Category Archives: millipedes

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

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


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, 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: Nightmares, Explosions and Geographic Origins.

Before I get to expedition updates and cool biodiversity stuff, I must say I had recent cause to reflect on why I originally titled this blog, “Island Biodiversity Race.”  As you know, major changes are coming to the islands, and we are desperately trying to learn as much as we can about their unique fauna and flora before these changes occur.

Photo by Polly Anderson, June 24, 2010

This thing appeared off the dock of my friend Ned Seligman’s house in Sâo Tomé  town on June the 23rd.  So far as I know, it is still there; Ned and his people do not seem to be able to learn what a floating oil rig is doing there, who brought it, etc.  He emailed this image to me entitled, “Your Worst Nightmare.”

For blog newcomers, the exclusive economic zone of the Republic of Sâo Tomé and Príncipe impinges on oil-bearing ocean floor, and the tiny island nation is in partnership with Nigeria to exploit these resources.

Exclusive economic zone of The Republic of Sao Tome and Principe.]

Without going into the record of environmental destruction associated with this industry, one can say at the very least that a large influx of wealth can be anticipated, along with the sorts of “development” activities that follow capital influx; and these activities almost always have a negative environmental effect.  That’s why we are in a race to find out what’s there, so that we can inform the citizens (and the world) of what they have.  Hopefully, when they have to make decisions down the road, they will have some knowledge of the potential environmental costs.

There are already plans on the big island for a deepwater harbor and a significant extension of the runway at the airport.  This is not rumor; on GG IV I spoke with several South Africans who are members of the group that is involved in these projects.

Aerial photo of Laguna Azul by Joacquim Wirth, 2010

In my view, a truly distressing plan is that of a different South African development group for Laguna Azul, one of the most beautiful and accessible sites on the big island.  This cove on the northwest part of Sâo Tomé not far from the captol is as its name suggests, famous its beauty.  It is the only spot on the islands where the baobab tree is found (Adansonia digitata), and I have watched Sao Tomean families picnic there.   My understanding is that this development group plans to reroute the main road away from Laguna Azul, then construct properties and sell them off as kind of a gated community.

A Sao Tomean Picnic at Laguna Azul [Weckerphoto, GG III]

There are rather frightening rumors about how permission for this project was obtained but as they are rumors only, I will not repeat them here.

Time for critter news.  A few weeks ago, our entomology department sent off the specimens from our millipede transacts to Dr. Rowland Shelley of the North Carolina State Museum. Recall that he had kindly agreed to try to identify them for us.  He has a wonderful website on myriapods: So far as we know, only one short paper on the island millipedes has been published; curiously, this paper, written in 1993, was based on a small collection made on the road to São Nicolau in the vicinity of Nova Moca, where our biologist friend Ricardo Lima lives. By chance our first GG IV millipedes came from nearly the same spot, under a log shared with a cobra bobo (Schistometopum thomense)!

Ricardo Lima at Nova Moca with cobra bobo from under our first millipede log. [ R Ayres phot.  GGIV]

Over the month, we collected millipedes in ten different locations ranging in elevation from sea level to 500+ m on Príncipe (first collections ever?) and 1500 m on Sâo Tomé.  Dr. Shelley sent me a preliminary report that in his opinion, our specimens included at least one undescribed species from the big island and perhaps two from Príncipe.  This is not surprising news, given what we have been finding over the past ten years, but what is really exciting is the fact that Dr. Shelley believes all our six + species of millipedes belong to the same family (Spirostreptidae) and genus, Globanus!  This lack of taxonomic diversity might seem disappointing but it is not; in fact, it is rather intriguing.  We may be learning more from what is not present, than what is.  The specimens are now going to Dr. Didier Van den Spiegel of the Royal Museum for Central Africa in Tervuren, Belgium,  a specialist on African myriapods.  If Dr. Shelley’s preliminary identifications are confirmed by Dr. Van den Spiegel, the modern oceanic island  millipede fauna may be a result of what is known as a “species explosion,” or a “species swarm.”

A Spirostreptid millipede from the Grenadines [M. de Silva phot.]

Imagine an “ancestral” form of Globanus somehow reaching (by raft?) Príncipe and/or Sâo Tomé at a time when the islands were devoid of any other millipedes.  Such a situation would present lots of empty niches into which, in the absence of any competition, the ancestral lineage would radiate, perhaps explosively.. Nature abhors a vacuum! A widely used example of this phenomenon is the swarms of different species of cichlid fishes in Lake Tanganyika, all descendants from a single colonizing species.

Species swarms have apparently occurred on other Atlantic islands, Madeira being a prime example, with earwigs (Perirrhytus), true bugs (Chinacapsus), nine species of the beetle genus, Trechus and a woodlouse genus, Procellio.

Non Gulf of Guinea Earthworm. photo-Kent Simmons from www.]

Another update:  samples of earthworms collected on Sâo Tomé in 2002 by my colleage, Dr. John Measey, of the South African National Biological Institute were sent to Dr. Csaba Csuzdi, a Hungarian oligochaete specialist in Budapest (one of the things John Measey studies is what caecilians eat).  Several years later, Dr. Csuzdi, using both his own and John’s specimens, published a comprehensive treatment of the earthworms of Sâo Tomé, recognizing 18 species.  Six of these (33%) are endemic to the island, and all six of them belong to the same genus, Dichogaster. As one might predict, all six of the endemics are high elevation species, which suggests they are present in habitats up and out of the reach of the 500 years of agricultural activities. Conversely, all of the widespread non-endemic species appear to be confined to the cultivated lowlands.  All of this suggests that the six endemic earthworms are the result of a species swarm or explosion.  Currently, we know less about the millipede fauna but what we do know suggests they may a species swarm as well.

Dr. Tom Daniel with first ST & P holosaprophyte, Rio Papagaio, Principe [RCD phot. GG IV]

In the May blog, I reported that we had found the first saprophytic plant on Sâo Tomé and Príncipe; Sciaphila ledermanni is a very rare plant indeed, and Dr. Tom Daniel’s publication on the discovery will soon be published. Saprophytes are plants that lack chlorophyll and, in this case, depend on fungi to provide sustenance from dead or decaying matter.  S. ledermannii is otherwise known only from Nigeria and Cameroon, well to the north of our islands; our discovery of this plant along two small rivers on Príncipe is intriguing, in part because of the possibility of the Niger River as its colonization pathway.

Giant Dispersal Raft, with Burrowing Endemics. [art by R. E. Cook, D. Lin phots, GG II] ]

In an much blog I discussed how my colleagues and I hypothesized that the most likely explanation for how the ancestors of the current endemic species colonized the islands was by rafting; our reasoning is based on a combination of the existence of the the two huge rivers that enter the Gulf (the Congo and Niger), the directions of the main oceanic currents in the Gulf (probably unchanged since the Cretaceous) and the fact that surface salinity drops significantly during rainy periods.

The inset in the image of the raft above includes pictures of the endemic burrowing reptiles and amphibian found only on the islands; these species do not swim or climb trees, their presence strengthens our large floating raft hypothesis.   In this 2007 publication, we did not suggest one river or the other as being the likely dispersal pathway to the islands; we just presented the evidence that either was possible.  In the case of our newly found Sciaphila, it would seem that the Niger region might be the most likely source area for this species.

An old friend of mine, Angus Gascoigne, just sent me a series of emails regarding another species with a distribution that is also very suggestive of the region of origin.

Angus Gascoigne, Contador Valley, Sao Tome. [RCD  phot. GGI]

Angus first came to the islands as an English volunteer (VSO), then worked for many years as chief technician for the Voice of America in Sâo Tomé. He currently teaches at the Instituto Superior Politecnico and is a lead biologist for ABS, the new association of Sao Tome biologists. He is a gifted naturalist, has lived on Sâo Tomé for over 20 years and published on everything from snails to Begonias.

Sao Tome Whip Spider. [J. Uyeda phot. GG II]

Angus wrote of finding a whip spider, or amblypygid, in the garden of his home in the town of Trinidade on Sâo Tomé.  To some, whip spiders might be the source of nightmares, looking vaguely like flattened, fast-moving spiders or scorpions. In fact, they are neither; they have their own order within the arachnids, the Amblypygi.  They are non-venomous, usually nocturnal predators on other arthropods . We herpetologists do a lot of work at night, and we have encountered them fairly frequently on rock walls, and bridges on Sâo Tomé.

[Sao Tome Whip Spider, waterfall en route Sao Nicolau. [J. Uyeda phot. GG III]

Angus and his colleague, Hugulay Maia, also of the ABS, have begun a study of these odd creatures, and they inform us there are two species known from the islands so far: Charinus africanus, which is considered an endemic to all three oceanic islands, and Damon tibialis. D. tibialis is currently known only from Sâo Tomé, the Democratic Republic of the Congo, and northern Angola, near the mouth of the Congo River.

Contador Valley Whip Spiders. [D. Lin phot. GG I]

As we must always bear in mind, it is quite possible that D. tibialis was inadvertently brought to the island by the Portuguese, and we cannot know for certain until we can look at their genetics.  But assuming this whip spider got to the island by natural dispersal, its current distribution might represent an example of the Congo as a source path as opposed to the Niger.

Two Possible Distribution Pathways. [C. Pfeiffer construct; RCD GG IV and J. Uyeda GG II phots]

Another possible Niger contribution is the ancestor of Africa’s largest treefrog, Leptopelis palmatus, found only on Príncipe.  It’s distribution and that of its presumed closest relative is pictured below.

Leptopelis palmatus and L. macrotis Distributions. [L. Scheinberg constr.]

I have little doubt that both major rivers have contributed to the founding populations of the marvelous endemic biota of the Republic of Sâo Tomé and Príncipe.

Before signing off, here is an update on our opening “nightmare” image, sent by our friend Polly Anderson, who took the photo.  I just received it as I was concluding this posting.

We met some guys at Pestana hotel on Friday. They work on the big oil rig in the bay, but surprise – its not an oil rig! It’s the same platform, but there are 2 cranes on it instead. Apparently it is resupplying here and waiting for the arrival of its crew. The guys were flown in here and then will go to the rig and on the 10th it is leaving for Luanda. The rig is going to lay pipes underground in the ocean for BP off the coast of Angola. It has nothing to do with São Tomé, it was just hanging out here to get supplies and its crew

The Parting Shot:

An Endemic Bird Nightmare at Nova Moca [R. Ayres phot. GG IV]


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, 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 Loloumb of Monte Pico and Faustino Oliviera, Director of the botanical garden 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, Velma and Michael Schnoll, and Mr. and Mrs. Robert M. Taylor for helping make these expeditions possible. Tax-deductable donations can be made to “CAS-Gulf of Guinea Fund.

The Race: GG IV– The Second Week

OK, first an update on our special “questing beasts”: last weekend after Roberta Ayres, our education officer, finally arrived, we had a traditional Sao Tome dinner at Nova Moca, a former plantation high on the eastern slopes of the mountain. This is where Ricardo Lima lives, our graduate student colleague who first brought my attention to the supposedly endemic Sao Tome shrew and our mystery “charroco,” the fish that does not appear on our fish checklist.

Roberta Ayers at Nova Moca (Ricardo Lima in back). RCD phot. GG IV

Ricardo gave me the preserved shrew he had already caught, a piece of its tail preserved separately, and the tail of a second shrew, removed by machete by one of Ricardo’s assistants (I should add that shrews are mean critters to handle, and some species have a toxic bite)… the rest of the shrew escaped. Again, we are anxious to test the DNA of this species to see if it is in fact a real endemic or was brought to the islands recently (like during the last 500 years of Portuguese comings and goings). We are unsure of what preservative Ricardo used for the material (he doesn’t remember), so we will be uncertain as to the value of what we already have until we get back to the lab. But I have left a couple of liters of 85% with him, so we know we will get DNA from the next shrew that is encountered

Millipede from Bom Sucesso. RCD phot. GG IV

Another questing beast is the millipede. To our knowledge, this group (the myriapods) has not been sampled on the islands, and our colleague at North Carolina State Museum, Dr. Rowland Shelley (a specialist on the group) expressed doubt that there was much present. Sure enough, almost the first log we turned over had a bunch of them underneath, and we also have a sample from 1100 m higher (including the above). Obviously we will search many more localities, both here and on Principe. I must also add that it is wonderful to have an expert colleague willing to examine stuff like this and give us some answers… the Academy has a lot of experts, but we don’t do everything.

The botany guys, Tom and Jim are doing wondrous things:

Pico Cavalio from the north. T. Daniel phot.GG IV

The picture above is of Pico Cavalio, one that must be climbed en route to the ultimate Pico de Sao Tome at 2,000 meters. This one is a brute and our guys have collected the top of it. So far, this is the highest locality that any of our expedition members has ever reached at 1566m. Also, the guys have found a bunch of new trails through really good forest that earlier teams missed, especially around Lagoa Amelia (in earlier blogs)– some of the results have been pretty spectacular.

Begonia macambrarensis, A Sao Tome endemic. T. Daniel phot. GG IV

This yellow begonia is known only from Sao Tome, and Tom and Jim found it on the way up to Pico Cavalio. This is our first collection of this endemic and quite exciting.

Brachystephanus occidentalis 2 more to go! T. Daniel phot.GG IV

Although the begonia is exquisite, Tom got even more excited about the critter above. Tom just published a monograph of the Acanthaceae of Sao Tome and Principe. By the end of GG III, he had found all but three of the endemic species present on the islands – now the list is down to two! And we have tissue samples for comparison with members oft his family on the mainland and other parts of the world.

Jim Shevock is in “seventh heaven;” he is up to 400 collections. Many of you don’t know Jim yet (he is new to the CAS faculty)—Jim is not only tireless, but totally ebullient! Nothing bothers him (and Tom and I have been trying!)

Jim Shevock collecting new trail. T. Daniel phot.GG IV

One of the real pleasures of leading these expeditions is learning how differently we do the same thing—collecting and preparing specimens. Collecting for Jim just looks like scraping away at rocks, or tree trunks and other substrates (of course, you have to know where to look, and that can take many years). And basically that is what he is doing.. However, the way he prepares his collections is really different.

Jim with prepared specimens of Leucobryum RCD phot. GG IV

Jim does not press plants like Tom does—the stuff Jim collects is already pretty flat! Instead, he has special archival paper which has, already printed on each sheet, a table that includes blanks for the number of the specimen as well as for the whole range of possible environmental conditions under which the specimen was collected. He fills out the blanks (duplicates everything on the computer, of course), then folds the specimen into the paper in such a way that each packet stands up while the specimen is drying within. Our room (all 3 of us together) frequently looks like a major regatta on San Francisco Bay.

Regatta in room #10. RCD phot. GG IV

It is probably obvious to anyone reading this that we could not do any of this work without approval from the authorities, especially the Ministry of the Environment. The Director is Arlindo Carvalho, a delightful man. One of the other gentlemen with whom we work is Vitor Bonfim, who is head of Conservation on the islands. I am including a picture of Vitor because his nephew is a FaceBook friend of mine.. a student in Los Angeles!

While the guys are having all this fun, Roberta Ayers and I have been pursuing the education proposal/dream/idea. We have had an amazing series of meetings with all sorts of people here on the islands; it is my intention that whatever we come up with will have the enthusiasm of the citizens at all levels.

Roberta Ayres and Roberta dos Santos. Omali Lodge. RCD phot GG

A very dear old friend of mine is Roberta dos Santos. Roberta works with Ned Seligman in his NGO, STeP UP. In fact, Ned met Roberta when he was running the Peace Corps here, years ago. But more than that, she has been in education her entire life and her degree (in education) is from the US. (Imagine a young girl born and raised on Sao Tome—on the Equator—landing in Buffalo, New York in mid January!) We have contracted with Roberta to assist our CAS Roberta set up meetings, etc. And Roberta knows absolutely everybody. The “Dos Robertas” are a wonderful team. Our CAS Roberta has the perfect personality as well as the educational know-how for what we are trying to accomplish, and I am much impressed. She “gets it,” and is fun.

After some panicky moments yesterday, our flights to Principe, the much older, smaller island, have been confirmed. I must confess I will be glad to get back out into the field. While I do have this dream for raising awareness of the unique biodiversity here, I really am a bush guy at heart, and meetings pale in comparison to stomping around in forests and swamps. CAS Roberta has all she needs to carry on during the week we are gone and is totally comfortable. Tom, Jim and I take off on Sunday and will return a week later, after Roberta has flown home. With fingers crossed, we are off to see if “cobra bobo” exists on Principe.

The Parting Shot:

Dinner at Nova Moc. Ricardo Lima and RCD. T. Daniel phot. GG IV


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, 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 Loloumb of Monte Pico and Faustino Oliviera, Director of the botanical garden 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 and Mr. and Mrs. Robert M. Taylor for helping make these expeditions possible. Our work can be supported by donations to CAS, Gulf of Guinea Fund