Guest post: The fascination of trees


[Editor’s note: This guest post is by Lane Davis. Lane is a former School for International Training student who spent a semester in Ecuador and did her independent study project with us. She then won a Fulbright scholarship to return to set up three research plots in our Cerro Candelaria Reserve,  at 2000m,  2500m, and 3000m. By identifying every tree in each plot, she has generated data which can help us quantify and understand not only the diversity of our forests, but also the important differences in composition between our forests at different altitudes, and between the Cerro Candelaria forests and others locally and globally. This kind of data provides a much-needed step towards understanding the deeper underlying causes  biodiversity – LJ]

[Traduccion a Espanol abajo]

Photos courtesy Lane Davis unless otherwise noted.

“And this?” Javier asks with anticipation as he opens the folded newspaper sheet labeled #47. I open my warped, mud-covered Rite in the Rain field notebook and look up the number. “Canopy tree, no latex or odor but the bark slash oxidized from white to brown. Do you want to see the live photos?” I ask. Javier shakes his head no and picks up a hand lens. I do the same and we each lift into the light a pressed and dried branch and examine it with our hand lenses.

Under the 30x magnification, the underside of the leaf shimmers with thousands of little scales. “What is it?” I ask him. Javier shrugs his shoulders almost jubilantly, muttering “Incredible,” and places the sample in a growing stack of unidentified plants. Later, we will scour Alwyn H. Gentry’s cinder block of a book “A Field Guide to the Families and Genera of Woody Plants of Northwest South America” and Walter Palacio’s “Árboles del Ecuador” (Trees of Ecuador) for families and genera of dicots with simple, opposite, alternate leaves; entire margins; and peltate trichomes (those shimmery scales) that could match sample #47. In this way, we will shrink the unknown stack, labeled “Desconocidos,” moving each plant we identify instead to piles of taxonomically related plants. But we will only make significant headway into the “Desconocidos” stack when we meet with another botanist, Walter Palacios. Yes, the same Walter Palacio’s I mention above who quite literally wrote the book on identifying trees in Ecuador. Javier and Walter are friends. Ecuador is a small country and its scientific community smaller, so pretty much all botanists know one another (which made it a little embarrassing when I asked for Walter’s signature on my copy of “Árboles del Ecuador,” but it was worth it).

But for now, Javier plunges back into the samples we haven’t reviewed at all yet. He grows more incredulous yet ecstatic each time he peels open one of the newspapers in which I have carefully pressed and dried tree clippings.  Sometimes he takes one look and proclaims the tree’s family, “Fabaceae” or “Lauraceae,” or even the genus, “Inga” or “Ocotea,” and I record this proclamation in my Microsoft Excel database and in the corner of the newspaper. But around half the time the sample remains with only a number to identify it.



One of my samples identified to family level. This is a member of one of the largest neotropical plant families, the Melastomataceae.


This uncertainty thrills Javier, a talented botanist, biologist, and the Executive Director of Fundación EcoMinga, the conservation organization I am affiliated with for my Fulbright work and which owns the forest where my dried tree clippings once grew. He has spent an unknowable number of hours traipsing through Ecuadorian forests; if he doesn’t recognize the plant, it must be at least somewhat rare. Javier also gets excited any time my pile of pressed plants yields a species he hasn’t seen in my samples yet, regardless of whether or not he knows what it is. With the discovery of each unique species, tree diversity goes up. The diversity of my plot, the 40m x 40m section of the forest where I gathered my plant samples, goes up in an absolute sense – one definition of diversity is simply the number of species present in a given area. But the implied diversity of the forest surrounding my plot shoots up even faster. My small plot cannot possibly capture the full diversity of the cloud forest, but we can use my data to estimate it. This calculation is based on the number of singletons, or species for which we have found only one individual tree in the plot. If singletons make up a large portion of the data, then we know the data isn’t representing the forest’s diversity well and there must be many yet undiscovered species outside of my plot. (For more on these calculations, see Chao and Jost 2012 and Chao et al. 2014).

For my part, the identifications and repetitions of plant groups are just as exciting as the unknown and new species; with each familiar sample and identifiable family characteristic, my own ability to identify cloud forest trees expands and solidifies. Unlike Javier, I have spent a knowable number of hours in the Ecuadorian cloud forest – to date, about 275 (not including evening and night hours when I slept in the field). Almost all of this time I spent collecting the plants piled in front of us, or walking to one of my three plots to do so.

During data collection, I lived in the 250-person village of El Placer at the base of Cerro Candelaria, the forest reserve owned by Fundación EcoMinga where I collected the now pressed and dried tree samples. Each morning I set out at 7:00 am, often but not always accompanied by a guardabosque (a forest ranger), and hiked to one of my three plots in the reserve. When I wrote my Fulbright grant proposal to study the vulnerability of Andean cloud forest trees to climate change, I planned to do so by learning about the altitudinal distributions of different trees species using eight different 10m x 100m plots ascending the mountain slope in Candelaria. Species growing in only a narrow altitudinal band will likely have a tougher time keeping up with their ideal growing conditions – as climate change shifts those conditions upslope – than species that are adapted to the conditions in a large geographic range.  It quickly became clear that I would not have enough time in the 10-month grant period to take data in such a large area, and Javier and I decided to modify our methodology to match that of the Evaluación Nacional Forestal (National Forest Evaluation) taking place in 2018, which uses square plots. That way, the Ecuadorian Ministry of the Environment could use our data in their study, too.



Views from and of El Placer

As a result, each morning I left El Placer to arrive at one of three 40m x 40m plots, located at 2000 m (6562 ft), 2500 m (8202 ft), or 3000 m above sea level (9843 ft).  Beginning from 1400 m (4593 ft), my commute required 2hrs and a very steep 1969 ft elevation gain to my first plot, 3 hrs and a crushing 3609 ft gain to my second plot, or 6 hrs and a demoralizing 5250 ft gain to my third plot. Consequently, I often camped in the field when I worked at my second plot and always did at my highest plot.



Campsite in and views from my plot at 3000 meters (nearly 10,000 ft above sea level).

I hiked through the Andean cloud forest, which usually meant hiking through a forest submerged in clouds. Cloud forests exist on mountains near lowland sources of atmospheric moisture – usually the ocean but in this case the Amazon Rainforest. Prevailing weather patterns push this moisture up the slopes, where it cools and condenses into low-level clouds, mist, or rain, leading to the frequent presence of precipitation in one of these forms.



Clouds in the cloud forest.

When I first began taking data in the cloud forest, I had no idea how to identify the trees around me, and with good reason. Though I took Field Botany at Williams College and identified plants as part of my senior Biology thesis, there are only a little over 70 species of trees in the state of Massachusetts (Butler 2016). In comparison, 131 different species of trees exist in the 4,000 square meters (slightly less than 1 acre) of cloud forest I have examined. Working to identify these trees using my dried samples, photos, books, the internet, the collections at the National Herbarium (a library of preserved plant samples), and significant help from professional botanists, I have slowly learned to recognize the defining characteristics of my plots’ most common families, genera, and species. Now when I walk through the forest, morphological features of plants capture my attention, often provoking a scientific name to come to mind. Large conical stipules, ring scars, and latex scream Moraceae; interpetiolar stipules insinuate Rubiaceae; and petiolar sheaths with a sweet soapy smell proclaim their identity – Hedyosmum.


Photo: Fausto Recalde/EcoMinga.





Cloud forest diversity is not confined to its trees. Far from it. For example, in the past 10 years, around 40 new species of orchid and 10 new species of frog have been discovered in EcoMinga’s reserves in one relatively small section of the Ecuadorian cloud forest. Above, a few photos of the incredible non-tree diversity of the cloud forest.

These trees and the billions of organisms that live on, under, and around them, ranging from soil microorganisms to Howler monkeys, as well as the inorganic features of the landscape like rocks and soil, make up the cloud forest ecosystem. This intricate network provides critical services to the human populations that make their home in the Andes Mountains. For example, cloud forest soil and epiphytes (plants that live on other plants and draw water and nutrients from the air rather than the soil) filter and regulate the flow of the glacial water which services millions of people in rural and urban Andean communities (Anderson et al. 2011). The extensive cloud forest root system helps hold soil in place, preventing erosion and landslides (Anderson et al. 2011). Climate change will disrupt these and other services, threatening human and ecosystem health and safety. For example, more intense rains combined with tree die-offs will increase erosion and landslides, which threaten human safety and water supply. In Quito in 2017, a landslide blocked the city’s main water channel, leaving 600,000 people without water for several days (Manetto 2017). In El Placer landslides occasionally cover pipes and cut off water; in my six months living there, this occurred once. [Editor’s note: See my previous post.]

Disruption of water supply is just one example of the myriad potential ways climate change and the resulting deterioration of the cloud forest ecosystem may affect El Placer and other similar communities nestled in Andean valleys. Better understanding the cloud forest’s fate under climate change will allow for targeted approaches to climate change preparation, for instance by creating emergency water delivery systems. Given the imminence of climate change, however, it is critical to implement strategies that decrease vulnerability to a wide range of climate change outcomes. I recently wrote a paper for Fulbright’s Regional Enhancement seminar on the how Fundación EcoMinga and El Placer’s partnership may do just that. I argued that EcoMinga bolsters El Placer’s climate resiliency by providing economic activities to the community that are less likely to be impacted by climate change than those that are otherwise available to them.

The main way EcoMinga does this is by employing community members as forest rangers in its reserves. The forest rangers build and maintain trails and cabins, assist visiting scientists and students with their research, and serve as keen eyes that often discover new species and other interesting biodiversity. My own work would have been out of reach (literally) without the help of Darwin Recalde, Jesús Recalde, Tito Recalde, Santiago Recalde, Jordy Salazar, and Andy Salazar. These men climbed 30-meter tall trees to reach leaves and flowers at the very top – those same leaves and flowers that now sit preserved in the National Herbarium in Quito and that make up the rows of my datasheets with which I will try to say something about the forest’s future.


Darwin Recalde climbing a tree to cut a sample of its leaves.

In fact, this goal – to assess the forest’s future under climate change – has morphed throughout my grant period. As with any interesting scientific study, this one has produced more questions than it will answer. Based on the calculations I mentioned earlier, though I took samples from 73 different tree species in my lowest altitude and most diverse plot, these represent less than half of the total number of species in the forest at that altitude.  What other species does the forest in this area contain? What allows the most common species I found to thrive? How will climate change affect its strategy? How will the forest’s response to climate change compare with my predictions? Will adaption differ in different locations within the cloud forest? Do these responses correspond with different microclimates? How do other aspects of the tree’s environment, like soil type and slope, affect forest adaption?

Many of these questions will only be answerable with a long-term research project. I have recently learned that my work will become part of just that. Fundación EcoMinga and the Instituto Nacional de Biodiversidad (National Institute of Biodiversity, or INABIO) are beginning a long-term forest monitoring collaboration. The study will comprise a network of plots in the Ecuadorian cloud forest including my three, a few other existing plots in EcoMinga’s reserves, and several more yet to be established. Tree growth, climate, and forest composition will be monitored regularly in these areas, and the data from my 2017-2018 study will form the baseline to which future measurements from my plots will be compared. While EcoMinga and INABIO are still determining details, the research will shed light on many of the questions my study has produced. In addition to providing baseline data, there are other ways I can help move this project forward. For one, I am striving to make the R (a statistical program) code I am writing to analyze my own data easily reproducible so other researchers and students can use it for quick analysis of data from all the plots.

This is an aerial view of Lane’s Plot 1 at 2000m elevation in our Cerro Candelaria Reserve. We fly over the 40m x 40m plot in the first few seconds, and then continue down the ridge to hover above our research station. Video by Lou Jost.

This is an aerial view of Lane’s Plot 2 at 2500m elevation in our Cerro Candelaria Reserve. We break through the clouds and fly straight to the 40m x 40m plot in the first few seconds, heading upslope. Then we turn around and float slightly downslope over and past the plot. Video by Lou Jost

I can also help by recruiting more students to continue the study. So much exciting work remains to be done. In addition to expanding and monitoring my plots, ample opportunities to personalize the project exist. For instance, you (yes, you!) could explore using drone imagery to identify trees from the air, investigate the role of rodents in seed dispersal, study the timing of tree sexual reproduction (phenology), or look at the genetics of cloud forest tree diversity – and how each of these impacts the forest’s adaption to climate change. All of these are areas in which EcoMinga currently works or would like to pursue. Whatever interests you, you will find enthusiastic scientists in Ecuador to support you. And if none of this attracts you but you know of others who it might, please send this post along to them.

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Example of aerial images of my plots that could be used to identify trees. Thanks to Lou Jost and his excellent drone piloting for these images! 

Finally, we can all support EcoMinga, its work conserving the cloud forest, partnership with El Placer, and scientific collaboration with INABIO by donating to the Foundation through the Orchid Conservation Alliance (US), the World Land Trust (UK) and Rainforest Trust (US). (Make sure you specify that the funds are for EcoMinga.) Contact Lou Jost ( for more information about donating.

Thank you for reading! If you are interested in continuing this work and/or in hearing more about it, please do not hesitate to contact me:

Lane Davis

(404) 805-2234 (WhatsApp or iMessage only until I am back in the US on May 11, 2018)

The opinions and information reported here are my own and do not represent those of the Fulbright Ecuador Commission, the Fulbright U.S. Student Program, or the U.S. Department of State.



Anderson, E.P., Marengo, J., Villalba, R., Halloy, S., Young, B., Cordero, D., Gast, F., Jaims, E., and Ruiz, D. Consequences of Climate Change for Ecosystems and Ecosystem Services in the Tropical Andes. In Climate Change and Biodiversity in the Tropical Andes; Herzog, S.K., Martinez, R., Jørgensen, P.M., Tiessen, H., Eds.; Inter-American Institute           for Global Change Research (IAI): MOtevideo, Uruguay; Scientific Committee on Problems of the Environment (SCOPE): Amstelveen, The Netherlands, 2011; pp 1-19.

Butler, B. J. 2016. Forests of Massachusetts, 2015. Resource Update FS-89. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 4 p.

Chao, A., Gotelli, N.J., Hsieh, T.C., Sander, E.L., Ma, K.H., Colwell, R.K., and Ellison, A.M. 2014. Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecological Society of America 84 (1): 45-67.

Chao, A. and Jost, L. 2012. Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93:2533−2547.

Manetto, F. 2017. Un derrumbe deja a 600.000 personas sin agua potable en Quito.” El Pais, December 8. america/1512681483_601181.html.


Post de invitado: La fascinación de los árboles
Nota de editor: Este post de invitados es realizado por Lane Davis. Lane es una ex estudiante de la escuela de Capacitación Internacional que pasó un semestre en Ecuador e hizo su proyecto de estudio independiente con nosotros. Después ganó la beca Fullbright para regresar a configurar las parcelas de investigación en árboles en nuestra Reserva Cerro Candelaria, a 2000 m, 2500 m y 3000 m. Identificando cada árbol en cada parcela, ella ha generado datos que nos pueden ayudar a cuantificary entender no sólo la diversidad de nuestros bosques, pero también la importante diferencia en composición entre nuestros bosques a diferentes altitudes, y entre los bosques del Cerro Candelaria y otros local y globalmente. Este tipo de datos proveen un paso muy necesario para comprender las causas subyacentes más profundas de la biodiversidad. -LJ]
Fotografía de cortesía Lane Davis a menos que se indique lo contrario.
¿Y esto?, pregunta Javier con anticipación en cuanto abre la hoja de noticias #47. Yo abro mi cuaderno de campo Rite in the Rain deformado y cubierto de lodo, y busco el número. “Árbol de dosel sin látex ni olor, pero la corteza se oxida de blanco a marrón. ¿Quieres ver las fotos en vivo?” pregunto. Javier niega con la cabeza y toma un lente de mano. Hago lo mismo y levantamos cada uno hacia la luz una rama prensada y seca y la examinamos con nuestras lentes de mano.
Bajo la magnificación 30x, el envés de la hoja brilla con miles de pequeñas escamas. “Qué es?” Le pregunto. Javier se encoje de hombros casi jubilosamente, murmurando “increíble”, y coloca la muestra en una creciente pila de plantas no identificadas. Después, limpiamos  el bloque de ceniza de un libro Alwyn H. Gentry’s “Guía de campo de las Familias y Géneros de de plantas maderables del Noroeste de Sudamérica” y “Árboles del Ecuador” de Walter Palacios para familias y géneros de dicotiledóneas con hojas simples, opuestas, alternas, margenes enteros; y tricomas peltados (aquellos con escamas brillantes) que podrían empatar con la muestra #47. En esta forma, encogeremos las pilas desconocidas, etiquetadas como “Desconocidos” moviendo cada planta que identificamos en lugar de pilas de plantas relacionadas taxonómicamente. Pero solo avanzaremos significativamente en la pila de “Desconocidos” cuando nos reunamos con otro botánico, Walter Palacios. Sí, el mismo Walter Palacios que mencioné antes, quien literalmente escribió el libro de identificando árboles en Ecuador. Javier y Walter son amigos. Ecuador es un país pequeño y su comunidad científica aún más, casi todos los botánicos se conocen (lo que hace un poco vergonzoso cuando pregunto por la firma de Walter en mi copia de “Árboles del Ecuador”, pero valió la pena).
Pero por ahora, Javier vuelve a sumergirse en las muestras que no hemos revisado del todo. El se vuelve más incrédulo pero extático cada vez que abre uno de los periódicos en los que he presionado y secado cuidadosamente los recortes de árboles. A veces toma una mirada y proclama la familia del árbol “Fabaceae” o “Lauraceae” o incluso el género,  “Inga” o “Ocotea,” y yo registro esa proclamación en mi base de datos de Microsoft Excel y en la esquina del periódico. Pero al rededor de la mitad del tiempo, esta muestra se mantiene sólo con un número para identificarla.
Una de mis muestras identifican al nivel de familia. Este es miembro de una de las plantas neotropicales más grandes, la Melastomataceae.
Esta incertidumbe emociona a Javier, un talentoso botánico, biólogo y Director Ejecutivo de la Fundación EcoMinga, la organización de conservación a la que estoy afiliada con mi trabajo en Fullbright y a la cual pertenece el bosque donde mis recortes de árboles secos crecieron una vez. Él ha pasado un gran número de horas recorriendo los bosques ecuatorianos, si el no reconoce la planta, debe ser algo raro. Javier también se emociona cualquier momento que mi montón de plantas prensadas producen una especie que aún no ha visto en mis muestras, independientemente de si sabe o no lo que es. Con el descubrimiento de cada especie única, la diversidad de árboles aumenta. La diversidad de mi parcela, la sección de 40x40m del bosque donde recogí mis muestras de plantas, aumenta en sentido absoluto: una definición de diversidad es simplemente el número de especies presentes en un área dada.
Pero la diversidad implicada del bosque que rodea mi parcela de 40 x 40 m del bosque donde recogí mis muestras de plantas, aumenta en sentido absoluto: una definición de diversidad es simplemente el número de especies presentes en un área determinada. Pero la diversidad implícita del bosque que rodea mi parcela se dispara aún más rápida. Mi pequeña parcela posiblemente no puede representar toda la diversidad del bosque nuboso, pero podemos usar mis datos para estimarlo. Este cálculo esta basado en el número de singletons, o especies para las cuales hemos encontrado solo un árbol individual en la parcela. Si los singletons hacen una gran porción de datos, entonces sabemos que los datos no están representando bien la diversidad del bosque y debe haber muchas especies sin descubrir fuera de mi parcela (Para más de estos cálculos, ver Chao y Jost 2012 y Chao et al. 2014).
De mi parte, la identificación y repetición de los grupos de plantas son tan emocionantes como las especies nuevas y desconocidas; con cada muestra familiar y de característica familiar identificable, mi propia habilidad para identificar los árboles de bosque nublado se expande y solidifica. A diferencia de Javier, yo he invertido muchas horas en el bosque nublado ecuatoriano – a la fecha, cerca de 275 (sin incluir horas de tardes y noches que dormí en el campo). Casi todo este tiempo, lo utilicé colectando montones de plantas frente a nosotros, o caminando en uno de mis tres parcelas para hacer eso.
Durante la recolección de datos, viví en la villa de 250 personas El Placer en la base de Cerro Candelaria, la reserva de bosque propiedad de la Fundación EcoMinga donde colecté las muestras ahora prensados y secas. Cada mañana me pongo en camino a las 7:00 am, a menudo pero no siempre acompañado de un guardabosque, y escalamos a una de mis tres parcelas en la reserva. Cuando escribí mi propuesta para la beca Fullbright para estudiar la vulnerabilidad de los bosques andinos para el cambio climático, planeé hacerlo por aprendizaje de la distribución altitudinal de diferentes especies usando ocho parcelas diferentes de 10 x 100 m ascendiendo las faldas de la montaña en Candelaria. Las especies que crecen en sólo una banda altitudinal estrecha tendrán más dificultades para mantenerse al día con sus condiciones ideales de crecimiento – a medida que el cambio climático cambia esas condiciones cuesta arriba – que las especies que están adaptadas a las condiciones en un grán área geográfica. Este rápidamente se vuelve claro de mono que no tengo suficiente tiempo en los 10 meses del periodo de subvención para tomar datos en un área tan grande, y Javier y yo decidimos modificar nuestra metodología para que coincida con la Evaluación Nacional Forestal (National Forest Evaluation) que se llevará a cabo en 2018, que utiliza parcelas cuadradas. De este modo, el Ministerio de Ambiente de Ecuador podría usar nuestros datos en su estudio también.
Vista de y desde El Placer
Como resultado, cada mañana lejo El Placer para llegar a una de las tres parcelas de 40 x 40 m, localizados a 2000 m (6562 pies), 2500 m (8202 pies), o 3000 m sobre el nivel del mar (9843 pies). Empezando de 1400 m (4593 pies), mi conmutador requiere 2 horas y una ganancia de elevación muy empinada de 1969 pies a mi primera parcela, 3 horas y unos  aplastantes 3609 pies para ganar mi segunda parcela, o 6 horas y una  desmoralizante ganancia de 5250 pies a mi tercera parcela. Consecuentemente, a menudo acampo en el campo cuando trabajo en mi segunda parcela y siempre lo hice en mi trama más alta.
Lugar de acampada y vista de mi parcela a 3000 metros (Cerca de 10 000 metros sobre el nivel del mar).
Escalé a través de los bosques Andinos, los cual usualmente significa escalar a través de un bosque sumergido en nubes. Los bosques nublados existen en las montañas cerca de las tierras bajas de humedad atmosférica – usualmente el océano pero en este caso el Bosque lluvioso amazónico. Los patrones climáticos predominantes empujan esta humedad por las laderas, donde se enfría y se condensa en nubes de bajo nivel, niebla o lluvia, lo que conduce a la presencia frecuente de precipitaciones en una de estas formas.
Nubes en el bosque nublado.
Cuando empecé a tomar los datos en el bosque nublado, no tenía idea de como identificar los árboles a mi alrededor, y con buena razón. Aunque tomé Field Botany en Williams College e identifiqué plantas como parte de mi tesis de biología, solo hay un poco más de 70 especies en árboles en el estado de Massachusetts (Butler 2016). En comparación, 131 especies diferentes de árboles existen en los 4000 metros cuadrados (ligeramente menos de un acre) de bosque nublado que he examinado. Trabajando para identificar estos árboles, usando mis muestras secas, fotos, libros, el internet, las colecciones del Herbario Nacional (una biblioteca de muestras de plantas preservadas), y la gran ayuda de botanicos profesionales, he aprendido lentamente a reconocer las características definitorias de las familias, géneros y especies más comunes de mis parcelas. Ahora, cuando camino por le bosque, las características morfológicas de las plantas captan mi atención, a menudo provocando que se me ocurra un nombre científico. Las estípulas largas y cónicas, escamas en anillo, y látex gritan Moraceae; las estípulas interpeciolares insinúan Rubiaceae; y vainas peciolares con un olor dulce y jabonoso proclaman su identidad: Hedyosmum.
Diversidad del bosque no se restringe a los árboles. Lejos de eso. Por ejemplo, en los pasados 10 años, cerca de 40 nuevas especies de orquídeas y 10 nuevas especies de ranas han sido descubiertas en las Reservas de EcoMinga en una sección relativamente pequeña del bosque nublado Ecuatoriano.  Encima, unas pocas e increíbles fotos de la diversidad de plantas no arbóreas del bosque nublado.
Estos árboles y los billones de organismos que viven en, abajo, y alrededor de ellos, que van desde microorganismos del suelo hasta monos aulladores, así como las características inorgánicas del paisaje como rocas y suelo, forman un ecosistema nuboso. Esta red intrincada provee servicios críticos para la población humana que hace su hogar en las montañas andinas. Por ejemplo, el suelo de los bosques nublados y las epífitas (plantas que viven en otras plantas y recogen agua y nutrientes del aire en lugar del suelo) filtran y regulan el flujo del agua glacial que sirve a millones de personas en las comunidades andinas urbanas y rurales andinas (Anderson et al 2011). El extenso sistema de raíces del bosque nublado ayuda a mantener el suelo en su lugar, previniendo la erosión y los deslizamientos de tierra (Anderson et al 2011). El cambio climático interrumpirá este y otros servicios, amenazando la salud y seguridad humana y del ecosistema. Por ejemplo, las lluvias más intensas combinadas con la muerte de los árboles aumentarán la erosión y deslizamientos de tierra, lo que amenaza la seguridad humana y el suministro de agua. En Quito, en 2017, un deslizamiento de tierra bloqueó el canal principal de agua de la cuidad, dejando a 600 000 personas sin agua por muchos días (Manetto 2017). En El Placer, los deslizamientos de tierra ocasionalmente cubren tuberías y cortan el agua, en mis seis meses viviendo ahí, esto ocurrió una vez.
La interrupción del suministro de agua es sólo un ejemplo de innumerables formas posibles en que el cambio climático y el deterioro resultante del ecosistema del bosque nuboso pueden afectar a El Placer y otras comunidades similares enclavados en los valles andinos. Mejor entendimiento del destino del bosque nuboso bajo el cambio climático permitirá enfoques específicos para la preparación del cambio climático, por ejemplo, creando sistemas de suministro de agua de emergencia. Dada la inminencia del cambio climático, sin embargo, es crítico implementar estrategias que disminuyen la vulnerabilidad de un amplio rango de resultados de cambio climático. Recientemente escribí un paper para el seminario Mejora Regional Fullbright acerca de cómo Fundación EcoMinga y El Placer pueden hacer exactamente eso. Argumenté que EcoMinga refuerza la resiliencia climática de El Placer al proporcionar actividades económicas a la comunidad que tienen menos probabilidades de verse afectadas por el cambio climático que aquellas que de otro modo estarían disponibles para ellos.
La principal forma en que EcoMinga hace esto es mediante los miembros de la comunidad como guardabosques en sus reservas. Los guardabosques construyen y mantienen senderos y cabañas, ayudan a los científicos y estudiantes visitantes con su investigación y sirven como ojos agudos que a menudo descubren nuevas especies y más biodiversidad interesante. Mi propio trabajo hubiera estado fuera de alcance (literalmente) sin la ayuda de Darwin Recalde, Jesús Recalde, Tito Recalde, Santiago Recalde, Jordy Salazar y Andy Salazar. Estos hombres escalan árboles altos de 30 metros de alto para alcanzar hojas y flores en la cima – aquellas mismas hojas y flores que ahora se preservan en el Herbario Nacional en Quito y eso forma las filas de mis hojas de datos con las cuales trataré de decir algo sobre el futuro del bosque.
Darwin Recalde escalando un arbol para cortar muestras de sus hojas.
De hecho, este objetivo – evaluar el futuro del bosque bajo el cambio climático se ha transformado a lo largo de mi periodo de beca. Como en cualquier estudio científico interesante, este ha producido más preguntas de las que responderá. Como en cualquier estudio científico interesante, este ha producido más preguntas de las que responderá. Con base en los cálculos que mencioné antes, aunque tomé muestras de 73 diferentes especies de árboles en la parcela de menor altitud y mayor diversidad, esto representó menos de la mitad del número total de especies en el bosque a esa altitud. ¿Qué otras especies contiene el bosque en esta área? ¿Qué permite a las especies más comunes prosperar? ¿Cómo afectará el cambio climático a su estrategia?  ¿Cómo se comparará la respuesta del bosque al cambio climático con mis predicciones? ¿Difiere la adaptación en diferentes localidades dentro del bosque nublado? ¿Estas respuestas se corresponden con diferentes microclimas? ¿Cómo afectan otros aspectos del entorno del árbol, como el tipo de suelo y la pendiente, la adaptación del bosque?
Muchas de estas preguntas sólo son respondidas con un proyecto de investigación de largo tiempo. Recientemente he aprendido que mi trabajo será parte de eso. Fundación EcoMinga y el Instituto Nacional de Biodiversidad (INABIO) están comenzando un monitoreo colaborativo del bosque de larga duración. El estudio incluirá una red de parcelas en el bosque nublado ecuatoriano incluyendo mis tres, otras pocas existiendo parcelas en las reservas EcoMinga, y muchas mñas aún por establecerse. El crecimiento de árboles, clima y composición del bosque será monitoreada regularmente en estas áreas, y el dato de mi estudio en 2017-2018 será la línea base para la cual las futuras medidas de mis parcelas serán comparadas. Mientras EcoMinga e INABIO están determinando detalles, la investigación arrojará luz sobre muchas de las preguntas que ha producido mi estudio. En adición a proveer la linea base hay otras vías por las cuales puedo ayudar a avanzar este proyecto. Por ejemplo*, me esfuerzo por hacer que el código R (un programa estadístico) que estoy escribiendo analice mis propios datos facilmente reproducibles de modo que otros investigadores y estudiantes pueden usar para análisis rápidos de datos de todas las parcelas.
Puedo ayudar reclutando más estudiantes para continuar el estudio. Mucho trabajo empocionante falta por hacer. En adición a expander y monitorear mis parcelas, existen amplias oportunidades para personalizar el proyecto. Por ejemplo, tu (sí, tu!) puedes explorar usando imágenes de dron para identificar árboles desde el aire, investigar el rol de los ratones en la dispersión de semillas, estudiar la sincronización de la reproducción sexual de árboles (fenología), o mirar la genética de la diversidad de árboles de bosque nublado – y cómo cada uno de estos impacta la adaptación del bosque al cambio climático. Todas estas son áreas en las cuales EcoMinga trabaja normalmente o le gustaría adquirir. Cualquier interés que tengas, encontrarás científicos entusiastas en Ecuador para apoyarlo. Y si nada de esto te atrae pero conoces a otras personas a las que podría atraerles, envíales esta publicación.
Finalmente, podemos apoyar a EcoMinga, su trabajo conservando el bosque nublado, en asociación con El Placer, y colaboración científico con INABIO donando a la Fundación a través de la Alianza para la Conservación de Orquídeas (US), the World Land Trust (UK) y Rainforest Trust (US). (Asegurate de especificar que los fondos son de EcoMinga). Contacta a Lou Jost () para mas información sobre donaciones.
Gracias por leer! Si estás interesado en continuar este trabajo o en escuchar más acerca del mismo, por favor no dude en ponerse en contacto conmigo:

Lane Davis

(404) 805-2234 (WhatsApp o iMessage sólo hasta que regrese a los EEUU en Mayo 11, 2018)

Las opiniones e información reportada aquí, son de mi propiedad y no representan aquellas de Fulbright Ecuador Commission, the Fulbright U.S. Student Program, o la U.S. Department of State.

Traducción: Salomé Solórzano Flores


Carnegie Airborne Observatory visits our area


Carnegie Airborne Observatory image of rainforest trees; different colors represent different spectral fingerprints. Click picture to enlarge. Image: Carnegie Institution for Science.

The Carnegie Institution for Science is a unique private organization devoted to advanced study of the earth, life, and the universe. The pioneer cosmologist Edwin Hubble (“Hubble constant”), geologist Charles Richter (“Richter scale”), geneticist Barbara McClintock, and many Nobel laureates from several different disciplines are or were Carnegie investigators. The institution has instruments orbiting Mercury, is a lead partner in constructing the world’s biggest telescope in Chile, and has one of the world’s most sophisticated ecological monitoring devices, the Carnegie Aerial Observatory (CAO). This is a two-engine 20-passenger plane that Greg Asner and colleagues has fitted with millions of dollars worth of specially-designed lasers and spectrometers. It can sample hundreds of thousands of hectares of forest per day, using LIDAR to build a 3-dimensional model of the forest’s trees with 8 cm resolution. At the same time as it acquires LIDAR data, it also samples the spectral properties of light reflected from the vegetation, gathering reflectance information at hundreds of different wavelengths (colors). This spectral data gives information about the chemical and physical properties of the leaves, and also provides a spectral fingerprint that can later be matched to field-collected spectral fingerprints from known species of trees. Some  trees have such distinctive fingerprints that they can be identified to species with this data; more commonly, they can be identified to genus, though sometimes only to family. The detailed structural, chemical and taxonomic data acquired by the CAO would be impossible to gather at the landscape level by any other method, and Greg’s work is dramatically expanding the range of questions that ecologists can ask about forest ecosystems.


Carnegie aerial observatory rainforest image: 3-D Lidar combined with spectral signal. Image: Carnegie Institution for Science.

Last year Greg had planned to use our mosaic of forests as reference sites for a study of Andean forests on different geological substrates and elevations.  Greg and his partner Robin Martin visited our Rio Zunac Reserve, his flight plans got approved by the Ecuadorian authorities, and everything seemed ready to go, but in the end he was not allowed to bring the plane into the country. This year, however, Greg was able to bring the plane in for a more modest ten-day study of Amazonia. The plane’s home for those ten days was the military base in Shell, a town in the upper Rio Pastaza watershed near our Rio Anzu Reserve. One of the CAO’s flight transects covered a two-kilometer wide strip from west to east (high to low) through our area, perhaps including parts of up to four of our reserves. This will be a very valuable data set that will teach us a great deal about the structure and diversity of these forests. However, it will take about a year to fully process the data, so we’ll have to be patient.


The Carnegie Airborne Observatory parked at the Shell military base. Our reserves are in the mountains in the background. Click picture to enlarge. Photo: Matt Scott.

The president of the Carnegie Institution for Science, Matt Scott, is a well-known geneticist and serious photographer. He came t0 Ecuador last month to fly with Greg, but first he wanted to visit some of our reserves. Our endangered Black-and-chestnut Eagles (Spizaetus isidori) were nesting again in our Rio Zunac Reserve after last year’s tragic nest failure, so this was a once-in-a-lifetime opportunity to observe the species as it went about its business.

I picked him up in the Quito airport. The trip from Quito to Banos was picturesque as always. The glacier of Cotopaxi was covered in a layer of fresh volcanic ash, and small puffs of ash and vapor were still rising up from the crater as we drove past it.


Cotopaxi’s glaciers covered in fresh ash. Click picture to enlarge. Photo: Matt Scott.



Close to sunset as we neared Banos after passing through a rainstorm. Click picture to enlarge. Photo: Matt Scott.

The next day we had an appointment with the Black-and-chestnut Eagles at 10am-11am. Our guards told us the parents  usually brought prey to the baby at that time, but were otherwise rarely seen around the nest. The nest is about 3-4 hours away from the road, after a forty minute drive from Banos, so we had to get up early and rush out there. It was hard to keep up  a good pace, since beautiful things kept distracting us. Still, we managed to get to the nest observation spot at almost exactly 11:00, and sure enough, there was the adult in the nest, along with the chick and something dead. The adult flew off almost immediately but shortly returned to feed on the prey item while the sated chick slept. The other adult was also nearby and both called frequently. We spent an hour watching them. It was a wonderful thing to see.


This was the view when Matt got to my house to start our trip to the Rio Zunac. Volcan Tungurahua with a lenticular cloud against a crystal sky, a great way to start the day. Click picture to enlarge. Photo: Matt Scott.


Morning fog over the Rio Pastaza. Click picture to enlarge. Photo: Matt Scott.


Black-and-chestnut Eagle (Spizaetus isidori) at its nest in our Rio Zunac Reserve. Photo: Matt Scott.


We saw several Highland Motmots. Photo: Lou Jost/EcoMinga.


Torrent Ducks on the Rio Zunac distracted us throughout the day. Click picture to enlarge. Photo: Matt Scott.


We found this crazy katydid at the end of our walk. Click picture to enlarge. Photo: Lou Jost/EcoMinga.


Butterflies and hesperids taking salts from the sand along the Rio Zunac. Photo: Lou Jost/EcoMinga.


Matt chills out in the Rio Zunac after our hike. Click picture to enlarge. Photo: Lou Jost.


The next day we went to our Rio Anzu Reserve near the Shell airport and the CAO. That reserve is not very rich in big stuff, but there are so many interesting small things that it is hard to take ten steps without stopping for photos. We eventually got to the Rio Anzu river and the magnificent fossil-bearing limestone formations capped with ladyslipper orchids (Phragmipedium pearcei). Though it was getting late, Matt asked to stay longer. I always like to hear that from a visitor!!


Matt photographing the limestone. Click picture to enlarge. Photo: Lou Jost/EcoMinga.


The limestone formations along the Rio Anzu, covered with orchids. Click picture to enlarge. Photo: Matt Scott.


Phragmipedium pearcei, a ladyslipper orchid, on the limestone. Click picture to enlarge. Photo: Matt Scott.


Riodinid butterfly in the Rio Anzu Reserve. Click picture to enlarge. Photo: Matt Scott.


Large hairy caterpillar. Click picture to enlarge. Photo: Matt Scott.


Me in bamboo forest along the Rio Anzu. Click picture to enlarge. Photo: Matt Scott.

Then we went to the military base to see the CAO. Security was tight and the military were not eager to let a pair of muddy rubber-booted gringos walk through their installations. Nevertheless we were able to talk our way through the multiple layers of officials who scrutinized us. But we didn’t want to ruffle any feathers so when we finally got to the plane, we just took a quick look at it and went back (still under military escort, but actually a very friendly one).


CAO at the military base. Click picture to enlarge. Photo: Matt Scott.

By the time we got to Greg and Robin’s hotel in nearby Puyo it was already dark. Greg was sitting at a table outside working on maps in his laptop, and he showed me the transects he had flown so far. I went back to Banos that night but Matt stayed and got to fly in the CAO over the following days. Lucky man!


Matt (left) and Greg happy to be in the air. Click picture to enlarge. Photo: Matt Scott.


The Rio Pastaza broadens and meanders as it leaves our mountains and enters Amazonia. Click picture to enlarge. Photo: Matt Scott.



The Amazon basin from the CAO. Click picture to enlarge. Photo: Matt Scott.



More of the Amazon basin from the CAO. Click picture to enlarge. Photo: Matt Scott.

Matt, thanks very much for your visit! It was an honor for us to show you our forests.

Lou Jost

Fundacion EcoMinga



Earth Day: High school students from Aldo Leopold’s alma mater spend a week in our Cerro Candelaria forest

Aldo Leopold’s 1949 book, “A Sand County Almanac”, was one of the first voices of the environmental consciousness that began to awaken in response to the post-World War II rise of man’s destructive power. The founder of Earth Day, Wisconsin Senator Gaylord Nelson, was deeply influenced by his writings.

Aldo Leopold wanted humanity to develop a land ethic, one that respected plants and non-human animals. He wrote:

“When god-like Odysseus returned from the wars in Troy, he hanged all on one rope a dozen slave-girls of his household whom he suspected of misbehavior during his absence.”

“This hanging involved no question of propriety. The girls were property. The disposal of property was then, as now, a matter of expediency, not of right and wrong. Concepts of right and wrong were not lacking from Odysseus’ Greece: witness the fidelity of his wife through the long years before at last his black-prowed galleys clove the wine-dark seas for home. The ethical structure of that day covered wives, but had not yet been extended to human chattels. During the three thousand years which have since elapsed, ethical criteria have been extended to many fields of conduct, with corresponding shrinkages in those judged by expediency only.”

“…There is as yet no ethic dealing with man’s relation to land and to the animals and plants which grow upon it. Land, like Odysseus’ slave-girls, is still property. The land-relation is still strictly economic, entailing privileges but not obligations.”

Perhaps (just perhaps) our ethical sphere has been extended a bit since Leopold wrote those words, but we have a long way to go. We spend less and less time in nature, to the point where most people today do not even know what real nature is. Intact ecosystems are now so rare that the vast majority of people will never experience them, much less fall in love with them. This visceral love of nature is the only thing that can drive people to sacrifice their own comforts to protect it.

Dr John L. Clark, who holds the Aldo Leopold Distinguished Teaching Chair at The Lawrenceville School, Aldo Leopold’s alma mater in New Jersey, is as much in love with nature as anyone I know. He has started a program to bring his high school biology students (ranging in age from 15-18 years old) to our reserves in Ecuador, to try to ignite this passion for real nature in the next generation.

John is an old friend of mine who used to be a Peace Corps volunteer here in the 1980s. He is now a famous botanist specializing in gesneriads, the African Violet family. He has published several monographs on gesneriad genera and has discovered many new species. Two years ago, as a professor at the University of Alabama, he brought a college biology class to our Rio Zunac Reserve to set up two quarter-hectare plots, in which every tree bigger than 10 cm in diameter was sampled, tagged, and identified. Dr David Neill from the Universidad Estatal Amazonica helped set up that plot and identified the trees. In the process they found what turned out to be two new species of Magnolia trees, and John discovered a new gesneriad in the genus Columnea.

Now in his new position at The Lawrenceville School, he has done the same thing with a dozen of his high school students, joining with David Neill again to set up a quarter-hectare plot in our Cerro Candelaria Reserve last month. It was a daring project, very unusual for an American high school.

Some of his students wrote about their experience. Here is Kaimansa Sowah’s essay, which she titled “Botanizing!”:

“Never had I seriously considered ecology or botany or even entomology as a field of interest until our trip to Cerro Candelaria on the eastern slopes of the Andes in Ecuador. Arriving in Quito on a Saturday morning with many missionary groups crowding the lines at immigration, I questioned if our work in Ecuador would have any real impact on the community. How could plant identification transcend traditional community service? It would not be until I was sitting around a fire at our high camp sipping tea made from recently collected crushed foliage of a Lauraceae we had found earlier, barely communicating sufficiently in my middle school Spanish that I managed to realize the profound importance of our trip to Ecuador.”

“The hike up to camp was brutal to say the least. Many of us had never hiked before and mounted on our backs were 50-pound packs with silica gel for preparing museum specimens, M&Ms (which would be our lunch for several days), and personal belongings. Our frequent stops for “Botanizing!” only heightened the difficulty level. Our expedition leader Dr. John Clark lights up at a fallen Gesneriaceae leaf, so throughout the hike and the trip as a whole, he was never short of excitement as our paths were lined with rare and new species. Fortunately, the view of mountains perfectly scattered, parting only for the rapids leading to and from waterfalls, fuelled our strenuous walk to the camp. The view never ceased to amaze us, and many of us still fail to believe its reality.”

“It was not until we began work on the plots that each of our own individual love for botany and plant life was established. Divided into groups of three, we established and inventoried tree diversity in a 0.25-hectare permanent plot. With the help of Tito, our guide, friend, and resident tree climber, we identified trees based on vegetative features (e.g., leaf patterns, leaf arrangement, smell), recorded DBH (diameter breast height), tree height, and tagged each tree with an aluminum label. Our field journals appeared something like this: “tree 4, subplot 5, 25 meter height, 18 cm DBH, simple-alternate leaves with milky sap (Moraceae?).” On the first day we found a cherry tree (Prunus sp.) that had never been observed by our resident scientist and tree expert, Dr. David Neill who is a professor of biology at the Universidad Estatal Amazónica. Many of the trees were challenging to identify, which only further affirmed how much biodiversity surrounded us. During a lunch break, we played a plant identification game where we were divided into teams and given Al Gentry’s book “A Field Guide to the Families and Genera of Woody Plants of North west South America.” Each team was timed in their ability to identify foliage to family. All of us being extremely competitive, we quickly held our leaves to the light using our hand lenses, crushing and smelling, and rapidly blurting out names like “Piperacae!” Euphoribacae!” “Melostomatacae!””

“Along with our own Dr. Clark were resident entomologists and ecologists who shared their love of biology. We also met the director and founder of the EcoMinga foundation, Lou Jost who is a theoretical mathematician, ecologist, and botanist who specializes in the study of orchids. We were surrounded by vast amounts of unique talent, which greatly sparked our own interests. Besides the fieldwork, we were able to connect and talk with our guides. They soon became our friends, and it was through conversations with them that we realized how grateful they were for our interest in visiting their reserve. No, as a sixteen-year-old girl, I had never thought of biodiversity research as one of my interests. And I cannot say whether it was our guide giving us hints during the scavenger hunt with his ability to identify plant families from meters away, or the sheer look of ecstasy when “Ranger”, also known as Dr. Clark, and Dr. Neill sat around their pressed leaves dumbfounded at a new species, or Darwin [Recalde]’s ability to navigate the maze-like mountains and carting us up steep hills. Nonetheless, this trip has piqued my interest and I suspect that botany and biodiversity will play a large role in my future.”

Eloise White wrote of her experience:

“…When I first signed up to travel to Ecuador with the School, I expected a week of light hiking, bonding with new friends, and great food, all coupled with the occasional botanical reference. While the food was indeed fantastic, the intensity of the trip took us all by surprise on the first day in the field, when we embarked on a challenging four-hour hike to our camp. It was not until after we finished showering in the beautiful waterfall and sat down at dinner to prepare our field notebooks for our work in the tree plots the next morning that I realized the importance of the work that we would accomplish during our time in the forest.”

“When we reached the plots bright and early the next day, we received instructions, and my group quickly fell into a rhythm of tagging trees with bright orange tape and communicating with our local guides who were climbing to the canopy of the trees, a task that gradually became easier as our Spanish improved. Each time that our guide, usually some 30 feet high in a tree, would cry “Ten cuidado!” the three students in my group would jump back and wait for an unidentified specimen to come crashing to the ground. That first day, in the moments that I spent with Dr. Clark, tagging and pressing plant samples into pages of newspaper, his excitement surrounding new and rare species was absolutely contagious. I found myself eager to memorize the names of plant species, to identify which types of bark had latex, and to distinguish simple leaves from compound leaves. Even now, I find myself so grateful to Dr. Clark and the other scientists accompanying us in the forest because they showed me what it means to be passionate about a specific field of study, something that I hope to do as I move forward in my Lawrenceville career, the college process, and my life.”

“…My Spanish teachers at The Lawrenceville School have always stressed the importance of experiencing the language abroad in order to truly further my understanding… Between trying to ask our guides to scale a certain tree to obtain a specimen and sitting around our campfire late in the night, telling ghost stories and jokes with Jordi and Darwin, I was constantly speaking Spanish. The pure exposure to the language coupled with the locals’ willingness to help me practice provided me with a unique opportunity to further an area of interest which I had not previously devoted much attention to. Furthermore, partially overcoming the language barrier opened the group up to an irreplaceable chance to form lasting friendships with locals, a memory that I will forever treasure. Lawrenceville constantly stresses the importance of expanding our horizons, and I can attest that in communicating with and working alongside unfamiliar faces, the twelve of us expanded our own world views significantly.”

“Before embarking on our journey, our teachers made it clear that our accommodations would be far from luxurious. We were told us that we would be perpetually damp, sweaty, and dirty, all of which later proved true. However, I will be the first to say that the view from our wooden cabin base camp without windows, doors, or even walls was extraordinary, rivaling that from any mountain getaway or island. When we summited Cerro Candelaria (3800+ m), while it was extremely challenging and put both our bodies and minds to the test, the breathtaking outlook from the top instantly made our hard work worth it.”

“Overall, my work and experiences in Ecuador were once-in-a-lifetime opportunities. They opened my eyes up to an entirely new scope of interests, people, and awareness. For example, as I previously planned on dropping out of Spanish for my senior year, I have changed my mind and will continue to advance my understanding of the language, hopefully into college. As I begin the college search, I have been relentlessly pestering my counselor about which schools have the best programs to study abroad while working with the science department. I attribute these shifts in my interests to my recent experience in Ecuador.”

Vivienne Gao expresses the very real physical challenges of this trip:

“Honestly, if I had known our expedition to Ecuador involved so much hiking, I probably would not have signed up. I’ve always been more comfortable in the water; I prefer swimming over running and am generally more athletic when I am not on land, so the minute I found out that our first hike to low camp would take roughly four hours, I definitely had my doubts…The day was hot, but not unpleasant, but I still kept my hair in braids to keep it off my neck. Once we began our hike however, the physical exertion made the heat borderline unbearable. We all carried large Osprey backpacks with our personal belongings, and these bags were not only heavy but also didn’t breathe well. Sweat happily gathered between my back and my pack, soaking through my shirt so that when I finally peeled the pack off, my shirt still clung to me as a dog’s fur clings to it, dripping, rinsed after a soapy bath in the backyard. The hike was mostly uphill, but the terrain varied. We were slopping through mud, climbing over rocks, and wading through streams, sometimes on level ground and sometimes on downhill slopes, but everything led us upwards eventually.”

“I remember seeing the cabin for the first time after three or so hours of hiking and thinking that this was the best moment of my life. I had fallen behind with a couple friends, so the rest of the group was already in the cabin waiting for us. As I slowly trudged up the hill, humoring the impressive cramp in my right calf that had formed over the duration of the hike, I congratulated myself for completing the hike, a feat that I considered the most difficult thing I’ve ever done. Little did I know that in the days to come, I would experience hikes many times more difficult than this one, a prospect far beyond my wildest imaginations…”

“Twenty minutes before we reached low camp, my small group of hiking companions and I had come across waterfall, the same waterfall that would host our daily shower and laundry trips. After reaching camp, everyone, me included, was excited to wash the salt and dirt off their bodies. The idea of showering in a waterfall enticed me, but my legs caved at the thought of hiking another twenty minutes to the waterfall, yet I went anyways. The waterfall became my favorite place and I went everyday after that.”

“I came back from Ecuador having learned more about my physical and mental limits, surprised at how hard I could actually push myself. I lost eleven pounds but earned so much more in experience and memories. The trip is something I will never forget, and who knows, maybe someday I’ll return, ready to face the challenges I faced this time and conquer them.”

As his students noticed, John Clark was at least as excited as they were:

“I am often asked how I know when something I come across is a new species. It is important to note that describing a new species is a process that is collections-based, requires several formal criteria outlined by the International Code of Nomenclature (ICN), and is contingent on a peer-reviewed publication. It is considered by some biologists (e.g., L.E. Skog who co-chaired my PhD committee) as “bad botanical etiquette” to say something is new without data. Nevertheless, outlined here are four species that I am confident have not been previously described. My doctoral dissertation resulted in a monographic revision of Glossoloma (Clark 2005). This is a group of plants that I dedicated more than a decade studying and when finished, I expected that there would be an occasional new species that would represent something that was not included in the monograph (Clark 2005). For example, Karyn Cichocki observed a new species of Glossoloma in 2007 when assisting me on an expedition in Ecuador. An additional new species was described with a student as a result of an expedition in Colombia (Rodas & Clark 2014). What I did not expect to find in Cerro Candelaria was a new species of Glossoloma every 500 meters in elevation change. I found three new species of Glossoloma between our base camp and the high camp. We also discovered a an undescribed species of Drymonia, which is a group that Laura Clavijo and I have studied together for more than eight years. I directed Laura’s dissertation committee (2007 to 2015) and together we have published more than eight papers on Drymonia. Thus, the four undescribed species featured in Figure 1 [below] are based on ongoing studies of museum specimens, extensive fieldwork, and comprehensive review of taxonomic literature. The remarkable discovery of biodiversity featured in Figure 1 is an example of the urgency and need for additional studies in the Neotropics.”

“There are also rare species from Cerro Candelaria that I did not expect to find. Two collections represent populations that were not previously known. The rarest plant that we found was Columnea bivalvis (photo below, D and E), which was previously only known from a single population (Amaya-Márquez & Clark 2011). [Note added by LJ: That original population was found in what is now our Rio Machay Reserve.] Drymonia ignea (photo below, A and B) is endemic to the eastern slopes of the Andes and was previously only known from 5 populations (Clark 2013). Never have I seen more than a few individuals of Drymonia ignea growing together and along the ridgeline there were multiple areas of ten or more individuals.”

The Lawrenceville School students not only gave us their friendship and enthusiasm but also brought the gift of electricity to our research stations. I’ll save that story for a separate post.

The Lawrenceville School staff who visited us: Baptiste Bataille, Jennifer Mayr (her husband is related to famous evolutionary biologist Ernest Mayr!) and John L. Clark.

The Lawrenceville School staff who visited us: Baptiste Bataille, Jennifer Mayr (her husband is related to famous evolutionary biologist Ernest Mayr!) and John L. Clark.

The essay excerpts used here are from John’s fuller version of this story which will soon be published by the magazine “Gesneriads”. They are used here with John’s, the school’s, and the magazine editor’s permission. Thanks John, and thanks Lawrenceville School students, for a wonderful cultural exchange and exciting scientific discoveries! Your enthusiasm and that of your students inspires us and makes our work feel worth the trouble. Lawrenceville School students, you literally walk in the footsteps of the great Aldo Leopold, and I hope that like him, some of you can help the earth face the challenges that your own generation will witness.

EcoMinga also thanks the World Land Trust and their donors Puro Coffee, Naturetrek, and PricewaterhouseCoopers for funding the Cerro Candelaria Reserve, and their donor Noel McWilliam for the funds to build the research station where these students, and many other students and scientists, stayed.

The World Land Trust’s “Forests in the Sky” appeal continues to expand the protection of this area.

Lou Jost

Estimating the diversity of an ecosystem based on an incomplete sample

The genus Lepanthes is one of the most diverse plant genera in our area, with more than 100 species just in the upper Rio Pastaza watershed. We keep discovering more new species, even after twenty years of intense surveying. This is typical of tropical floras and faunas. It is a real challenge to try to estimate the diversity of a rich group of organisms like this. We need the help of clever statistical methods! Painting: Lou Jost.

The genus Lepanthes is one of the most diverse plant genera in our area, with more than 100 species just in the upper Rio Pastaza watershed. We keep discovering more new species here in our reserves, even after twenty years of intense surveying. This is typical of tropical floras and faunas. It is a real challenge to try to estimate the diversity of a rich group of organisms like this. We need the help of clever statistical methods! Painting: Lou Jost.

Ecologists and conservation biologists often need to quantify the diversity of some community of animals and plants. In the tropics, where there are many rare species, this can be hard to do, because there are so many rare species; a random sample from the population will miss many species. My friends Phil DeVries, Tom Walla, and Harold Greeney were still finding additional species even after five years spent systematically sampling the butterfly diversity of a single research site in the Amazon basin of Ecuador. This is typical of tropical habitats. So biologists need the help of mathematicians and statisticians to help estimate the true diversity based on incomplete samples.

The person who has done the most to advance this aspect of statistical theory is Anne Chao, who I am lucky to have her as a friend and coauthor. She has figured out how to make use of information in the sample to tell us how incomplete it is, even when we don’t know how many species there are in the actual population. I have written about this subject before, when she and I figured out how to compare the diversities of two or more ecosystems based on incomplete samples from each of them (with a version in Spanish here). This is a common task, but biologists had always been doing it wrong. That post was about estimating the number of species in each community. However, there are many reasons why that number is not the best measure of biological diversity or complexity. Not only is it the hardest number to estimate accurately (because there are always unseen species), it also ignores the relative abundances of species, and those abundances have an important effect on the ecosystems’s complexity. An ecosystem with one very abundant species of birds and nine very rare species is less complex than an otherwise-similar ecosystem with ten equally common species of birds. An individual bird in the first ecosystem can be quite sure that the next bird it meets will be a member of the very abundant species. A bird in the second ecosystem will be completely uncertain about which of the ten species it will encounter next.

The amount of uncertainty in the species identity of the next individual encountered is actually something that can be quantified exactly if we know the relative abundances of each species in the ecosystem. In 1948 Claude Shannon, the inventor of information theory, showed that this uncertainty is equal to the entropy function that had been derived by the physicists Boltzmann and Gibbs in the late 1800s. Shannon’s discovery led to the wide use of entropy in other disciplines, including ecology. Shannon’s entropy measure became the most commonly used abundance-sensitive measure of biodiversity. The entropy is a simple function of the relative abundances of each of the species present in the population. Ten years ago I showed that this entropy function needs to be transformed by taking its exponential before it can be interpreted as diversity (Jost 2006, 2007, 2010). But whether we are interested in entropy or its exponential, in biology we still have to estimate these quantities from incomplete samples, so we don’t know the true relative abundances of the species in the population, and we don’t even know exactly how many species there are in the population.

Anne had been thinking about this problem for more than thirty years, and recently came up with a beautifully elegant solution for estimating entropy from small samples. The solution makes clever use of the information contained in the sample about the unseen species in the population. I wrote briefly about it here. This month she was asked by the journal Methods in Ecology and Evolution to write a blog post about her solution. You can read her full blog post here, complete with essential links to more information. Here are some excerpts, with my notes in brackets:

Estimating Entropy from Sampling Data

In practice, the true number of species and their relative abundances are almost always unknown, so the true value of Shannon entropy must be estimated from sampling data. The estimation of this seemingly simple function is surprisingly difficult, especially when there are undetected species in the sample. It’s been proven that an unbiased estimator for Shannon entropy doesn’t exist for samples of fixed sizes.

The observed entropy of a sample or the ‘plug in’ estimator, which uses a sample fraction [the abundance of the species in the sample, divided by the size of the sample] in place of the [true] relative abundance of species [in the population], underestimates the entropy’s true value. The magnitude of this negative bias can be substantial.

For incomplete samples, the main source of the bias comes from the undetected species, which are ignored in the plug-in estimator. An enormous number of methods/approaches have been proposed in various disciplines to obtain a reliable entropy estimator with less bias than that of the plug-in estimator. The diversity of the approaches reflects the wide range of applications and the importance of bias-reduction.

My Introduction to Alan Turing’s Statistical Work

Around 1975 (when I was a graduate in the Department of Statistics, University of Wisconsin-Madison) my advisor at the time, Bernard Harris, suggested that an “attractive and absorbing” (his original description) PhD thesis topic would be to develop an ‘optimal’ entropy estimator based on sampling data. He thought Alan Turing’s statistical work might prove to be useful and hoped that I could tackle this estimation problem.

Alan Turing memorial statue in Sackville Park, Manchester, UK ©Lmno

Alan Turing memorial statue in Sackville Park, Manchester, UK ©Lmno

However, at that time I didn’t even know who Alan Turing was! Although I started to read two background papers by I. J. Good (links below) about Turing’s statistical work, I couldn’t fully digest the material in the short time available. So, I didn’t work on the entropy estimation problem for my PhD thesis; instead, I derived some lower bounds for a variety of diversity measures. Ever since then, however, entropy estimation has fascinated me and has been in my mind/thoughts, and I regarded it as my ‘unfinished thesis’ topic.

The Building Blocks of My Entropy Estimators

According to I. J. Good (Turing’s statistical assistant during World War II), Turing never published his wartime statistical work, but permitted Good to publish it after the war. The two influential papers by Good (1953) and Good and Toulmin (1956) presented Turing’s wartime statistical work related to his famous cryptanalysis to crack German ciphers. After graduation, I read these two papers many times and searched for more literature. It took me a long time to fully understand these two papers especially Turing’s statistical approach to estimating the true frequencies of rare code elements (including still-undetected code elements), based on frequencies in intercepted ‘samples’ of code.

“Mathematics, rightly viewed, possesses not only truth, but supreme beauty, a beauty cold and austere, like that of sculpture.” -Bertrand Russell ©Lmno

“Mathematics, rightly viewed, possesses not only truth, but supreme beauty, a beauty cold and austere, like that of sculpture.” -Bertrand Russell ©Lmno

The frequency formula is now referred to as the Good-Turing frequency formula. Turing and Good discovered a surprisingly simple and remarkably effective formula that is contrary to most people’s intuition. The formula proved to be very useful in my development of entropy estimators.

One important idea derived from the Good-Turing frequency formula is the concept of ‘sample coverage’. Sample coverage is an objective measure of the degree of completeness of the intercepted ‘samples’ of code elements. The ‘sample coverage’ of a sample quantifies the proportion of the total individuals in the assemblage that belong to sampled species. Therefore, the ‘coverage deficit’ (the complement to sample coverage) is the probability of discovering new species, i.e. the probability that a new, previously-unsampled species would be found if the sample were enlarged by one individual. Good and Turing showed that for a given sample, the sample coverage and its deficit can be accurately estimated from the sample data itself. Their estimator of coverage deficit is simply the proportion of singletons (in this case species with only one individual) in the observed sample. This concept and its estimator play essential roles in inferring entropy.

A Novel Entropy Estimator

A species accumulation curve (SAC) shows the cumulative number of species as a function of sample size. In the figure below we see the expected curve when individuals are sequentially selected from a community with a given number of species, with relative abundances.

The first breakthrough in my search for an estimator of Shannon entropy was the realization that entropy can be expressed as a simple function of the successive slopes of the SAC.
[Anne is probably the only person in the world who would have noticed this!] The curve’s successive slopes show the rates at which new species are detected in the sampling process. I had found a novel way to estimate entropy via discovery rates of new species in a SAC and these rates or slopes are exactly Turing’s coverage deficits for varying sample sizes!

The statistical problem was then to estimate the expected slopes or coverage deficits for any hypothetical sample size. Good and Turing’s approach provided the coverage deficit estimator for the expected slope of the sample that has been taken. All of the expected slopes for smaller sample sizes can be estimated without bias from statistical inference theory. However, there is no unbiased estimator for the expected slopes for sample sizes greater than the sample taken. These slopes are usually dominated by rare undetected species whose effect on entropy cannot be ignored. So, the burden of entropy estimation is shifted onto the estimation of the expected slopes for sizes greater than our sample.

The second break-through step to solve this problem was also attributed to the wisdom of Turing and Good, who showed that the number of singletons carry much information about the number of undetected rare species. I slightly modified their idea to use both singletons and doubletons to estimate the hard-to-estimate slopes by my modified Good-Turing frequency formula.

With the collaboration of Lou Jost and the simulation/programming work of Y.T. Wang, we published in 2013 the novel entropy estimator based on the derived slope estimators [open access to full text]. Our extensive simulations from theoretical models and real surveys generally showed that the new estimator outperformed all the existing estimators. It took me over 35 years to derive the optimal estimator for my ‘unfinished thesis’, so I have been calling it my entropy ‘pearl’. (The novel entropy estimator along with other related estimators can be calculated online.)

Doing Research is like Carving Jade

…As the old saying goes: “doing research is like carving jade, we are never satisfied with what we have until it is perfect”. This is also my advice to anyone starting their career in academia. The topic of entropy estimation has attracted and absorbed me for more than 35 years, and hopefully the novel estimator did yield an ‘optimal’ solution, if it’s still not perfect.

In her blog post Anne also explains how this method generalizes to the estimation of a wider class of diversity measures based on generalized entropy, a problem she and I had been working on for ten years. See her original blog post for details.

Anne’s contributions to the mathematics of biology are one of the reasons why Sebastian Vieira and I recently named a new orchid after her. Thanks, Anne, for your very fruitful collaborations over the years!

Lou Jost


Chao, A. and Jost, L. (2011). Diversity measures. Sourcebook in Theoretical Ecology (eds. A. Hastings and L. Gross). Berkeley: University of California Press.

Chao, A. and Jost, L. (2012) Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93: 2533-2547.

Chao, A. and Jost, L. (2015) Estimating diversity and entropy profiles via discovery rates of new species. Methods in Ecology and Evolution 6: 873–882.

Chao A, Wang YT, Jost L (2013) Entropy and the species accumulation curve: a novel entropy estimator via discovery rates of new species. Methods in Ecology and Evolution 4: 1091-1100.

Jost, L. (2006) Entropy and diversity. Oikos 113: 363–375.

Jost, L. (2007) Partitioning diversity into independent alpha and beta components. Ecology 88: 2427–2439.

Jost, L. (2009) Mismeasuring biological diversity: Response to Hoffmann and Hoffmann (2008). Ecological Economics 68: 925–928.

Two more new frogs discovered in our Rio Zunac Reserve

Pristimantis sacharuna. Photo: Mario Yanez.

Pristimantis sacharuna. Photo: Mario Yanez.

Our Rio Zunac Reserve has been an endless source of new discoveries of plants and animals. I’ve written in the past about the discovery of new magnolia species, new melastomes and orchids, and new frogs. Last month herpetologists Juan Pablo Reyes (who is also our reserve manager), Carolina Reyes, Maria Perez L., and Mario Yanez, (who is also an EcoMinga director and head of the National Institute for Biodiversity), have published two more new frogs from this reserve. One of the new species, Pristimantis pinchaque, was discovered at 1600m elevation in the immediate vicinity of the scientific station that we built there some years ago with the help of the IUCN-Netherlands and the Netherlands Postcode Lottery, while the other new species, Pristimantis sacharuna, was discovered farther up the trail from the station, at 2200m. That makes four new species discovered so far in this reserve, joining Pristimantis ardyae and Osornophryne simpsonii.

Pristimantis pinchaque. Photo: Mario Yanez.

Pristimantis pinchaque. Photo: Mario Yanez.

Pristimantis pinchaque is apparently very rare; it has not been seen since the first two specimens were found in 2008, even though many herpetologists have visited the site since then. It is named after the Mountain Tapir, Tapirus pinchaque, a charismatic and endangered mammal which lives in the same forest. Pristimantis sacharuna is also apparently very rare, with only two specimens found in four years of investigation. It is named after the “duendi” or mythic forest man of indigenous legends.

The story of these discoveries was covered nicely by the national press. The country’s largest newspaper, El Comercio, even made an interactive article that lets readers see the diagnostic traits of each frog by clicking on different parts of its anatomy, and in a special feature for their “Planet” section, they also published a nice diagram to help readers distinguish the frogs:

And there is still more to come! We have two more new frog species being described right now, from survey work supported by a donation from Henri Botter and Ardy Van Ooij of the Netherlands.

These investigations are collaborative efforts between EcoMinga and the National Institute of Biodiversity, the Zoological Museum of the Pontificia Universidad Católica del Ecuador, and the Fundacion Oscar Efren Reyes. We’re excited to have such distinguished collaborators, and we are eager to see what surprises still await us in these very special forests.

Lou Jost