The Race: Odds and Year’s Ends

A female leatherback sea turtle (Dermochelys coriacea) on Praia Grande, Príncipe island heading back to sea after laying her eggs overnight. Phot. V. Schmitt.

At up to 3 m (9.8 ft) and 932 kg (2,055 lb), the leatherback is the worlds 4th largest reptile (after three crocodile species) and by far the largest turtle.

Along with two other marine species (above), the green (Chelonia mydas – inaturalist phot), and the hawksbill (Eretmochelys imbricata – Brittanica phot.), it breeds annually on certain São Tomé and Príncipe beaches.

Mouth and pharynx of Leatherback. Pinterest phot.

Leatherbacks feed almost exclusively on coelenterates (jellies or jellyfish) and swallowing them is added by long inward-pointing fleshy projections in the throat and pharynx (above). All of the worlds marine turtles are endangered species and the Gulf of Guinea Ids are important sites, guarded and studied by a number of NGO’s including the Príncipe Trust and ATM: the Association for the Protection, Research and Conservation of Sea Turtles in Lusophone countries.

The birds (avifauna) of the two islands are quite remarkable. Combined, São Tomé and Príncipe are only about 1000 km² and yet they are home to 28 unique endemic bird species of many different families. This is the highest density of bird endemism by area in the world. We have also discussed gigantism and dwarfism (above), poorly understood phenomena occurring frequently in island plants and animals.

São Tomé Grosbeak, Neospiza concolor, M. Melo phot and at right.

There have been several recent studies on the islands’ endemic bird species. Dr. Martim Melo of CIBIO, foremost latter-day student of this avifauna has shown that the genetic relationships of the seemingly extremely scarce São Tomé Grosbeak (above) reveal that it is actually a canary, not a grosbeak. It is thus now the world’s largest canary and a true island giant. It is 50% heavier than the next largest related species. As an island giant, it joins other São Tomé giant endemics like the Giant Sunbird, Dreptes thomensis, and the Giant Weave,r Ploceus grandis, the two largest species of their families in the world. These have been featured in earlier blogs. Two additional, less famous São Tomé species are the largest species of their respective families in Africa (above).

São Tomé Speirops, Xosterops lugubris. Phot.  Lars Petersson]

SãoTomé Thrush,  Turdus olivaceofuscus, Phot.  Lars Petersson]

Under different systems, all of the endemic bird species could be considered rare and endangered since each unique species can only have a range of significantly less than 1000 km², which is the area of the two islands combined. In a sense, these unique species could be considered among the rarest in the world. One of the fathers of the “modern synthesis” (evolution + genetics), Prof. Ernst Mayr, wrote a paper in the magazine Science examining the levels of bird endemicity in relationship to island size and distance to mainland source. The extraordinary density of the endemic birds of our islands when compared with predicted levels (below) yield a separate third curve; a phenomenon he simply illustrated but with no attempt to explain it.

The status of nine of the São Tomé endemics is “threatened” according to modern ornithologists; until recently, three of these have been considered “critically endangered”: the São Tomé Grosbeak (above), the Dwarf Olive Ibis and the São Tomé Fiscal Shrike (below)

São Tomé Fiscal Shrike, Lanius newtoni. Phot. L. Petersson

Dwarf Olive Ibis, Bostrychia bocagei. Phot. L. Petersson

A recent paper by Dr. Ricardo de Lima (below) and colleagues is an examination of the ranges and population status of these three endemic birds. They found that the south-west central region of the island, most of which is included in the São Tomé Obô Natural Park, has the highest potential for the Critically Endangered birds, and that indeed all three species were associated with native forest. The ibis prefers high tree density, while the fiscal selects low tree density and intermediate altitudes. “Despite very restricted ranges, population sizes seem to be larger than previously assumed. These results suggest that the fiscal and grosbeak might be better classified as ‘Endangered’, while the ibis should maintain its status under different criteria, due to a very restricted range during the breeding season.”

Drs. Rayna Bell, Smithsonian Institution (left) with Dr. Ricardo Lima of University of Lisbon (right)

The colorful and varied endemic bird fauna of São Tomé and Príncipe is a flamboyant if somewhat enigmatic example of speciation and evolution over time. As the graph above illustrates, there are more endemic species by unit area than one would predict, even in light of the near proximity of the African mainland. Although the unique species are now reasonable well-described compared with other elements of the fauna, there is still a great deal to be learned.

The only venomous snake on the islands is the Forest Cobra “Cobra preta”, of São Tomé, fairly common in most inland habitats, and the source of some controversy over the years (see blog, May 2009). Whether it is naturally occurring i.e., the descendent of a disperser like many of the terrestrial uniques, or whether it was brought by man in some manner has been a point of discussion for some years. For a long time the few scientists working on the islands have referred to this species as Naja melanoleuca, a large cobra which is widespread on the African mainland (below).

From blog, May 2008: Within the House of Slytherin II. (left) Naja melanoleuca. Cowyeow photo]

A recent publication by Dr. Luis Ceríaco et al. (National Museum of Natural History and Science) suggests that the São Tomé cobra is a naturally occurring endemic; they have named it Naja peroescobari, after one of the Portuguese navigators who discovered the island in 1471. The authors distinguish Cobra preta from the mainland species by slightly larger size, a different color pattern underneath, and the arrangement of some scales in the throat area.

The São Tomé cobra, Naja escobari. Phot. Tiziano Pisoni, and see Dec. 2013 blog- Another New Species…

While on the subject of reptiles, recent expeditions have afforded us with more knowledge, along with some outstanding new photos of related island endemics:

São Tomé “jita”. Boaedon (Lamprophis sp). Phot Lars Petersson

Príncipe “jita”. Boaedon (Lamprophis sp). Phot. Andrew Stanbridge

We know these two island species are distinct from one another and from close mainland relatives on the basis of pattern and molecular evidence. The big island species (above) has a pattern of lines along the back; that of Príncipe is a series of blotches. A group of us led by Dr. Luis Ceríaco is in the process of assigning new names to these island two populations.

Greef’s Giant gecko, Hemidactylus greefii of São Tomé. A. Stanbridge phot. Note greenish eyes.

Príncipe Giant gecko, H. principensis. R. Bell phot. Note golden eyes.]

Hugulay Maia (insert, below) is a young São Toméan from the big island who has been interested in the natural history of the Gulf of Guinea Islands for most of his life. He and his mentor, the late Angus Gascoigne, have appeared in this blog a number of times over the years, and Hugulay is currently a PhD candidate in marine ecology at the University of Santa Catarina in Brazil. He has just produced a durable guide to the marine species most frequently encountered by local fishermen, pictured below. As a scientist he will be a strong, valuable asset to the islands’ fauna and flora and to the government, especially with respect to the conservation of its natural marine resources.

Quintino Quade (below left) and Genevieve Chase on Macambrara Ridge, São Tomé 2017.

Quade of STeP UP has been a bulwark of our field work and education program since the beginning in 2001. He is a translator, teacher of English and many other useful things.  Chase is a consultant from Washington D.C., examining possibilities for expansion and modification of our Gulf of Guinea Project in the future.

The Parting Shot:

Me at Jockey’s Bonnet, Principe Id. This is an ancient island off the east coast that harbors a currently recognized endemic subspecies of finch.

PARTNERS
Our research and educational expeditions are supported by tax-deductable donations to the “California Academy of Sciences Gulf of Guinea Fund*.” We are grateful for ongoing governmental support from the Republic of São Tomé and Príncipe, and especially to Arlindo de Ceita Carvalho, Director General, Victor Bonfim, and Salvador Sousa Pontes of the Ministry of Environment and to Faustino de Oliviera of the Department of Forestry for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos, Anita Rodriguez and Quintino Quade of STePUP of Sao Tome, our “home away from home”. GG IX, X and XI were funded in part by a generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Rod C. M. Hall, Mr. and Mrs. Henri Lese, Mr. and Mrs. John L. Sullivan Jr., Mr. and Mrs. John Sears, in memory of Paul Davies Jr. and a heartening number of Bohemian friends. We are grateful for the support of Roça Belo Monte (Africa’s Eden-Príncipe) for both logistics and lodging.
*55 Music Concourse Drive, Golden Gate Park, San Francisco CA 94118 USA

 

 

 

 

 

The Race: “Once More Unto the Breach, Dear Friends…” – (the troops)

Ghosts of Past Bondage and Present Beauty (unless otherwise indicated, images by our photographer, A. Stanbridge.)

Our next expedition, GG XI, departs in November. Since returning from GG IX last year, we have been involved in two subsequent expeditions: that of graduate student Matthias Neumann (University of Kassel) whose work on island flatworms we are supporting, and GG X, our second marine expedition led by Dr. Luiz Rocha, of the Academy. As a result, GG IX has perhaps received less “blog attention” than usual, so I am including a few more images below.

Upon arrival on Príncipe Island, we always pay our respects to President Jose “Tose” Cassandra, in order to inform him of our intentions. He has been a strong supporter of our scientific and educational work on the island since early days; as can be readily seen, a visit to his office is always a pleasure. To my left is Dr. Maria Jeronimo, Portuguese entomologist.

Above, the team is hiking up to the rim of Lagoa Amelia, a crater lake on São Tomé at over 1400 meters. The giant bamboo is  an invasive or it was introduced for some reason; it is not native to the islands. One of the major joys of being a field biologist is that one often finds oneself working in wonderfully beautiful places like these.

Kids watching a video of themselves dancing in an abandoned roça (plantation house).

At about 700 m in elevation in the Contador Valley on the northwest side of São Tomé, there is a kilometer-long tunnel/aqueduct that is a great locality for bats, amblypygids,  geckos(below), and other normally night-time critters of interest. The team in route to the tunnel: Dr Luis Mendes, Dr Rayna Bell, Lauren Scheinberg, Drewes, Dr Maria Jeronimo and K. B. Lim.

Lauren Scheinberg and Rayna Bell in the tunnels.

 

Drewes, Quintino Quade and K. B. Lim in the tunnels.

The giant, four-fingered gecko of Sao Tome, Hemidactylus greefi.

The tunnel entrance is behind us; we are examining specimens just collected . To the left is Scheinberg, Drewes, K.B.Lim, manager of the local power plant, Dr. Luis Mendes (foreground) and Quintino Quade.

The water of the Rio Contador is eventually directed to the country’s only brewery,  far below in the town of Neves. Here ROSEMA, the local beer, is produced; we feel this is a noble enterprise and support it frequently and enthusiastically.

Dr. Luis Mendes with Quintino Quade on the hunt.

The prey: a silverfish, one of Luis’s academic specialties.

Below, Dr. Rayna Bell of the Smithsonian Institution with a São Tomé giant treefrog, Hyperolius thomensis. Rayna has been studying the genetics and evolution of the unique tree frogs of the islands for a number of years.

Above, three members of the team work an unnamed waterfall on the west side of Príncipe Island; from left to right: Dr. Maria Jeronimo, Rayna and Lauren Scheinberg. Shortly after this image was taken, Rayna Bell collected a large female Príncipe giant tree frog (Leptopelis palmatus).

Not only is it rare to find such an animal during daylight hours, this is also the only all-black African tree frog specimen I have ever seen in a lifetime of studying them. Tree frogs always protect their under surfaces against water loss by evaporation; thus the frog is perched (above) on the largest smooth surface available nearby- our indomitable photographer, Andrew Stanbridge.

Andrew is the veteran of six expeditions and has provided us with invaluable photographic documentation of our past six years of fieldwork.

Drs Jeronimo and Luis Mendes, our GG IX entomologists, examine the latest butterfly capture (above) and below, Roberta Ayres, coordinator of all of our education efforts examines the oddest vertebrate on São Tomé, the unique legless amphibian or caecilian (Schistometopum thomense) known to the locals as “Cobra bobo” and greatly feared as well. Its nearest relative is found thousands of kilometers to the East in Tanzania and Kenya. Caecilians are harmless.

Roberta Ayres  near São Nicolao, São Tomé.

The eleventh expedition will run from mid-November to mid-December and will include nine scientists, two of them new to the project.
Our botanists, Drs Tom Daniel and Jim Shevock, will be joined for the first time by Dr. César Garcia (below left) of the University of Lisbon, Museum of Natural History.  César is  a bryophyte specialist who has already worked and published with shevock; together they will continue to survey the moss, liverwort and hornwort flora of the islands. This year the botanists will again attempt a survey of the remote Pico Mesa on Principe.

Dr. Lauren Esposito (above, right, with an American crocodile in hand – Crocodylus acutus) is a new member of the faculty of CAS and a specialist on arachnoids. She and returning entomologist Maria Jeronimo will continue with our ongoing broad survey of the insect faunas of the islands. Roberta Ayres, our education head, Rayna Bell, Andrew and I round out the members of GG XI. While on Príncipe we will hopefully be joined in the field by Felipe Spina a bee biologist with the Príncipe Trust.
Each year, we look forward to seeing our local collaborators and friends such as Quintino Quade, his wife Anita Rodrigues and Roberta dos Santos all from the NGO STeP UP, Arlindo Carvalho of the Ministry of the Environment, our “friend on the mountain,” Henrique Pinto da Costa and many, many others.

We are very fortunate to have the support of both the Omali Boutique Hotel on São Tomé and Roca Belo Monte on Príncipe (above left and right respectively) who provide us with logistical support during our expeditions; this has been vital to the success of our Gulf of Guinea projects over the years.
Here’s the parting shot:

Storm brewing on the southeast coast near Rebeira  Peixe, Sao Tome.

PARTNERS:
Our research and educational expeditions are supported by tax-deductable donations to the “California Academy of Sciences Gulf of Guinea Fund.”*  On the islands, we are grateful for ongoing governmental support, and especially to Arlindo de Ceita Carvalho, Director General, Victor Bonfim, and Salvador Sousa Pontes of the Ministry of Environment of the Republic of São Tomé and Príncipe for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos and Quintino Quade of STePUP of Sao Tome, our “home away from home”. The upcoming GG XI has been funded in part by a generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Rod C. M. Hall, Timothy M. Muller, Mr. and Mrs. John L. Sullivan Jr., Mr. and Mrs. John Sears, and a heartening number of “Coolies”.  Once again we are deeply grateful for the support of the Omali Lodge (São Tomé) and Roça Belo Monte (Príncipe) for both logistics and lodging, and to the Príncipe Trust for partial sponsorship of our ongoing primary school education program.

*California Academy of Sciences
55 Music Concourse Dr.
San Francisco, CA 94118
USA

 

The Race: Updates and Progress in Paradise

The new header image of Príncipe Island from the east (above) was made by our friend, Jan Fourie, of Africa’s Eden; Príncipe is 31 million years old and was much, much larger in the Oligocene.

 

Revisiting the famous “Bode of Bombaim” with cobra skin in hand, along with our indefatigable photographer, Andrew Stanbridge (left). This area of central São Tomé seems to harbor sizable numbers of forest cobras (Naja nigricollis), the islands’ only venomous snake species, thought to have been introduced by early Portuguese settlers. We have extracted DNA from Bode’s skins to test this hypothesis.

3/4ths of the members of GG IX. Dr. Rayna Bell, UC Berkeley; Lauren Scheinberg, CAS; Maria Jeronimo, Gulbenkian U; and Dr. Luis Mendes, Nat. Hist. Mus., Lisbon. Absent are Roberta Ayres (CAS), Andrew Stanbridge, photographer and me (CAS).

Some more updates from GG IX: Dr. Luis Mendes is completing his monograph on the butterflies of the islands. He informs us that he collected about 400 specimens during GG IX. 

His collections represent 40 species of six families from both islands with new records and observations of endemics.

Jim Shevock (above) of CAS, is a veteran of three past Gulf of Guinea expeditions, and has just published a sixth scientific paper on bryophyte flora of the islands. It is plain that the bryophyte flora of the islands is much more diverse than had been thought and Jim has many more species to be and new ones to describe especially. Jim will be a participant on GG XI in November.

Dr. Rayna Bell (above left), now of the Smithsonian Institution, continues her work with the treefrog genus Hyperolius. The opportunity arose for us to sample the southern part of the Obo Natural Forest on São Tomé where, it turns out, the giant tree frog (H. thomensis) is much more easily found and observed. There are some intriguing biological issues involving genetic interaction between these two species which are so different in size and color (above right), and Rayna continues her studies of them and the giant tree frog (Leptopelis) of Príncipe.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Drs Bell and Ricardo Lima in the forest of São Tomé, inland of Angolares.

On the academic side of things, the Island Biology Conference held in July at the University of the Azores on Terciero Island was a great success with over 400 scientists and students in attendance for a week. At the first such meeting held in Hawaii there were only two presentations on Gulf of Guinea science; in the Azores, we had a day-long symposium featuring talks on many aspects of island biogeography and conservation.

Dr. Bell presents an amphibian paper at the Gulf of Guinea Symposium, Terceiro Island, Azores.

Such meetings facilitate useful interactions between scientists and students, allowing them to avoid overlap of effort and at the same time promoting cooperation; we were very heartened by the increase in the number of people doing research and educational activities on the islands.

Drs Mariana Carvalho and Ricardo Lima discuss various São Tomé/Principe
projects (above). Both are authorities on forest ecology,  the Gulf of Guinea bird fauna and the interactions of human populations with the environment. After several years of work in Mozambique, Mariana is returning to the islands where she will continue her work under the auspices of Birdlife International. Ricardo was one of the organizers of the symposium.

In several previous blogs I have mentioned Hugulay Maia, a Sao Tomean from the town of Angolares on the southeast coast. We first met Hugulay years ago through his mentor and friend, Angus Gascoigne, an accomplished resident naturalist on São Tomé. Tragically, Angus passed away a few years ago; he would have been very proud to learn that Hugulay  is now pursuing PhD research on the coastal fishes of the islands.

Hugulay diving on Príncipe, GG.X (left) and preparing specimens (right, with Dr.Ricardo Rocha (CAS), and graduate student,  Luisa Fontoura. (far right, U. Catarina, Brazil).

Maia was a member of the GG X marine team, as was his doctoral advisor, Dr. Sergio Floeter of University of Santa Catarina, Brazil. A few months ago in Lisbon (below), he presented part of his thesis work to the Gulbenkian Foundation in Lisbon, one of the major funders of his work on coastal fishes.

Our next terrestrial expedition, GG XI will be in November and will be the topic of the next blog.

PARTING SHOT.

Here at Praia Jalé in southeastern São Tomé is a leatherback sea turtle (Dermochelys coriacea), one of several species of ocean-going sea turtles that breed on the islands annually. Looking on are our old friends Bastien Loloum, his wife Delicia and kids Flora and David. Bas said: “The nesting turtle got surprised by sunlight and was just finishing up [laying eggs] as we arrived by her side. The picture was taken by a German tourist who was also staying at the lodge that same night.” This is the world’s largest turtle and the 4th heaviest reptile (after 3 monitor lizards). These giants can reach 2.13m (just under 7 feet) with a mass of 650 kg (1433 lbs)!

 

PARTNERS.
The research expeditions are supported by tax-deductable donations to the “California Academy of Sciences Gulf of Guinea Fund”* On the islands, we are grateful for ongoing governmental support, especially to Arlindo de Ceita Carvalho, Director General, Victor Bonfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos and Quintino Quade of STePUP of Sao Tome http://www.stepup.st/, our “home away from home”. GG VIII, IX , X and upcoming GG XI have been funded by a generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Mrs. W.H.V.“D.A.” Brooke, Thomas B. Livermore, Rod C. M. Hall, Timothy M. Muller, Prof. and Mrs. Evan C. Evans, Mr. and Mrs. John L. Sullivan Jr., Clarence G. Donahue, Mr. and Mrs. John Sears, and a heartening number of “Coolies”, and members of the Docent Council of the California Academy of Sciences. Once again we are deeply grateful for the support of the Omali Lodge (São Tomé) and Roça Belo Monte (Príncipe) for both logistics and lodging and to the Príncipe Trust for partial sponsorship of our on-going primary school education program during GG VII and GG VIII.

*California Academy of Sciences
55 Music Concourse Dr.
San Francisco, CA 94118
USA

 

THE RACE: As the (Flat)worm Turns

Dr. Tom Daniel, senior botanist, demonstrating impressive intrepidity in a Sao Tome river. GG VII A.  Stanbridge phot.

In the earliest blogs we discussed how islands are ideal for studying certain evolutionary processes and patterns; some of these phenomena like gigantism and dwarfism (below) are actually characteristic of islands. And, the results of these processes are much easier to see on islands because of their smaller size (versus, say, continents) and the smaller size of the plant and animal populations that inhabit them.

Worlds largest (left, Sao Tome) and smallest (right, Principe) Begonia. (RCD construct).

Invasive species are organisms that somehow become accidentally established in areas where they did not exist before; they can be hugely damaging to ecosystems especially on islands, that are made up of plant and animal species that have co-evolved in isolation over perhaps millions of years. In the absence of natural predators (checks and balances), invader populations can become numerous and spread rapidly, and this can have a devastating effect by exhausting the resources these species utilize in the local environment. Invasive animal species populations can burgeon hugely, and then frequently die off; by then, the damage is usually done.

(Phot. Miko Nadel, GG VII)

A few years ago, we received a photo of a brightly colored worm-like creature on São Tomé (above), taken by one of our graduate students, Miko Nadel, a lichenologist from San Francisco State University. This striped creature turned out to be a terrestrial flatworm, a member of a primitive phylum of invertebrates called the platyhelminthes. Those of us lucky enough to study biology backwhen students were given actual organisms to observe rather than plastic models or video clips will remember “planarians,” aquatic flatworms (below) noted for their amazing ability to regenerate.

Planarian (Dugesia) . Stock photo, Google photos.

Flatworm regeneration. (Univ. Heidelberg photo.

Terrestrial flatworms, also known as geoplanids have no anatomical or physiological mechanisms for retaining water and are thus very much tied to moist environments. They are voracious predators upon other soil invertebrates such as earthworms, slugs and, most importantly for us, snails. Geoplanids secrete a mucus that begins to digest and dissolve their prey externally (below), and all geoplanid species known feed using an extendable digestive tube or pharynx.

Geoplanid attacking a snail. (R. Lima, phots)

As above, at Macambrara, Sao Tome. [Phot S. Mikulane]

Once aware of these, we began to notice more of them as did our ecologist colleague, Dr. Ricardo Lima, and we all became rather concerned. Why? Readers may recall that about half to 60% of all of the species of terrestrial mollusks (snails) of both São Tomé and Príncipe are endemic; i.e., they are found nowhere else in the world.

[RCD construct-multiple photographers]

In fact, these snails have been isolated on the island and evolving for such a long period that scientists currently recognize six different genera and a unique snail family there! Given what we know about invasive species, if these geoplanids are indeed a new arrival then the unique snail fauna of the islands may well be in significant danger.

 

Another geoplanid [phot. F. Azevedo]

In 2012, we sent the original specimen to Dr. Ronald Sluys of Naturalis Biodiversity Centre in Leiden, Holland, one of the few specialists on flatworms. In the meantime, Dr. Ricardo Lima (below) has been conducting ecological research on the São Tomé forests for several years and has been able to collect many more geoplanids and send them to Dr. Sluys.

When the GG IX team visited Dr. Lima at Monte Café, São Tomé last year, he showed us pictures of a number of very different looking morphs of flatworms that he had collected and sent to Dr. Sluys. Were these different species or just variations on one or two species (morphs)?

Various geoplanids from Sao Tome [phots R. Lima, F. Azevedo, M. Nadel, R. Rocha]

Ron Sluys has been supervising a graduate student from the University of Kassel, Germany who is doing his MSc degree based on this material. His name is Matthias Neumann and as I write, he is on the island of São Tomé with Dr. Lima, studying the flat worms in situ and collecting more! We were able to fund his expedition with the Academy’s Gulf of Guinea Fund (see “Partners”, below).

Matthias Neumann and partner in the bush on Sao Tome. [R. Lima phot]

Matthias’ preliminary work suggests that there are indeed a number of species of geoplanids on the island: at least five new, undescribed species of the genus Othelosoma, and another previously known species, Bipalium kewense. B. kewense is an extremely widespread species, presumably carried about in the roots of plants; evidently it preys on earthworms rather than snails.

So far, little is known of these strange creatures. Neumann says that at least two of the undescribed species of Othelosoma are snail predators but even so, the presence of a number of species on the island rather than one dominant, rapidly spreading one might be taken as somewhat reassuring. If there are a number of closely related members of the same flatworm genus, it is more likely that the common ancestor of these species arrived a long time ago, speciated, and thus co-evolved with the endemic snail fauna. If this is so, than we would expect an ecological predator/prey balance in this system. If the flatworm fauna is in fact a radiation from a given colonizer, then it would mirror the status of the earthworm fauna as we understand it (below). So far, we are uncertain whether geoplanids are present on Príncipe.

 

 

 

 

 

 

 

 

 

 

We are in the planning stages for GG XI; see the next blog.

The Parting Shot:

 

 

 

 

 

 

 

A local teacher assists us in a binoculars demonstration on Sao Tome, GG IX. Dr. Luis Mendes in background.  [phot. A. Stanbridge]

Partners:

The research expeditions and the primary school education program are supported by tax-deductable donations to the “California Academy of Sciences Gulf of Guinea Fund”* We are grateful for ongoing governmental support, especially to Arlindo de Ceita Carvalho, Director General, Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos and Quintino Quade of STePUP of Sao Tome http://www.stepup.st/, our “home away from home”. GG VIII, IX , X and upcoming GG XI have been funded by a generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Mrs. W.H.V.“D.A.” Brooke, Thomas B. Livermore, Rod C. M. Hall, Timothy M. Muller, Prof. and Mrs. Evan C. Evans, Mr. and Mrs. John L. Sullivan Jr., Clarence G. Donahue, Mr. and Mrs. John Sears, and a heartening number of “Coolies”, and members of the Docent Council of the California Academy of Sciences. Once again we are deeply grateful for the support of the Omali Lodge (São Tomé) and Roça Belo Monte (Príncipe) for both logistics and lodging and for partially sponsoring our education efforts for GG VII and GG VIII.

*California Academy of Sciences
55 Music Concourse Dr.
San Francisco, CA 94118
USA

 

 

THE RACE: Homage to “The Prince”

The island of Príncipe is ancient… at 31 million years of age it is twice as old as São Tomé, yet biologically the two islands are unquestionably related. Along with documenting and describing hitherto unknown species of strange, endemic plants and animals that inhabit one or the other island (rarely, both), we attempt to understand the relationships of these species to each other and to their ancestral populations from the African mainland.

For instance among the striking Príncipe uniques is Leptopelis palmatus, the Príncipe giant tree frog (above). Females of this species attain dimensions such that they are largest tree frog in Africa! Males, first described by us, are usually less than half their length. The original specimen upon which the species description was based over a century ago was a single female of 110mm body length (excluding legs). Like all female frogs, they do not have an advertisement call and despite their great size, very few have been found and reported in the scientific literature.

(male)

(female)

A few days ago, Dr. Rayna Bell found the large female pictured above, pressed to the surface of a small flat rock on the ground on a steep, dryish slope in the northwestern part of the island. Females tend to be dark compared to males, but this is the first all-black specimen reported. There are three other tree frog species on the islands, but they are all closely related to each other and belong to a different frog family from the Príncipe giant.

Dr Bell has shown that these other three are most closely related to a species from the Ogooué and Congo River basins and thus likely of western Central African origin. Other work has shown that the nearest relative of Leptopelis palmatus of Príncipe is from west of the Niger River and thus the giant is  probably of northern (West African) origin, perhaps dispersing from the Niger River drainage..

Dr. Luis Mendes and Maria Jeronimo have continued to collect butterflies to fill in knowledge gaps with species for Luis’s book. Luis has collected a number of specimens that he cannot readily identify; on these poorly-known islands, this is particularly exciting.

Maria has actually been doing a lot of everything: collecting butterflies with Luis, joining us in the classrooms and going out at night collecting with Dr. Rayna Bell, Lauren Scheinberg and our photographer, Andrew Stanbridge. Considering that this is a “break” from her PhD dissertation work, her energy level is truly impressive.

Most geckos are nocturnal but the genus Lygodactylus is comprised of strictly daytime species in Africa and Madagascar. Readers may recall that the islands of Annobon, São Tomé and Príncipe are each inhabited by a single endemic species usually distinguishable by different black markings under the chin; other than these markings, the island species are usually a combination of grey and black. Luis Mendes captured an adult male on Príncipe that appears to be in breeding coloration of a sort I have not seen before in this genus. There is one species endemic to a small forest in Tanzania that is a beautiful blue, several others in East Africa that have yellow heads, and one species in Zambia that has a yellow belly. The male collected on Principe has a bright yellow head and the body that is a striking shade of light green. Not only is this the first time I have observed a green individual, I am also unaware of any literature describing temporary (usually hormonal) color intensities associated with breeding activities in this group of lizards.

Many of the photos we post on social media might well suggest that our work is being carried out in some sort of paradise; in some ways it is exactly that but is by no means easy!

 

The work we do here is only possible with the support of local entities; this is especially true on Príncipe. On our first two expeditions years ago we had difficulty finding suitable accommodations (reliable power for our equipment, etc) and logistics; there were not many available vehicles on the island, and we had little access to the really interesting higher elevation areas of the island or more remote southern areas. Since that time, our efficiency has increased hugely due to the generous support of several organizations on the old island. First and foremost is the Office of the Regional President (Tose Cassandra-he is also head of the recently created Principe World Biosphere Reserve, and also Daniel Ramos, head of the Príncipe Obo Natural Park.

For a number of years, we have been able to stay at Bom Bom Island resort which has also helped with needed transportation, both vehicular and marine. This year we were invited to stay at the new Roça Belo Monte (Africa’s Eden) who also provided transportation and assistance. At one point we planned a boat trip to explore the remote southeastern part of the island but the skipper, Bobby Bronkhorst, of Makaira Lodge fell ill.

Our work is now also in cooperation with the newly formed Príncipe Trust; the Trust played a major supporting role in the production of this year’s biodiversity bird field guide/coloring book and binoculars! Our biodiversity education efforts were concluded for Gulf of Guinea IX here on the old island with our return to the 3rd grade classes of the same schools we have been visiting since 2011. Usually after class visits, we see 3rd graders out in the bush peering through their new binoculars but frequently backwards! This may well be more fun for them.

Parting Shot:

 

PARTNERS:
We are most grateful to Arlindo de Ceita Carvalho, Director General, Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos and Quintino Quade of STePUP of Sao Tomehttp://www.stepup.st/, our “home away from home”.We gratefully acknowledge the support of the G. Lindsay Field Research Fund, Hagey Research Venture Fund of the California Academy of Sciences for largely funding our initial two expeditions (GG I, II). The Société de Conservation et Développement (SCD) and Africa’s Eden provided logistics, ground transportation and lodging (GG III-V), and special thanks for the generosity of private individuals who made the GG III-VII expeditions possible: George G. Breed, Gerry F. Ohrstrom, Timothy M. Muller, Mrs. W. H. V. Brooke, Mr. and Mrs. Michael Murakami, Hon. Richard C. Livermore, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. Robert M. Taylor, Velma and Michael Schnoll, and Sheila Farr Nielsen; GG VI supporters include Bom Bom Island and the Omali Lodge for logistics and lodging, The Herbst Foundation, The “Blackhawk Gang,” the Docent Council of the California Academy of Sciences in honor of Kathleen Lilienthal, Bernard S. Schulte, Corinne W. Abel, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. John Sears, John S. Livermore and Elton Welke. GG VIII was funded by a very generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Mrs. W.H.V.“D.A.” Brooke, Thomas B. Livermore, Rod C. M. Hall, Timothy M. Muller, Prof. and Mrs. Evan C. Evans, Mr. and Mrs. John L. Sullivan Jr., Clarence G. Donahue, Mr. and Mrs. John Sears, and a heartening number of “Coolies”, “Blackhawk Gang” returnees and members of the Academy Docent Council. Once again we are deeply grateful for the support of the Omali Lodge (São Tomé) and Roça Belo Monte (Príncipe) for both logistics and lodging and for partially sponsoring part our education efforts for GG VII and GG VIII.

Our expeditions can be supported by tax-deductable donations to “California Academy of Sciences Gulf of Guinea Fund”

THE RACE: GG IX – RETURN TO THE BIG ISLAND

The seven members of GG IX all met up in the Lisbon airport on September 18 and arrived the next day in São Tomé. Two new collegues on this expedition are from Portugal. Dr. Luis Mendes, a butterfly expert from the Natural History Museum in Lisbon is finishing a major book on the butterflies of the islands and is checking certain localities for species that have not been seen for many years.

Maria Adelina Jeronimo, a PhD candidate from the Gulbenkian Institute in Portugal, also studies butterflies but specifically the genetics of certain novel morphological characters. Maria is a matter of months from finishing her doctoral dissertation.

Dr. Rayna Bell has returned for her third trip and is continuing her studies of the interesting hybridization phenomenon that seems to be occurring between the two endemic São Tomé tree frog species, reported in a major publication earlier this year. She has also discovered that the little green tree frogs of Principe, while seemingly nearly identical to those of São Tomé, are in fact a separate species. This does not surprise us really; see earlier blogs on geckos and snakes!

Rayna is being assisted by our third new member,  Lauren Scheinberg of the Herpetology Department of the California Academy of Sciences.

Our brilliant (and very tall) photographer, Andrew Stanbridge, has joined us for the fifth time (also as co-leader).

The education team is, as usual, Roberta Ayers and myself, plus our long-time São Toméan colleagues, Roberta dos Santos, Anita Rodriguez and Quintino Quade Cabral. Maria Jeronimo has been assisting. A new cycle begins this year, starting again with third grade.

There have been several interesting discoveries already. A couple of years ago we published a study of the giant geckos of the islands, describing a new species from Príncipe (Miller, et al, 2012). We suggested that the large endemic species of São Tomé, Hemidactylus greefi, was only found in natural settings, having been out-competed in the towns by common, widespread recent colonizers. During our first week the group found an adult Greef’s gecko at sea level on a door in Angolares, the second largest town on the big island. I suspect that if competition with widespread common immigrants accounts for the absence of this gecko in the capitol city, there must be fewer of the former in Angolares. This is understandable as Angolares is still quite small and is not by any means a port city; that has always been São Tomé city.

 

As in every year, we went to the Olea tree at Macambrara (1100 m) to check on the known population of the giant São Tomé tree frog, Hyperolius thomensis. Long-time readers will recall that two holes in this enormous tree are the only place we have consistently found this colorful species. While we have conjectured that the giant tree frog must be widespread in the higher elevation forests (we can hear it call from far above in the canopy), we have been unable to find another locality or tree with appropriate breeding holes… until now!

With our colleague, Dr. Ricardo Lima of University of Lisbon, we were able to gain access to the primary forest above the huge southern oil palm plantation of Agripalma. At 350 m, above an abandoned roça called Monte Carmo, Bell, Scheinberg, Lima and Stanbridge found large numbers of the giant São Tomé tree frog breeding in pockets of water on fallen logs. This southern-most locality indicates, as we suspected, that this flamboyant frog is widespread in the relatively undisturbed forest and that while not restricted to high elevation (Macambrara), it does indeed seem to breed in pockets of water such as tree holes, rather than standing water like its close relative.

 

Here, there appears to be no hybridization as the two tree frog species are separated by oil palm rather than less biologically hostile agricultural fields or plots. In fact, so far as I know, hardly any endemic vertebrates or native plants are able to survive in oil palm.

This area is also the last bastion of a remarkable endemic bird, the Dwarf ibis, the smallest species of ibis in the world; it is severely threatened, both by hunting and by habitat destruction (oil palm). Notably, there are now two charismatic endemic species, a bird and a frog, endangered by human activities on this remote and fascinating island.

So far, this year’s activities have included working in a number of new localities including the central massif and in the far south. High in the mountains above Roça Agua Izé, one of the larger of the coastal colonial cacao plantations, we got our first decent view of a Giant weaver, Ploceus grandis. One of many endemic island giants on São Tomé and Príncipe, this colorful weaver is the largest in the world.

As always, our educational efforts are aimed at raising the children’s awareness of the unique aspects of the island flora and fauna. We do not preach conservation per se. but rather try to show the young students how special their islands are and thus how special they are as owners.

This year, each third-grader gets our coloring book about endemic bird species on both islands, a box of colored pencils and a pair of plastic binoculars (which work!!) These are not just handed out…. we present them personally to each student in each classroom, along with enthusiastic instructions for use, and the reasons  we come each year; we involve the students, the teachers, even school principals, and it is great fun. At the end, 10 of our stick-on logo patches are given to the teacher to reward good work. Each of us is involved at one time or another, as voices begin to suffer after 3 or 4 classroom visits.

At this point, I must mention some wonderful folks who have been vital to the education effort this year: Alice and Wayne Settle conceived of and sponsored the acquisition of the small binoculars; Jim Boyer of the California Academy of Science once again produced a booklet that qualifies as a work of art, and help with producing the bird books is coming from the Príncipe Trust.

In the next blog, I will report on our progress on the smaller geologically ancient island of Príncipe.

Meantime, here’s the parting shot:

 

All photos by Andrew Stanbridge

 

 PARTNERS:
We are most grateful to Arlindo de Ceita Carvalho, Director General, Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos and Quintino Quade of STePUP of Sao Tomehttp://www.stepup.st/, our “home away from home”.We gratefully acknowledge the support of the G. Lindsay Field Research Fund, Hagey Research Venture Fund of the California Academy of Sciences for largely funding our initial two expeditions (GG I, II). The Société de Conservation et Développement (SCD) and Africa’s Eden provided logistics, ground transportation and lodging (GG III-V), and special thanks for the generosity of private individuals who made the GG III-VII expeditions possible: George G. Breed, Gerry F. Ohrstrom, Timothy M. Muller, Mrs. W. H. V. Brooke, Mr. and Mrs. Michael Murakami, Hon. Richard C. Livermore, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. Robert M. Taylor, Velma and Michael Schnoll, and Sheila Farr Nielsen; GG VI supporters include Bom Bom Island and the Omali Lodge for logistics and lodging, The Herbst Foundation, The “Blackhawk Gang,” the Docent Council of the California Academy of Sciences in honor of Kathleen Lilienthal, Bernard S. Schulte, Corinne W. Abel, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. John Sears, John S. Livermore and Elton Welke. GG VIII was funded by a very generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Mrs. W.H.V.“D.A.” Brooke, Thomas B. Livermore, Rod C. M. Hall, Timothy M. Muller, Prof. and Mrs. Evan C. Evans, Mr. and Mrs. John L. Sullivan Jr., Clarence G. Donahue, Mr. and Mrs. John Sears, and a heartening number of “Coolies”, “Blackhawk Gang” returnees and members of the Academy Docent Council. Once again we are deeply grateful for the support of the Omali Lodge (São Tomé) and Roça Belo Monte (Príncipe) for both logistics and lodging and for partially sponsoring part our education efforts for GG VII and GG VIII.

Our expeditions can be supported by tax-deductable donations to “California Academy of Sciences Gulf of Guinea Fund”

The Race: The Blog Returns with a Science Update

“The Race” has been silent for a while; a sabbatical accompanied by computer glitches at both sites (Wildlifedirect.org; calacademy.org) led to it, but this was not meant to signal a pause in our island work by any means! We will be returning to the islands for two more expeditions later this year.

During the past nine months or so, some important scientific papers have been published by expedition members; these continue to illustrate the unique nature of the island fauna and flora.

Ricka Stoelting (D. Lin phot, GGI)

Ricka Stoelting was on the islands for a solid two months during GG I in 2001. The research she did on the unique Sâo Tomé caecilian was the basis for her MSc degree, and this has just been published; this paper is the first on the population genetics of a caecilian species,  and she has shed light on a number of issues involving this strange Sâo Tomé endemic.

(left) Cobra bobo, Schistometopum thomense (A. Stanbridge phot. GG VII); (right) from Stoelting, et al., 2014, PLoS One.

Ricka discovered that there are four distinct populations of the cobra bobo that are genetically different from each other – not different enough to be considered separate species, but different enough to suggest that these populations were isolated from each other in the distant past (recall that evolution is genetic change accumulated in isolation over time). She also found that these different populations were probably separated from each other through major geological changes in the environment. Note (above right) that the western populations (green) and northern populations (red) are associated with volcanic landscapes less than 1 million years old, while the southern populations (blue) are associated with older volcanic soils of about 2.5 million years in age. This suggests that the formerly widespread caecilian species was wiped out to the north by volcanism about 1 million years ago, then later repopulated from the surviving southern population. The yellow population is very young, perhaps only about 36 thousand years. Currently Ricka is pursuing a PhD at the University of Wisconsin.

Dr. Ricardo Lima with caecilian, Schistometopum thomense. (R. Ayres phot. GG. IV)

While on the subject of the endemic Sâo Tomé caecilian, I can report that our colleague, Dr. Ricardo Lima (above), just found three  caecilians while climbing the Pico de Sâo Tomé, and one of these establishes an altitude record for this strange worm-like amphibian species at 1504 meters (4,600+ feet).

Dr. Rayna Bell on Sao Tome.  (A. Stanbridge phot, GG VI)

Last September, Rayna Bell, a participant on both GG VI and GG VII expeditions, completed her PhD degree at Cornell University; her doctorate was based on her work on the Hyperolius tree frogs of the islands, and the first part of it has just been published.

(left) after Bell, R.C. et al. 2014. Journal of Biogeography. (right). Nearest mainland relative of island Hyperolius tree frogs. (RDC phot)

We knew from earlier work that the two tree frogs of Sâo Tomé were each other’s nearest relatives.  But where did they come from originally?  Dr. Bell’s research indicates that the nearest relative of both the endemic oceanic treefrog and the Sâo Tomé Giant treefrog is a member of a large group of species in West Africa, Hyperolius cinnamomeoventris (above right); in particular, a subset of this group she terms “clade A,” (indicated by green stars above left) appears to share joint ancestry with the island species.

Hypothesized large “riverbank” raft.  (artwork by R.E.Cook)

Her research suggests that the common ancestor of the island treefrogs reached Sâo Tomé first, in a single colonization event, probably by rafting (above). This event likely occurred between 9 and 3.5 million years ago, and the original colonists probably originated from the Ogooué or Congo Rivers. On Sâo Tomé, these original “pioneers” differentiated into a giant highland form (now H. thomensis) and a smaller lowland species. Then, between 1.1 million and 270 thousand years ago, members of the lower elevation species, the oceanic treefrog, Hyperolius molleri dispersed to the much older island of Príncipe, where they again became isolated from the parent population on Sâo Tomé and began to accumulate genetic change (“speciation”). A second scientific paper by Dr. Bell on these species and their evolution on the Gulf of Guinea Islands is due out soon.

The status of the Sâo Tomé shrew (Crocidura thomensis) has been somewhat problematic, for scientists at least. Shrews are very poor overwater dispersers, and since it is possible for the shrew to have been transported from the mainland by man fairly recently, there have been questions about its validity as a true endemic species. Did it reach Sâo Tomé by natural means?  This strange insectivore was recorded only nine times after its original description in 1887 and until recently, it has also been considered quite rare. (Someone should have asked the locals!)

The Sâo Tomé  shrew (phot. Dr. Mariana Carvalho).  (left) Dr. Ricardo F. de Lima.

In a recent publication, (below) our colleague, Dr. Ricardo Lima has added twenty-three new locality records for this fascinating creature, indicating that it is not rare. It has just been overlooked. An excellent ecologist, de Lima has assessed the conservation status of this important species.  Moreover, he and his colleagues have provided tissue so that we have been able to examine its DNA to determine whether or not it arrived on the island by natural means or was more recently brought by man. It appears to be a true endemic species (to be reported in a future publication), and as such it is the only non-flying endemic mammal species on the island of Sâo Tomé.

From de Lima, R.F., et al]. 2015 Fauna & Flora International. Oryx. Grey dots denote localities known in 1996. Black dots are new localities.

During GG II (2006), we collected two shrew new-borns on the island of Príncipe which at the time we assumed were a mainland species, Crocidura poensis. Dr. Luis Ceriaco, who is now an adjunct member of our CAS faculty, subsequently obtained many more adult individuals on the island (it is much more common than C. thomensis) and surprisingly this species is also new and endemic.

From Ceriaco, et al. 2015. Mammalia

Dr. Ceriaco has just formally described it as Crocidura fingui, its name in the local Creole. So the only non-flying endemic mammals in the Republic of Sâo Tomé and Príncipe are shrews! I do not think any biogeographer would have predicted this (see August 2010 blog: the “Magic of Molecules).

Here’s the parting shot:

Rotula deciesdigitatus, one of the rarest sand dollars in the world; in the islands, known principally from Praia Morrao, west coast of Sao Tome. (Weckerphoto, GG III)
PARTNERS

We are most grateful to Arlindo de Ceita Carvalho, Director General, Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for their continuing authorization to collect and export specimens for study, and to Ned Seligman, Roberta dos Santos and Quintino Quade of STePUP of Sao Tomehttp://www.stepup.st/, our “home away from home”. We gratefully acknowledge the support of the G. Lindsay Field Research Fund, Hagey Research Venture Fund of the California Academy of Sciences for largely funding our initial two expeditions (GG I, II). The Société de Conservation et Développement (SCD) and Africa’s Eden provided logistics, ground transportation and lodging (GG III-V), and special thanks for the generosity of private individuals who made the GG III-VII expeditions possible: George G. Breed, Gerry F. Ohrstrom, Timothy M. Muller, Mrs. W. H. V. Brooke, Mr. and Mrs. Michael Murakami, Hon. Richard C. Livermore, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. Robert M. Taylor, Velma and Michael Schnoll, and Sheila Farr Nielsen; GG VI supporters include Bom Bom Island and the Omali Lodge for logistics and lodging, The Herbst Foundation, The “Blackhawk Gang,” the Docent Council of the California Academy of Sciences in honor of Kathleen Lilienthal, Bernard S. Schulte, Corinne W. Abel, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. John Sears, John S. Livermore and Elton Welke. GG VIII was funded by a very generous grant from The William K. Bowes Jr. Foundation, and substantial donations from Mrs. W.H.V.“D.A.” Brooke, Thomas B. Livermore, Rod C. M. Hall, Timothy M. Muller, Prof. and Mrs. Evan C. Evans, Mr. and Mrs. John L. Sullivan Jr., Clarence G. Donahue, Mr. and Mrs. John Sears, and a heartening number of “Coolies”, “Blackhawk Gang” returnees and members of the Academy Docent Council. Once again we are deeply grateful for the support of the Omali Lodge (São Tomé) and Bom Bom Island (Príncipe) for both logistics and lodging and for partially sponsoring part our education efforts for GG VII and GG VIII.

Our expeditions can be supported by tax-deductable donations to “California Academy of Sciences Gulf of Guinea Fund”

The Race: On Rocket Frogs and Millipedes

First, some great news on the academic front.  One of the graduate student participants on GGI back in 2001 just completed his PhD at the University of California, Berkeley. Meet Dr. Joel Ledford, newly-minted world authority on spiders and explorer of Gulf of Guinea biodiversity.  In the picture below, he is holding “bubba,” one of the three endemic tarantula species of São Tomé.

DR. Joel Ledford with Hysterocrates apostolicus.   D. Lin phot. GGI

Readers of this blog already know that when we talk about biodiversity, we are talking about everything living, not just the big fancy stuff like birds and giant begonias.  Many of the secrets of island evolution are to be unlocked through the study of small organisms.   I have just received some preliminary news from Dr. Didier Van den Speigel of the Royal Central African Museum in Belgium.  After Dr. Rowland Shelley of the North Carolina State Museum did a preliminary analysis of our GG IV millipede specimens, we sent them to Didier, a specialist on this group in the Old World.  Rowland had concluded that we had at least one new species of the genus Globanus from each island.

A millipede (not Globanus) phot.  from cephalopodiatrist.com

Didier has examined material from other museums and has concluded that, in fact, the genus Globanus itself is endemic, found nowhere else in the world but the islands of São Tomé and Príncipe.  We are still unsure of how many species our GG IV material represents, but what seems evident at this time is that they are all each other’s closest relatives.  Drs Van den Speigel and Shelley are in agreement that this turns out to be the case, it would represent a “species swarm,” much like the endemic earthworms of São Tomé (see July 2010 blog “Nightmares….”, for an explanation).  The work continues……

In my memorial to Abade last month, I described one of our early unsuccessful  searches for Newton’s rocket frog during GG I.

Newton’s rocket frog, Ptychadena newtoni.  D. Lin phot.  GGI

This widely distributed genus of about 50 species is found throughout sub-Saharan Africa and distinguished by a sharp snout, paired vocal sacs (lower arrow),  distinctive glandular ridges on the back (upper arrow) and very long legs.  In fact a member of this group from South Africa holds the world record frog jump of over 33 feet (10m)!  P. newtoni is one of São Tomé’s classic “island giants; at 76 mm (not including legs) a  São Tomé female is much larger than any specimens of mainland species on record.

After days of visiting known localities mostly in and near the town of São Tomé and finding them dry, heavily disturbed and frogless, one rainy evening two young boys led us to a vacant lot less than 200m from where we were living, and there were the frogs!   Ultimately, genetic analysis of these frogs established that they were, indeed, full endemic species, but also led John Measey, currently of South Africa, and a group of us to publish our rafting hypothesis in the Journal of Biogeography (2007 – see earlier blogs).

Our difficulty in finding this species in the northern lowlands of São Tomé (all of the known localities at the time) suggested to me that this may be the only endemic amphibian species on São Tomé that might be endangered due to human development.

Series of Ptychadena newtoni larvae from Java, Sao Tome.  RCD phot. GG II

However, during GG II we found a series of tadpoles at Java (elevation 595m) which we later determined belonged to this species (although no adults were seen).  Tadpoles are typically identified by various external characteristics, but especially by fine structures of the mouthparts. The drawings below are taken from a nearly completed manuscript that attempts to technically describe the tadpoles (larvae) of all the endemic island frog species; it has not been published because, even after all these years, we have still not found the larvae of the Príncipe giant treefrog, Leptopelis palmatus!

P. newtoni mouthparts from unpublished manuscript.

P. newtoni left lateral view from unpublished manuscript.

Our discovery of the Java larvae indicated that Newton’s rocket frog is not necessarily present only in the heavily developed northern lowlands.

Recently, a young biologist, Hugulay Maia, whom we first met during GG IV has found some new P. newtoni localities.

Hugulay Maia of ABS, doing tree work.  unknown photographer]

Hugulay is a member of Associação dos Biologos (ABS), a local group of biologists involved in biodiversity efforts on São Tomé. The group is led by Dr. Alzira Rodrigues of the Polytechnic Institute; other members you have met in this blog are Angus Gascoigne and Victor Bomfim.

Current P. newtoni localities: green = to 1992; pink = to date

Now, thanks to Hugulay’s observations (and photographs) we have a somewhat better idea of the distribution of Newton’s rocket frog.  Earlier known localities are in green and were published by a Swiss worker in 1992; our GG II Java locality and Hugulay’s new localities are in pink.  Hugulay’s data confirm that the species is not confined to the north.  He has observed it at Colonia Açoreana (labeled) and two more southerly spots, Angra Toldo Cavaleite and Roça Alinhança.

The data are still thin, but we can at least infer that Ptychadena newtoni is more widespread than originally thought.  Almost all of the mainland species breed in relatively still or slow-moving water, and it is reasonable to assume this is the case with Newton’s rocket frog.  All of the old localities (in green) are associated with lowland reaches of major water courses: the city localities are in the Agua Grande drainage; Hugulay Maia’s new records are all from the Ribeira Afonsa drainage, and the Diogo Vaz locality (green symbol in the NW) is from the small Agua Anambo, which parallels the larger, much faster Rio Maria Luiza to south.  Java our highest locality is on the Rio Abade, but the tadpoles were collected in a man-made pool in a roadside, partially dry creek bed, not in the river itself.   To assess the actual status of Newton’s rocket frog, I think we just need to look more closely in bodies of slow or still water along major rivers throughout the island.

The Parting Shot:

Dr. Joel Ledford: Spider hunters in repose. D. Lin phot.   GG I

PARTNERS

We gratefully acknowledge the support of the G. Lindsay Field Research Fund, Hagey Research Venture Fund of the California Academy of Sciences, the Société de Conservation et Développement (SCD) and Africa’s Eden for logistics, ground transportation and lodging, STePUP of Sao Tome http://www.stepup.st/, Arlindo de Ceita Carvalho, Director General, and Victor Bomfim, Salvador Sousa Pontes and Danilo Barbero of the Ministry of Environment, Republic of São Tomé and Príncipe for permission to export specimens for study, the continued support of Bastien Loloum of Zuntabawe  and Faustino Oliviera, Curator of the Herbarium at Bom Sucesso. Special thanks for the generosity of private individuals, George G. Breed, Gerry F. Ohrstrom, Timothy M. Muller, Mrs. W. H. V. Brooke, Mr. and Mrs. Michael Murakami, Hon. Richard C. Livermore, Prof. & Mrs. Evan C. Evans III, Mr. and Mrs. Robert M. Taylor, Velma and Michael Schnoll, and Sheila Farr Nielsen for helping make these expeditions possible.  Our expeditions can be supported by donations to “California Academy of Sciences Gulf of Guinea Fund”.

The Race: Within the House of Slytherin (I. Lizards)

Our race to discover and describe the unique fauna and flora of São Tomé and Príncipe continues, and the six members of Gulf of Guinea Expedition III (B) are diving in the ancient waters of Príncipe as I write; they return to the Academy next week.  As I wrote earlier, Marta is sampling the sea slug fauna (nudibranchs), Gary, Bob and Dana are looking at coral and barnacles, having found a new species of the latter in waters off São Tomé during GG II, and John and David are looking at small marine fish, with emphasis on eels.  The group has an added goal, and that is to bring back some freshwater prawns (Macrobrachium) that abound in the São Tomé rivers. These specimens are for a young high school student named Alex Kim.

  

 A freshwater Macrobrachium prawn from Guinea (www.)

Alex is a senior at Thomas Jefferson High School of Science and Technology in Virginia.  He is doing an ambitious biogeography project on these prawns, relatives of which are found on both sides of the Atlantic.  Alex contacted me through this blog—you can read his comments at the end the November posting.  During a brief visit to DC over the holidays, I brought some preserved specimens we collected in GG I and GG II which I handed over to one of his advisors, Dr. Patrick Gillevet of George Mason University, and now the GG III (B) group plans to bring him some fresh material for DNA studies.  This is really fun academic stuff, and we are delighted to have the involvement of a young colleague.

 

 A Macrobrachium prawn from Cameroon. (www) 

Except for documenting our exciting hunt for Príncipe Jita, (see first May posting), I have not written that much about the endemic reptiles of these islands; in fact, there are quite a few of them, some rather spectacular.  While reptiles, especially geckos and skinks, are much better dispersers over saltwater than amphibians, snakes are not particularly good at it; moreover, like the amphibian caecilian, cobra bobo, a number of these endemics are legless species.  There are also some island species that may be endemic, but we are not sure…. we just haven’t studied them closely enough yet. In this posting I will show you the unique lizard species.  One readily identifiable endemic species is Greeff’s gecko, or the Giant gecko, Hemidactylus greeffii.  

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Greeff’s Gecko, Hemidactylus greeffii . A Sao Tome specimen. RCD phot. GG I   

Greeff’s gecko is an island giant; it is evidently much larger than other African member of the genus (and there are over 55 African taxa of Hemidactylus with likely many more to be discovered). Our longest specimen is over 200 mm in total length (including original tail); but longer specimens are known.  This gecko is not only very large it also differs from all of its African relatives in lacking a claw on the first (inner) finger and first toe. Somehow, this feature has been lost during the thousands, perhaps millions of years of isolation on the Gulf of Guinea Islands. Greeff’s gecko also has greenish eyes, which also distinguishes it from other nocturnal geckos on the island which, so far as we know, are not endemics.

 

 H. greeffii.  Note absence of claw on first thumb. ST specimen. RCD phot. GGI

 

H. greeffii with  greenish eyes.  ST specimen. D. Lin phot. GG II.

Greeff’s gecko occurs on both São Tomé and Príncipe; at least we think it does. Here’s what I mean: specimens from both islands look very much the same but a couple of years ago, a group of researchers from the University of Madeira and Portugal looked at the DNA of specimens from both islands and found that data from mitochondrial DNA suggested the two populations were very different, and that they may well be two distinct species in spite of their apparent anatomical similarity. These results were not confirmed by study of nuclear DNA however, so scientifically the “jury” is still out, and we call both island forms, Greeff’s gecko. This critter is quite common in rock walls, culverts, rock crevices on both islands and is strictly nocturnal. 

 

Principe specimen of Greeff’s gecko. D. Lin phot. GG II.  

A similar situation exists with a small terrestrial skink called Panaspis africana, or Gulf Leaf-litter skink. A daytime forager, this small uniform-brown skink is very common in the lowlands; it can be easily heard and seen scuttling through dried cacao leaves and it is almost always found on the ground on both islands; one of our largest gravid (with eggs) female specimens from São Tomé is about 100 mm in total length, but most of our examples are smaller.  

Gulf Leaf-litter skink. Panaspis africana; D. Lin phot. GG II. 

The same group of researchers from the University of Madeira studied the DNA of leaf litter skinks of both islands, and also Annobón, the last island in the chain and part of Equatorial Guinea.  They used, in part, tissues and specimens collected by us during GG I in 2001.  In this case they found clear evidence for three separate species, one on each island (the one on Annobón is already called P. annobonensis); this was supported by both mitochondrial and nuclear DNA sequence.  However, in one of those tragic, fortunately rare, occurrences in science, the specimens from which the tissue samples were taken were either lost in transit or misplaced.  Without voucher samples the results cannot be duplicated or tested nor can we demonstrate the results.  So for now, although there was evidence that Panaspis is two different species on São Tomé and Príncipe we cannot confidently describe the populations of the different islands nor give them scientific names.  Until the study can be redone with new material, the Gulf leaf-litter skinks remain known as simply Panaspis africana. 

Author working on Principe.  Weckerphoto. GGIII  

The way we collect these specimens is not sophisticated – we use our hands. We turn over logs, rocks and branches on the ground or sift through leaf litter with rakes; we climb trees and cliffs; we go out at night with flashlights and headlamps. After capture, the specimens are put in separate plastic bags for later processing.  

 

Dr. Iwamoto in Sao Tome H. greeffii habitat on Sao Tome. RCD phot. GG I 

Jens Vindum searching leaf litter on Sao Tome. D. Lin phot. GG II 

 

Principe day gecko in plasic bag. RDC phot. GG III.

Every specimen we collect gets a unique field number, which is the same used for photographs of it, recordings or tissues samples taken. 

 

My grad student, Ricka Stoelting,  processing specimens on Principe. RCD phot. GG I 

Certainly one of the oddest endemic lizards is the legless skink, unique to Príncipe Island, Feylinia polylepis.  There are about six species known in this genus, the remaining five found broadly distributed on the African mainland.   

 

Principe legless skink, Feylinia polylepis. brown  phase. D. Lin phot. GG I. 

 They appear in two different color morphs, a brown one and a pale gray one, regardless of size or sex.  The locals call them, Ozhgah (or at least the name  sounds like that). 

 

Principe legless skin – grey phase.  D. Lin  phot. GG II 

 

Feylinia polylepis head shot. D. Lin phot. GG II 

They can be found under almost anything on the ground provided the earth is slightly moist. Once exposed, they are very quick and can rapidly disappear into holes in the ground. They are conspicuously common in the Príncipe lowlands, and in this regard are reminiscent of the caecilians of São Tomé Island; the high density of their numbers in suitable habitats suggests predation may be low in these areas. 

Not all geckos are nocturnal.  In the Old World there are two large groups that are secondarily diurnal, although they, like all geckos, lack eyelids.  The genus Phelsuma is a group of numerous species of velvety green geckos found on Madagascar and the Indian Ocean islands; the other group, Lygodactylus are also present in Madagascar but also distributed throughout the Afrotropical region as well.  They are not brightly colored, and taxonomically rather poorly known.  The group as a whole is being studied by Dr. David Vieites (and his students) of Madrid and Dr. Adam Leache, of the University of Washington.  I have been involved as well but largely in studying the relationships of day geckos of the Gulf of Guinea Islands.  

 Lygodactylus thomensis. Sao Tome.  D. Lin  phot. GGI 

The Gulf of Guinea Day geckos are sun-lovers and strictly climbers, being fairly common on tree trunks and scuttling up walls even in São Tomé town and Santo Antonio, Príncipe.  They are very small, at about 70-80 mm total length.  The day geckos of the Gulf of Guinea islands (excluding the continental island of Bioko) have long been recognized as a distinct, endemic species, Lygodactylus thomensis, first discovered on São Tomé Island.  The day geckos on Príncipe and Annobón have been described as subspecies (or races, if you will) of the São Tomé species..  As you can see from the illustration below, one of the characteristics used to define species of day geckos is the throat pattern. 

 

 Day geckos of the Gulf of Guinea Islands.  RCD prep.

The throat patterns of the lizards on each of the three islands are quite consistently distinct from one another, and work by us and the University of Madeira suggest that they have been isolated from each other for a long, long time, and that each is a full species unique to its island. Work is continuing on these lizards.

 

 L. delicatus of Príncipe Island. RCD phot. GG III

 There are other conspicuous lizards on both islands but these are not considered endemics; i.e., they occur elsewhere and are probably just good over-water dispersers. The large speckled-lipped skink, Mabuya maculilabris, is common and widespread in the lowlands of both São Tomé and Príncipe. It is a good climber and is seen in a variety of habitats especially along the coast lines.  This species also broadly distributed on the African mainland.

 

 Speckle-lipped skink (Mabuya maculilabris) of the Gulf of Guinea. Sao Tome. D. Lin phot. GG II] 

 

M. maculilabris detail. D. Lin phot. GGII 

There are also non-endemic, nocturnal geckos on both islands. Most appear to be the widespread house gecko, Hemidactylus mabouia, also occuring nearly throughout Africa. 

 

House gecko, Hemidactylus mabouia.  D. Lin. phot.  GG II] 

Note that the eyes are not greenish and that this species does not lack claws on the inner toe and finger.  There is some confusion as to how many non-endemic species are present and what to call them. 

H. mabouia foot from beneath. note claws. Weckerphoto GG III.

Snakes are coming next. 

Here’s the parting shot:  

The thrill of discovery! Bom Bom Island, Principe.  Weckerphoto. GG III  

PARTNERS We gratefully acknowledge the support of the G. Lindsay Field Research Fund,  Hagey Research Venture Fund of the California Academy of Sciences, the Société de Conservation et Développement  (SCD) for logistics, ground transportation and lodging, STePUP of Sao Tome http://www.stepup.st/, Arlindo de Ceita Carvalho, Director General, Victor Bomfim, Salvador Sousa Pontes and Danilo Bardero  of the Ministry of Environment, Republic of Sao Tome and Principe for permission to export specimens for study, and the continued support of Bastien Loloumb of Monte Pico and Faustino Oliviera, Director of the botanical garden at Bom Sucesso. Special thanks for the generosity of private individuals, George F. Breed, Gerry F. Ohrstrom, Timothy M. Muller, Mrs. W. H. V. Brooke and Mr. and Mrs. Michael Murkami for helping make these expeditions possible.  

The Race: A Toad Less Traveled

 Sorry, I love titles like this… and I have more!  Actually, there are no toads (Bufonidae) on São Tomé and Príncipe; interesting in itself because seven other amphibian species of five different families have survived the ocean crossing during the many millions of years since the islands first emerged. Moreover, toads are common in almost every  conceivable terrestrial mainland habitat.  

My last two blogs have been a bit academic.  Having laid the biogeographical ground work, it is probably time to get back to the unique, endemic island critters.   The tiny, 31-million year old island of Príncipe is the only home of Africa’s largest treefrog, Leptopelis palmatus – the Príncipe Giant Treefrog.  It is one of the world’s rarest frogs, as well.

 

 L. palmatus – D. Lin phot. GG I 

Let me be quick to point out that the Príncipe critter is not Africa’s largest frog; that title belongs to Conraua goliath (below), which is found on the mainland in southern Cameroon and Gabon.  In fact, the goliath frog is the largest in the world but it is not related to any on São Tomé and Príncipe– no members of its family have made it across the saltwater gap to the islands, or if they ever did, they have not survived. 

  

Conraua goliath—J.-L. Perret phot. 

Leptopelis palmatus is the largest African treefrog (emphasis on “tree”) — frogs that are adapted for climbing with, among other features, enlarged finger and toepads.  As I have pointed out in earlier blogs, gigantism is a relative thing; the giant endemic plants, frogs, birds and lizards of São Tomé and Príncipe are bigger than all of their relatives but they are not necessarily so large you trip over them (like Galapagos or Aldabra tortoises); they are simply larger than all of their relatives.  The two images below put this frog in some perspective, and I think you will agree that this is one BIG treefrog. 

Me, with the first female. R. Stoelting phot. GG I 

 

The frog on Dong Lin, our photographer.  R. Stoelting phot GG I

This species was first described in 1868 on the basis of a single female specimen, housed in the Berlin Museum. At the time of GG I in 2001, the Príncipe giant treefrog was known only from this single type specimen and seven additional specimens, all females, collected by local Príncipeans for a Swiss colleague named Catherine Loumont.  The largest of Loumont’s specimens is 110 mm from snout to vent (we do not include legs when we measure frog sizes), and even after our years of work, this specimen remains the largest ever found – it is nearly 30 mm longer than the largest of its nearest mainland relative, Leptopelis macrotis, distributed from central Sierra Leone to Ghana. One of several differences between the two species is the striking deep-red eyes of our island endemic. 

The eye of the Príncipe giant treefrog. D. Lin phot. GG II 

This first specimen we found during GG I (first three treefrog images, above) was yet another female, 108 mm in length.  Our mammalogist, Doug Long, was led to the critter by some kids from the now-defunct plantation of Sundi in northwest Príncipe. Sundi may no longer function as a plantation but it is still inhabited by the descendents of former workers—lots of them, there is even a mayor. 

 

Doug Long and the Sundi kids. RCD phot. GG I 

The arrival of this frog was greeted with great enthusiasm by yours truly; here in my hands one of the rarest frogs in the world! And it was huge! I was not surprised to learn that it had been found on the ground, as it is hard to imagine something so bulky climbing around in bushes and trees. The male of this species was completely unknown, so far as we knew at the time,.  None had ever been collected, photographed nor described in the scientific literature.  So we also knew nothing about the species’ breeding biology, male advertisement call or tadpole.  At the time, we were unaware of a blog posted two years before our visit by Jonathan Bailey on the Gulf of Guinea Conservation Group website, entitled “One month in the Forest of  Príncipe.” Jonathan (now Dr.) Baillie described hearing the calls of male L. palmatus as “like a pop bottle being continuously opened.”  He heard them high up on Pico do Príncipe near a small stream at about 700 m and actually collected two of them which had been deposited in the Natural History Museum in London.  But during GG I, the male giant treefrog was terra incognita, so far as we were concerned.  

Second female from Rio Papagaio.  J. Ledford phot..  GG I 

During a second GG I visit to Príncipe a few weeks later, my then-graduate student,  Ricka Stoelting, collected another female along the Rio Papagaio, a large-ish river that flows through Príncipe’s only town, Santo Antonio.  It was also of a rather dull in color but with white spots. We have since learned that this is about as brightly colored as females get. 

Rio Papagaio in town, downstream. RCD phot. GG III

 

Ricka Stoelting, my graduate student on Sao Tome.  RCD phot. GG I.  

During this second visit, Ricka and Dr. Sarah Spaulding ascended Pico do Príncipe to the top and camped at nearly the same spot where Jonathan Baillie had been two years before.  There she found the males, lots of them, calling from bushes and branches at night near a very small creek.  

 

Tiny creek on the Pico.  J. Uyeda phot. GG II

 Ricka brought the series of males back down the mountain, and they were astounding.  Unlike the females they were very brightly colored and highly variable, in pattern, as well; this variability is rather unusual in frogs, although there are some species that are sexually dimorphic for color. And they were much, much smaller than the females, though we knew they were full-sized breeding adults. During later analysis we learned that the largest breeding males are only about 41% of the size of the largest females, a size disparity that is striking.

First series of live males (far right is a juvenile).  J. Ledford phot. GG I

  Ricka never heard them calling and anyway she had no way of recording them if they had.  One of the parameters we use in establishing relationships among frog species is analysis of the voice (or advertisement call.). Males call to attract females, and at the same time to advertise their presence and territory to other males.  The advertisement call is species- specific and obviously adaptive when there are other species utilizing the same water for breeding.   To really define Leptopelis palmatus, I needed a recording of the voice, and this was to become a priority in the future. Below is a preliminary analysis of the call of another Gulf of Guinea frog species which we think is present on both islands.  Here, we are comparing the advertisement calls of males from two different localities on both islands, and we can see that they are basically the same.

 

 Preliminary sonograms of Oceanic treefrog. Marshall/Drewes construct.  

 Back at the Academy, Ricka and I prepared the first formal description of male Príncipe giant treefrogs.  Now aware of Baillie’s blog, we read his word description of the advertisement call.  Although the Principeans insisted the frogs did call, it remained an open question, especially when I learned from anatomical study that the male frogs  lack vocal sacs and vocal sac openings, features that most calling frogs possess (including other members of the genus Leptopelis).   GG II in 2006 included Josef Uyeda as my student (now a PhD candidate at Oregon State University).  Josef was working on a different group of island endemics called puddlefrogs (see earlier blog: “We Find Jita”) but when we were on Príncipe, I sent him up the Pico with his friend Mac and the same guide, Manona, who had led Jonathan Baillie and Ricka years before. They were armed with my old Sony cassette recorder (my iPod had failed).  Bear in mind that the only known localities for males were at nearly 700 m, high on the Pico and while this made no biological sense, that’s where my stalwarts had to go. This is no small matter given the topography of the island, but graduate students are good at this sort of thing and anyway, they tend to be younger and more vigorous than their advisors!

Principe terrain. Pico do Príncipe is in the clouds to the left of the large Pico Papagaio. R. Wenk phot. GG III

 

 Josef Uyeda hunting for caecilians on São Tomé. D. Lin phot. GG II  

While in the same general area as earlier workers at about 700 m, Josef got a lot done but the party was caught in heavy rains.  He heard males and saw them calling but only managed some rather distant, poor-quality recordings (the conditions were miserable), but now at least we knew that the frogs did, indeed, call.  GG III, last spring, provided some answers, thanks in part to our friend Ramos of Bom Bom Island.  Ramos is assistant manager of the resort, a native Principean and a keen, observant naturalist.  See the photo of Ramos in the “We Find Jita” blog.  I described our past difficulties in trying to record the voice of the Príncipe giant treefrog to him, and he grinned and said, We will go to my roça (farm) on Pico Papagaio and at 5:30, we will get them!  I was highly skeptical…

Roça Papagaio, Ramos’s farm at 250 m. R. Wenk phot. GG III 

Ramos’s farm is in the forested area on the northern flanks of Pico Papagaio at about 250 m.  Just before you reach it on a dirt steep uphill road, you cross a tiny creek; this is where Ramos took us –  about 30 m up that small creek, thick with dense undergrowth, and there we sat, waiting for the forest cacophony of grey parrots, mona monkeys to subside.  Nothing much happened.  I had my iPod with recording head at the ready.  We waited in the gathering gloom for about 20, maybe 30 minutes, Ramos grinning throughout and occasionally exclaiming,  Just wait. We will get them! 

Me waiting, iPod in hand, for the giants to call.  T. Daniel phot GG III 

And sure enough, we began to hear frogs calling. I looked at my watch. It was 5:30. The call is certainly a strange one; it lacks resonance (remember males don’t have a vocal sac) and thus it is rather flat and unmelodius. Rather than my trying to describe it or arguing with earlier descriptions, you can listen to it yourself by clicking this link: 

Click here to listen

And here are a couple of photos of the male that was calling, taken by Wes. These are un-posed and taken before we collected it as a voucher specimen for the voice:

 Weckerphoto GG III

 Weckerphoto – GG III 

There are still great gaps in our knowledge of this most unique frog.  Obviously, the males are well-distributed in the lower elevations; we just have not been in right place at the right time.  We still cannot explain why females are dull and rather cryptic in coloration and usually found on the ground, while there appears to be no selection for color in males.  The dull color of females seems consistent, as a couple of months ago I found six additional females (no males) collected in 1988 at the Doñana Institute in Seville and they were clearly drab in life; my colleagues at Donana tell me they were collected on the ground in lowland localities at Rio Papagaio and Bela Vista.

Six female Seville specimens at Donana Institute.  RCD phot. 

We still have not observed breeding, nor have we ever seen tadpoles.  In this genus, Leptopelis, they are very distinctive, and I would predict the tadpole will look like this: 

 

A Leptopelis tadpole. Image courtesy of Dr. R. Altig 

Here’s the parting shot: 

 

Nezo, of Angolares, Sao Tome: artist, musician, restaurateur and worthy man – Weckerphoto GG III

PARTNERS We gratefully acknowledge the support of the G. Lindsay Field Research Fund,  Academy Research Venture Fund of the California Academy of Sciences, the Société de Conservation et Développement  (SCD) for logistics, ground transportation and lodging, STePUP of Sao Tome http://www.stepup.st/, Arlindo de Ceita Carvalho, Director General, and Victor Bomfim, Salvador Sousa Pontes and Danilo Bardero  of the Ministry of Environment, Republic of Sao Tome and Principe for permission to export specimens for study, and the continued support of Bastien Loloumb of Monte Pico and Faustino Oliviera, Director of the botanical garden at Bom Sucesso. Special thanks for the generosity of four private individuals, George F. Breed, Gerry F. Ohrstrom, Timothy M. Muller and Mrs. W. H. V. Brooke for making these expeditions possible.