botany to compare their photosynthesis performance with members of the same species that had grown up within the cave. This was to get some indication as to how plastic their response was.
Each plant was connected to a hand-held PAM chlorophyll fluorometer, an incredibly sensitive device that can measure several key outputs of photosynthetic reactions in the chloroplasts as they take place. By comparing our study plants to those growing outside of the cave and from the scientific literature we hope to see whether cave-dwelling plants differ from non-cave plants in some of those parameters, and whether those differences are dependent upon what kind of plant they are. These parameters include the efficiency of photosynthesis, that is how much of the light energy is harnessed by the photosynthetic reactions, how much is dissipated and how resilient the photosynthetic apparatus is to changing light intensity. If we find a difference between cave and non-cave dwelling plants then taken together these measurements can provide some indication of which group of photosynthetic reactions are leading to these differences.
Working with Royal Botanic Gardens, Edinburgh student Jia Dong and plant anatomist Louis Ronse De Craene has resulted in some exciting and thought-provoking images of nettle flowers. The aim of our collaboration is to understand how nettle flowers develop and in the process work out what parts they have in common and which they don’t. The samples used were from living collections at Edinburgh and RBG Kew, together with my own collections in alcohol made over several years. The results are some beautiful and very informative scanning electron micrographs which show that the part of the female flower which recieves pollen (stigma) and conducts it to the egg (style) is characterised by two classes of hairs, one comprising defensive tusk-like hairs (above) and the other receptive tubular like hairs (above & below).
Tubular hairs associated with the stigma are characteristic of all nettle flowers. They also appear very early in development. Combined this makes us think that they might have a role in pollination. Specifically in the reception of pollen. Being wind-pollinated, nettles don’t have a lot of control as to whose pollen reaches their female flowers and so there needs to be a way for them to control which pollen grains develop and fertilise the single egg. It seems likely that these hairs play a role and hopefully Jia will be able provide some more great images to test some hypotheses about this.
One important task for specialists in a particular plant group, in my case nettles, is to visit national or regional collections and not just rely on the collections of our own institutes, no matter how good they are. As part of a conference and field trip I have just spent a couple of days in the La Paz herbarium. My colleague Nicholas Hind will spend three weeks there identifying plants from the daisy family (Compositae) and running an identification course. There were 92 boxes of unidentified Compositae waiting for him when we arrived!This herbarium was founded by German botanist Stephan Beck in 1984 and currently houses over 400,000 herbarium specimens. The reason why such visits are important both for the specialist but also for the herbarium are that although there is an active inter-herbarium loan system for plants it relies on material being accessioned, mounted and identified. This can be a real challenge to achieve in a country where there are few funds and even fewer botanists. The visit of a specialist allows material that has not been mounted or accessioned, together with unidentified material, to be reliably (one hopes) identified and so used as reference for future identifications. Continue reading Herbarium visit to La Paz (LPB)→
Funded by the Synthesys project I am studying the Cuban nettle collections of the Berlin-Dahlem Museum. The aim is to finish my account of the nettle family for the Flora of Cuba project that I started five years ago. It might seem odd that Cuba has so many nettle species that I can still be working on it, albeit in a fragmented way, for five years. Also that Berlin should be an important repository of Cuban plants, but there is a reason.
Cuba has an exceptionally rich flora and is especially important for the nettle family, Urticaceae, which is represented by about 70 species in eight genera. Together the Greater Antilles, Jamaica, Haiti/Dominican Republic, Cayman Islands and Puerto Rico, is a centre of species-richness for one group of nettles in particular, the genus Pilea. There are over 150 species of Pilea in the Greater Antilles, 60 of which are native to Cuba. The reason for such high diversity is unknown but may have something to do with the age of the islands, preponderance of limestone substrates or something else that we haven’t thought of yet. It does, however mean that I have been spending a lot of time looking at herbarium spcimens of this genus in the Berlin herbarium.
The connection between Germany and the the Greater Antilles started with botanist Ignatz Urban, who did a lot of work documenting the plant diversity of the Greater Antilles. Tragically most of his collections were destroyed in World War Two when the herbarium was bombed. During the Cold War the connection that had established between Berlin and the Caribbean switched to East Germany and the Jena herbarium impulsed by the formidable Johannes Bisse who founded the National Botanic Garden of Cuba. This lead to the foundation of the Flora of Cuba project a collaboration between the many excellent Cuban botanists and their German counterparts, initially in Jena and then from 1993 in Berlin. So that is the reason that I am in Berlin looking at Cuban nettles!
Nettles are characterised, amongst other things, by having flowers with a single egg in their ovary and a single stigma, the structure which conducts the pollen to its target. Work by developmental biologists almost a century ago suggested that the ancestor of nettles probably had two eggs per ovary after discovering that at a very early stage of development nettle ovaries contain two eggs one of which disappears as the flower develops resulting in the single egged flower which characterises the family. It was therefore a great surprise when plant collections from Costa Rica examined in the 1990s were found to have flowers with two or three eggs and stigmas per ovary. These very unusual plants were described as a new species: Boehmeria burgeriana by colleagues Melanie Wilmot-Dear and Ib Friis. Continue reading Ghost flowers in the nettle family→
I first came across this tiny creeping herb in my garden where it had been planted as an ornamental. The bright green leaves, mostly less than 3 mm across form an attractive carpet. Until now I had never been able to spot its flowers despite having checked several times over the last few years. My guess is that this species has a relatively narrow flowering time in spring and the flowers are so tiny that they are only visible with a hand-lens. For several years the genus has intrigued me, not so much because of its small size and creeping habit but because of its distribution and evolutionary relationships.
Firstly because Soleirolia consists of a single species that in the wild is only known from the Mediterranean island of Corsica. This is the only genus of nettle I know that is restricted to a single island or to the Mediterranean and I am very keen to try and found out why this could be (the history of the Mediterranean basin is quite a turbulent one). Secondly Soleirolia has traditionally been grouped with the widespread Parietariaandintriguingly with Gesnouinia, which also includes a single species but is restricted to theCanary Islands in the Atlantic Ocean. Whilst they look very different as plants their female flowers share many similarities of form. It might be, therefore, that Soleirolia and Gesnouinia should be viewed as Parietaria species that have diverged morphologically as a consequence of being isolated on islands, a common phenomenon in evolutionary biology. I am currently testing this possibility using DNA sequence data and could have a better idea in a couple of months.
Last week I was lucky enough to be invited to present a seminar on the nettle family, the Urticaceae at the Royal Botanic Garden Edinburgh. I have been working on this group of plants for over fifteen years with various collaborators and am finally ready to publish a revised classification of the family. I had a good audience who asked some insightful questions as well as making some good suggestions for future research. You can see a pdf of my slides by clicking on the following link: Edinburgh Urticaceae 19-6. Afterwards I had a chance to talk to a number of colleagues that I would like to start collaborating with in the future.
For over a hundred years the genus Myriocarpa in the nettle family which comprises ca 15 tree species in South and Central America has been impossible to place within the family. This is largely because of the very unusual shape of the part of the female flower that receives pollen, known as the stigma (see image below) and the fact that neither of the two great experts could agree over whether the petal-like structures at the base of the flower were petals associated with the flower, or bracts associated with the stalk. Whilst this might not seem like the stuff to keep a botanist awake at night it has become of interest again as using DNA data we have identified as sister to another small group of trees, Gyrotaenia, found in Cuba, Hispaniola, Jamaica and Central America that has a flower which consists of the petals fused to form a tube which is fused to ovary.
I was therefore very curious to see whether Vladimir Blagoderov, Manager of the Museum’s Sackler Imaging Suite could help me generate an image that would help us resolve the mystery. He could! The two images above are each composed of about 20 images which ‘slice’ through the sample which was of young flowers collected in alcohol in Belize over 10 years ago. The resolution was amazing, each cell being visible. In fact you could even make out the rough crystalline structure on the surface of the hairs! Both of these images also helped us to answer the question, revealing that this flower does indeed consist of a tube composed of fused petals that is subsequently fused to the ovary. This we could see in both images where the clearly visible ovary is enveloped by another distinct tissue, as in the case of Gyrotaenia. It was also confirmed by the petal-like bracts at the base of the flower having stalked glands, structures not known to occur on the ovaries of Urticaceae. So a morning’s work and an idea of evolutionary relationships enabled me to answer a question that had been frustrating an albeit very small group of botanists for over 100 years!
Thanks to funding from the Bentham Moxon Trust and the Guilin Botanical Garden, myself and colleagues explored five caves for plants in October of this year (2014). There are likely thousands of caves in the limestone karsts of south-east asia which contain plants. Whilst of great interest botanically and for conservation they are also beautiful in their own right and each cave is unique. I thought I would provide a portrait of each one to show how varied they are in their form, where plants grow in them and their size.
The cave above was one of the few that we spotted from the road and then were able to get to. It is also one of the few that had little evidence of human disturbance, very few footprints and areas of pure white travertine that had fallen from the roof had not been walked on or collected. The main plant-bearing cavern of the cave was about 25 m deep and the roof 15 m high and it had a well developed flora, you can see plants from the african-violet family (Gesneriaceae) in the foreground and we collected five species of nettle here.
Xiangshuidong Tian Keng cave in Guizhou was one of the largest caves that we have collected in, but also one of the most impacted by tourism and use by local communities. The main cavern is about 250 m deep with a roof between 45 and 30 m high and it is set within a huge cliff forming the side of a mountain. It also has a waterfall and river running across the back of it. It is in this cave that we found a very rare and unusual form of Elatostema oblongifolium that has its male flowers borne on specialised shoots but overall the plant diversity of the cave was quite low, presumably because of the large numbers of local tourists and associated trampling of much of the ground available for plants
This was the first cave that we encountered on this field trip. We found it after first being taken to a hole in the ground as what must have been a mistranslation from Mandarin into the local dialect. The cave was relatively big and had a trail running inside. The main cavern shelves very steeply meaning that very little light penetrates into the cave. Despite this and the relatively high altitude, 1500 m, we collected seven species of nettle from here.
This cave should have been perfect as it shelved gently meaning that light penetrated quite deep, it also had plenty of places for plants to grow, such as boulders and rocks. We only collected four species of nettle here, probably because the cave was heavily impacted by farmers using it as a barn to keep their water buffalo in at night. Evidenced by lots of hoof prints and dung. This is a very common use of caves and the trampling of buffalo and their herders can have a significant impact on the plants in the caves (see below).
Nettles have unisexual flowers, that is each flower functions only as a male or a female. Counter-intuitively though the flowers still retain the non-functional and often much-reduced organs of the non-functional sex. These are called pistillodes in the case of the rudimentary female organs in male flowers and staminodes in the case of the rudimentary male organs in female flowers. Inpart of the nettle family the staminodes are put to good use: ejecting the seed from the fruit.
This can be best understood by considering that nettles have male flowers which open explosively, pollen being released in tiny clouds (they are also very small). In fact one species, Pilea microphylla, is commonly known as the artillery plant for this reason. The mechanism for the explosive opening of the male flower is that the stalks (filaments) of the pollen sacks (anthers) are folded in on themselves in bud. As they develop these stalks fill with water until they are all pressing against each other within the flower bud and ready to burst. At a certain point the pressure becomes too great for the thin petals of the flower bud and they rip leading to the stalks being able to straighten explosively. This has happens incredibly quickly and although it has not been recorded for nettles, in the closely related mulberry where this also happens, the flower can open in 25 millionths of a second, moving petals to velocities in excess of half the speed of sound.
In one group of nettles which includes seven genera this explosive ability of the pollen sacks has been harnessed to release the seed from the fruit. Below you can see a close-up of the fruit of Pilea japonica showing the staminodes flexed and ready to eject the seeds (dark coloured).
Alex Monro's blog about the documenting and conservation of biodiversity