Category Archives: flowers

The RACE: SIZE MATTERS!

It is a fundamental tenet of the science of island biogeography that more different species of plants and animals will be found on larger islands than on smaller ones. When we say “larger” in this regard, we really mean surface area. Note that in the graphic illustration below right, both islands have the exactly the same circumference, but the lower island has a mountain in the middle of it which markedly increases any measure of its overall surface area.

island size2

RCD construct.

The greater (and more varied) the surface area, the larger the number of niches for living organisms; hence with time and evolution there will be more living plants on animals on larger islands than smaller (above left). For “niches”, think of “jobs”; every living thing has a three-part job: 1. where it does what it does (spatial niche); when it does what it does (temporal niche) and how it gets its energy (trophic niche). No two living things can overlap on all three and coexist, hence size (area) matters! There are other factors of course, such as geological age and island distance from source, that affect the numbers and characteristics of species found on islands.

Our islands of São Tomé and Príncipe are classic examples of the area/species number relationship. Here are just a few examples:

BEGS

Begonia thomeana. T. Daniel phot. — GG

BUTT

African butterfly. Photo from  ARKives. Google Images

fros

New tree frog (Hyperolius) species from Principe Id. A. Stanbridge phot– GG VII.

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African dragonfly.  ARKive phot. Google Images.

The island area effect is even more convincing when the entire archipelago of four islands is included, from the largest (Bioko) to the smallest (Annobón).

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 Lacewing distribution in Gulf of Guinea. Dong Lin phot. GG I; RCD construct.

As one can see, there is an obvious correlation between island size and the number of lacewings present; however in this case it is also important to note that while Bioko is clearly the largest island, it is also geologically youngest and closest to the mainland, having been attached to the mainland multiple times during the Pleistocene. Such factors can have an important effect on these comparisons. While these correlations prove correct over and over again. However this does not mean that very small islands cannot house wonderful biological surprises, and we are learning that this is true in the Gulf of Guinea.

JB 1

Jockey’s Bonnet. A. Stanbridge phot, GG VII

Above is the Jockey’s Bonnet (or Ilha Caroço) so named for its obvious shape. This large rock is only about 3.5 km off the southeast shore of Príncipe, only 35 hectares in area but perhaps 100 m in height. It was undoubtedly once part of the main island, which readers will recall dates back to the Oligocene Epoch, so it is probably quite old geologically.

JB 2

Jockey’s Bonnet. A. Stanbridge phot. GG VII

Although small, the Jockey’s Bonnet houses at least two very intriguing species. The population of native oil palms (Elaeis guineensis) on the western shores of this tiny island have obviously been there for a long time as they have begun to accumulate change from the parent species on the main Island, but a few km away! While still clearly the same species, the Jockey’s Bonnet palms bear seeds (fruit) that is at least twice the size of the palms on Príncipe and São Tomé.

Elaeis guineensis

Bonnet oil palm seed. RCD phot. CAS botany specimen

For bird lovers, an even more exciting occurrence on the Jockey’s Bonnet is that of the Bonnet Seed-eater, a small brown passerine bird noticeably different from its relatives only a few kilometers away on Príncipe!

bonnet seed eater 2

Bonnet seed-eater. A. Stanbridge phot. GG VII

These unique birds are heavier, have longer, broader bills and shorter wings than their island neighbors and have been shown to be genetically distinct from them. They are extremely common (some 3,500 individuals at last estimate) and live exclusively in the oil palm forest pictured above. They have a specialized diet of palm oil and palm pollen, and it is tempting to speculate that there might be some relationship driving the evolution of the palms and the birds.

Isolation and evolutionary change within a population of birds separated from their nearest relatives by only 3 km may well seem counter-intuitive; after all, don’t birds fly? The answer is yes they do, but they don’t need to, they often don’t, at least not long distances! Flying is energetically expensive; if the habitat is relatively stable, suitable for survival and reproduction, why leave it? In spite of their ability to fly, most bird species tend to remain in specific kinds of habitats and areas. This is called philopatry.

Tnhosa grande

Tinhosa Grande.  A. Stanbridge phot. VII

Far to the south of Príncipe (ca. 20 km.) is a fascinating group of small islands known as the Tinhosas. The largest of these is Tinhosa grande (above) with a surface area of but 20.5 ha.

margins

RCD construct

Geologically the Tinhosas are of great interest because they mark the southernmost limit of the Oligocene Príncipe of over 31 million years ago. As we have noted in earlier blogs, Príncipe was once much, much larger but through millions of years of weathering, largely from the southwest, all that remains are the Tinhosas and Príncipe, along with its other islets. And again, Principe is twice as old as São Tomé.

birds

RCD phots, GG I and II  (right – bridled tern)

The Tinhosas are important rookeries for some sea birds such as Brown and Black Noddys, the Sooty Tern and Brown Booby and are recognized by Birdlife International as Wetlands of International Importance and official Waterfowl Habitat.

Tinhosa Grande is also inhabited by at least two different kinds of lizards, a skink species and a gecko species. These were observed and photographed by members of a recent ornithological expedition but specimens were not collected. Our colleague, Dr. Luis Ceriaco, of the Natural History Museum in Lisbon discovered that some of these skinks had been collected by a Portuguese expedition and deposited in that museum 45 years ago.

CeriacoDr. Luis Ceriaco with Principe giant tree frog..  phot from Facebook.

After analysis, Luis discovered that the Tinhosa Grande specimens represented a new species which he has described as Trachylepis adamastor. It is a very large skink differing from its nearest relatives in size, scales and coloration.

Tinhosas skink 2 (c) Ross Wanless

Tinhosa skink. (Trachylepis). Ross Wanless phot.

Members of the more recent bird expedition reported to Ceriaco that that the population of these skinks seemed very dense, and Ceriaco later speculated that there might be a trophic relationship between the numerous skinks and the nesting birds. Notice above that the skink is feeding on a recently broken egg (this photo appeared in the paper by L. Cericaco). Such relationships are not unknown.

cousin overall

Cousin Island. Google images; bridled tern RCD image, Cousin habitat RCD image; Mabuya wrighti James Warwick image)

Cousin Island of the Seychelles Archipelago in the Indian Ocean exhibits a strikingly similar situation that has been well-studied. This small island of 27 hectares supports enormous populations of two species of skinks: Mabuya wrighti, which is large, and Mabuya seychellensis, which is smaller. Studies revealed that in 1979 there were approximately 1,713 individual skinks per hectare, and that these were supported directly by nesting terns (60,000 pairs of Lesser Noddy terns alone) through broken eggs, feces and dropped fish. Such a situation may well exist on Tinhosa Grande.

The Tinhosa gecko remains a mystery. We have no examples of it and cannot examine its morphology or molecular relationships.

tin gecko2tin gecko

Tinhosa gecko. (Hemidactylus sp.) Photos by Nuno Barros, courtesy, Birdlife Int.

The photos are not of sufficient quality to determine whether this gecko is related to one of the unique island species (H. principensis, H. greeffii) or is a more widespread species.

We are preparing for GG IX in September. More anon.

The parting shot:

parting shot

A 4th grade Sao Tomean student with our biodiversity playing cards. A. Stanbridge phot. GG VII

 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: GG VII—We Reunite and Part Again

After two hectic weeks of education activities on São Tomé, Rayna Bell (Cornell University) arrived and the four of us joined the botanists, Tom Daniel, Jim Shevock, Miko Nadel, Tamas Szuts (our spider guy) and Andrew Stanbridge (our photographer) on Príncipe.   I  have asked Andrew, a veteran of three Gulf of Guinea expeditions, to illustrate some of what transpired while the group was divided.

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Our botany team, day one on Principe: Jim Shevock, Tom Daniel and Miko Nadel.

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Botany team en route to climb the mesa. Back left in the yellow hat is our guide Baloo.

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Jim on the “trail” to the mesa.

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Male Leptopelis palmatus found on the trail to the mesa. The females are the largest tree frogs in Africa.

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Tom discovers Principina, a unique sedge.

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Miko on top of the mesa

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Jim and Tom collecting specimens along the route to Roça Sundy.

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First Academy visit to the offshore island “Jockey’s Bonnet”.

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Bonnet seedeater, unique to the small island of “Jockey’s Bonnet”.

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Tom carrying specimens upriver.

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Tamas and “Bobby” Bronkhurst pooting spiders on Jockey’s Bonnet.   Here is the parting shot.

parting

  All images 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-V 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 VII has been 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 continued support of the Omali Lodge (São Tomé) and Bom Bom Island (Príncipe) for both logistics and lodging and especially for sponsoring part our education efforts for GG VII. Our expeditions can be supported by tax-deductable donations to “California Academy of Sciences Gulf of Guinea Fund”

The Race: Endemicity and the Gulf of Guinea VII Expedition (I. The Scientists)

Readers may recall that last March, prior to GG VI, I gave several lectures in Portugal on Gulf of Guinea island biodiversity. The last was an international colloquium on São Tomé and Príncipe held at the University in Lisbon. There I met a number of the participants, among whom were old friends and a delightful entomologist named Dr. Luis Mendes; Luis and I remained in contact, and he has just published and sent me the most up-to date survey of the butterfly fauna of the islands butterfly fauna.

BUTTERFLIES

Photo by Luis Mendes

As we have learned to expect, the endemicity (uniqueness) level is high. Luis and his colleague, Bivar de Sousa, report 111 species present on both islands, 29 of which are found nowhere else in the world. Thus, fully a quarter of the butterflies (26%) are endemics. This is further testimony to the great age of these islands, as we know that genetic change (evolution) occurs with isolation and time. Last month, another paper appeared by Loureiro and Pontes confirming the endemic status of a species of dragonfly, Trithemis nigra found only on Príncipe but not seen for many years.

best Trithemis_nigra_PI_NSL_0213

Photo of Trithemis nigra byNuno Loureiro 

The image below is a summation of our current knowledge of  some of the insect endemicity on the two islands; much of the data upon which this summation is based are very old, and so much more work needs to be done.

INSECTS

photo: www images:  CAS construct.

We are getting ready for GG VII (April-May), and below is our new logo for the expedition; note that the famous Cobra bobo, a legless amphibian found only on São Tomé has been joined by an endemic Príncipe snake, also called Cobra bobo but entirely unrelated. (The cartoons of both animals were made by my graduate student, Dashiell Harwood, and the layout was by a member of our Biodiversity Education Team, Michael Murakami.

logo

GG VII (2013) logo.

Jimmy

James Shevock of CAS; photo A. Stanbridge- GG VI

Jim Shevock, a world-class bryologist, will be joining us for the third time. As you can see from the data above, he has already greatly increased our knowledge of mosses and their relatives on the islands, and there are still many species to be found. For example, during GG VI last year, Jim returned to the same locality along the Rio Papagaio in Príncipe that he had collected during GG V; in GG VI and found many plants he did not find the first time, including 10 of them new to the country! Jim has worked a lot in Asia and his nickname on Taiwan is “Little Bear.”

Rayna

Rayna Bell at Caxuiera, Sao Tome. A.  Stanbridge phot – GG VI

Rayna Bell is a graduate student from Cornell University. During GG VI she studied possible hybridization between the two endemic São Tomé treefrog species Hyperolius thomensis and H. molleri and currently has a paper in press on her work with us last year. This year we will try to find the elusive tadpole (larva) of the Príncipe giant treefrog which remains undescribed. Leptopelis palmatus is the largest treefrog in Africa.  Speaking of herpetology, to date our CAS island specimens and tissues have been used in 33 scientific publications, internationally!

Tom

Dr. Tom Daniel, Lagoa Amelia, Sao Tome.  RCD phot, GG III

Dr. Tom Daniel is a veteran of GG III and GG IV. Our senior botanist, he is a specialist on the flower family Acanthaceae (shrimp plants); in the picture above, he is standing in Lagoa Amelia next to Heteradelphia, a genus we think is endemic to São Tomé. He has done a lot of work on ferns and other Gulf of Guinea plant groups as well.

Tamas final

Dr. Tamas Szuts with some of his critters – Tszuts photos

Dr. Tamas Szuts is an expert on jumping spiders of the family Salticidae. He was a post-doctoral fellow here at the Academy under Dr. Charles Griswold (GG I) and will be joining the team for the first time. He is now on the faculty of the University of West Hungary. Salticids are about the only spider  group I think are kind of cute, face to face!

Miko

Miko Nadel, Sao Tome.  A. Stanbridge photo. GG VI

Miko Nadel is a graduate student at San Francisco State University under Prof. Dennis Desjardin (GG II, GG III). After making a comprehensive lichen collection during GG VI, he has decided to focus his research on the lichen genus Usnea; these are the hanging, pendulous lichens known in the US as “old man’s beard.”

droo better

Andrew Stanbridge at Laguna Azul, Sao Tome.  A. Stanbridge photo. GG VI

We will once again be documented by the world’s largest photographer, Andrew Stanbridge, veteran of GG V and GG VI. Andrew was one of those who ascended the Pico do São Tomé last year (see last April blog). His obvious photographic skills are only part of what he brings to our expeditions.

bob-1

Dr. Bob Drewes with Regional President of Principe, Hon. Jose Cassandra.  A. Stanbridge phot.  GG VII

I will be leading the trip as usual and will attempt to answer the ongoing question: do I have to wear a tie to see President Jose, or do I not have to wear a tie? .. Never quite seem to get it right.

The second part of the blog will be focused on the education team and our plans for Gulf of Guinea VII

Here’s the Parting Shot:

incredible Principe

Incredible Principe Island. A. Stanbridge phot. GG VI

 

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 Tome http://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-V 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 VII has been 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 continued support of the Omali Lodge (São Tomé) and Bom Bom Island (Príncipe) for both logistics and lodging and especially for sponsoring part our education efforts for GG VII.
Our expeditions can be supported by tax-deductable donations to “California Academy of Sciences Gulf of Guinea Fund”.

 

 

 

 

 

The Race: Island Biologists in Training

Jens Vindum, Senior Collections Manager, Department of Herpetology. (phot D. Lin-GG I)

I need to add and addendum to last month’s blog, “Why We collect Specimens.” Our Senior Collections Manager, Jens Vindum (GG I, GG II) has just informed me that since 2003, there have been 33 international scientific papers published on our Gulf of Guinea reptile and amphibian specimens and/or tissue samples from them!

Clearly, the scientific world is beginning to hear about Sâo Tomé and Príncipe! At this point, I do not know how much of our material from other disciplines has been used but certainly our samples are in labs all over the world.

We have been extremely fortunate to have been able to bring a series of our graduate students with us on a number of our expeditions.  Not only have most flourished academically and many have published on their island projects, they represent a cadre of new young scientists who have an understanding of the uniqueness of the islands and the people who live on them.  All have interacted closely with local island citizens and as a result, function as young biology ambassadors for these fabulous islands.  Overall, the islands are still very poorly known to the outside world, but we are getting there!  Here are our young colleagues:


Lindsay Wilson on Bioko Island with bush viper.  RCD phot – 1998

Lindsay Wilson was a participant on our 1998 expedition to Bioko, the first island in the Gulf of Guinea chain. She completed her MSc on African treefrogs of the genus Hyperolius at San Francisco State University with highest honors.


Joel Ledford on Sao Tome.  D. Lin phot- GG I

Joel Ledford joined Gulf of Guinea I as the graduate student of Dr. Charles Griswold. He completed his MSc at San Francisco State and then his PhD in spider systematics at the University of California, Berkeley.


D. Lin phot – GG I

Also on GG I was Ricka Stoelting, my graduate student. She completed her MSC on the endemic caecilian of Sao Tome (she is holding one, above) and is currently a PhD candidate at the University of Wisconsin.  She is also working on the publication of her MSc work at San Francisco State (SFSU).


B. Van Syoc photo – GG III

Dana Carrison-Stone was a participant of the marine expedition, GG III as the graduate student of Dr. Bob Van Syoc.  Dana discovered two new species of barnacles from the islands and they are part of her MSc which she completed last year at SFSU.


D. Lin phot – GG II

Josef Uyeda was on GG II and again GG on III as an undergraduate at Willamette University and one of my Summer Systematics interns.  During his island work, he discovered and described a new species of frog from Sâo Tomé. As I write, he is defending his doctoral thesis (tomorrow!) at Oregon State University. Flash!! Josef finished his PhD today! (Oct 5)

 

                                                                                                             unknown phot.

Mac Campbell, also a Willamette undergrad, joined GG II as an assistant to our ichthyologist, Dr. Tomio Iwamoto.  He has since completed his MSc at University of Alaska, Fairbanks and is currently a PhD candidate in fish systematic at the same institution.


Weckerphoto – GG III

Rebecca Wenk joined GG III as the grad student of Dr. Tom Daniel one of our senior botanists.  Rebecca’s work resulted in her successful completion of her MSc at SFSU and also an excellent scientific publication on plants of the family Acanthaceae.  Tragically, Rebecca died of a serious illness last year.


A. Stanbridge phot – GG IV

Miko Nadel is a graduate student at San Francisco State, studying under Dr. Dennis Desjardin, the mycologist on GG II and GG III). Miko was a participant on GG VI doing the first comprehensive survey of lichens on the island.

A. Stanbridge phot. GG VI

Rayna Bell also joined us on GG VI, studying color variation in African treefrogs. Rayna is a PhD candidate at Cornell University.

The people above were or still are graduate students who have actually worked on the islands with us.  But they are not the only young academics studying our Gulf of Guinea Island material.  Here at the California Academy of Sciences we have a program known as the Summer Systematics Institute (SSI). This program is funded by the National Science Foundation, and undergraduate students can apply to work on scientific projects for the summer under the mentorship of a CAS faculty member. Here are those that have worked on Gulf of Guinea specimens. I have not included students who started as undergrad SSI interns and later became our grad students (Lindsay Wilson, Josef Uyeda and Ricka Stoelting).


unknown phot

Katie Marshall was an Occidental College undergrad and my SSI intern in 2006.  Katie studied the genetics of the Oceanic treefrog, Hyperolius molleri, the only Gulf of Guinea endemic frog that occurs on both islands.  Katie is currently a PhD candidate at the University of Washington, studying the genomics of marine bacteria.

RCD phot.

Lisette Arellano was an undergrad at the University of California, Santa Barbara when she joined us as my SSI intern in 2009. Lisette examined the morphology and genetics of cobra jita snakes (Lamprophis), long thought to be the same species on Sâo Tomé and Príncipe.  Lisette showed that in fact the two island populations are genetically quite different, also recognizable by color pattern as distinct.  Although we know each island is a different species, we have been unable to publish new names because the relationships of the same group on mainland Africa are still very unclear. Lisette is currently a PhD candidate in Biology at the University of Colorado.


RCD phot – 2010

One of the last vertebrates one would predict to be native to an oceanic island is a shrew, largely due to physiological constraints. During the SSI summer of 2010, Eden Maloney’s DNA work showed that the Sâo Tomé shrew, Crocidura thomensis, did arrive on the island naturally, probably many thousands of years ago and is a true endemic species. Its nearest relative is a different species found in eastern South Africa.  Eden has just graduated from the University of California, Los Angeles and is applying to graduate schools.  We are working on publishing her work.

unkown phot.

Lizzie Miller of the University of California, San Diego was my most recent SSI intern (2010). Lizzie has graduated and is now in graduate school at UCSD studying fish systematics.  Readers will already know from this blog that Lizzie discovered and described a new species of gecko from Príncipe, Hemidactylus principensis.

Lauren in Nigeria. D Blackburn phot – 2012.

Lauren Scheinberg is also a grad student at San Francisco State University. Although never an SSI intern nor has she been with us to the islands, she was my lab assistant on a long-term physiology project and now works as a curatorial assistant in our department.  She has become involved in a rather complicated taxonomic problem with the island skinks of the genus Afroablepharis. Like Lisette’s snakes, we know from the work of colleagues in Madeira and Portugal that the skinks are different species on Sâo Tomé and Príncipe.  Unfortunately, material we loaned them that formed part of the basis of this hypothesis was somehow lost in transit.  Lauren has analyzed our remaining material but collating the information generated by different labs can be extremely difficult.  But we are working on it.

Plans are already afoot for GG VII next year.

Here’s the parting shot:

Joy on the way to Rolas, Sao Tome.. B. Simison phot. – GG VI

PARTNERS
We gratefully acknowledge the support of the G. Lindsay Field Research Fund, Hagey Research Venture Fund of the California Academy of Sciences, (GG I, II), the Société de Conservation et Développement (SCD) and Africa’s Eden for logistics, ground transportation and lodging (GG III-V), STePUP of Sao Tome http://www.stepup.st/, Arlindo de Ceita Carvalho, Director General, and Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for permission to collect and export specimens for study. Special thanks for the generosity of private individuals who made the GG III-V 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 HBD of Bom Bom 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. Abell, Prof. & Mrs. Evan C. Evans III, John and Judy Sears, John S. Livermore and Elton Welke.
Our expeditions can be supported by tax-deductable donations to  “California Academy of Sciences Gulf of Guinea Fund”

The Race: Why We Collect Specimens!

Summer has been extremely busy.   Our irrepressible bryologist, Jim Shevock, comes into my lab almost weekly with new moss discoveries from GG VI.  He says a new paper on Fissidens (the largest moss genus on Sâo Tomé and Príncipe) is almost finished and will be submitted for publication as soon as he and his colleagues (from the US, the Netherlands and Lisbon) complete a key to identification of the species. Recall that Jim nearly doubled the number of collections he made during GG IV… He thought he had seen everything! The mesa on Príncipe will be a primary target for our botanists on GG VII, next year (see below).

 

Phrynobatrachus leveleve – RCD phot, GG VI

A nice surprise from GG VI was that we finally got some nice, un-posed  photographs of the Sâo Tomé puddle frog, Phrynobatrachus leveleve. Readers may recall that we described this new endemic species back in 2007 following GG II (leve leve means “take it easy” in the local language).  Obviously the way to get good shots of these critters is at night!

Ptychadena newtoni.  A. Stanbridge phot – GG VI

Another good find on a different night was Newton’s rocket frog (Ptychadena newtoni) at a new locality, Caxueira.

The creek at Caxueira.    A. Stanbridge phot. – GG VI

In earlier days I was concerned that this species, endemic to Sâo Tomé, was on the wane due to human development, but it appears to be more widely distributed than we thought (see also Feb 2011 blog). Caxueira is not far from the city center.

Why do we collect plant and animal specimens? Why do we bring them home euthanized and preserved (or in the case of plants, pressed and dried), and why do we organize and store them for posterity?  The easy answer is that we need to find out what they are, to identify them and describe them so we can communicate about them.  We certainly cannot conserve or preserve or even talk about species if we do not know they exist. This is particularly important in the tropics where so many different species have evolved, and especially in areas like Sâo Tomé and Príncipe that have never been fully explored by biologists.  An added note is that for a biologist to know that a species is new and undescribed, he has to know all the related species that it isn’t and then demonstrate it!

It is a fact that a lot of things in the tropics that look alike are not at all related; conversely, some critters that look radically different are, in fact, just variants of the same species.

The botanists of course confront similar questions. Below are two species of the genus Impatiens.

(l) T. Daniel phot – GG IV; (r) M. Nadel phot – GG VI

Both species are high elevation forms described a long time ago: I. manteroana is thought to be endemic to Príncipe, while I. thomensis is known only from Sâo Tomé. But are they really different species? And if so, are they each other’s closest relatives?  We do not yet have material of the former, but this is a question we can answer next year through DNA analysis. The specimen on the right was photographed high on the Príncipe mesa, which is one of the reasons it is a target area for next year.

Below is an island example of two species that look very much alike but are definitely not the same:

D. Lin phots: GG I, GG II]

These are photographs of small leaf-litter skinks of the genus Afroablepharus. The specimen above was collected on Sâo Tomé during GG I and the one below came from Príncipe (GG II).  While they look identical, they are actually two different species as shown by colleagues of ours who were working on the molecular level: extracting DNA from small bits of tissue (probably tail tips) the two species were shown to be genetically quite different.  The one from Príncipe was described over 160 years ago (A. africanus), while the one above, from Sâo Tomé, remains unnamed. This is most frustrating as even though we know they are separate species, we cannot describe the new species yet because the Sâo Tomé animals from which the DNA was analyzed were not collected.  It is a complicated situation that both groups of workers together are trying to resolve at this time.

Another example can be found in the island geckos about which I have written before.

From public presentation by E Miller. CAS Big Kahuna phot (same specimens from above and below).

For over one hundred years, the geckos from both islands that lack thumb nails were considered to be the same species, Hemidactylus greeffi, originally described from Sao Tome.  Our same colleagues noted that the two were genetically different but again failed to take whole samples and so could not describe the Príncipe species as new.  It was not until we closely examined specimens in our Academy collections from both islands that we found many morphological differences between the two, which strongly supported the genetic evidence of our colleagues.  The animal on the right is now  known as Hemidactylus principensis, yet another island endemic. As luck would have it, the paper was published while we were on Principe!

The smaller specimen on the right in both views is also what is known as the holotype; i.e., it is the single animal that is described in minute detail that becomes the “name bearer”.  All geckos collected from the islands and identified as H. principensis will be based on the description of this particular specimen; holotypes are the most important specimens in any collection.

In our collection, which is probably the fifth largest in the world, all holotypes are housed separately and identified by a blue ribbon.

Part of Herpetology collections rooms; holotypes above right, paratypes below right. RCD phots.

Another question often asked of museum scientists is “why do you have to collect so many?”  The answer is that species vary; no two members of the same vertebrate species are identical.  This is why we include additional specimens in a species description.  While the holotype or “name bearer” is usually a single animal in a standard description, other members of the same purported species, hopefully from the same place, are also described in some detail in order to account for individual variation.  These are usually designated as paratypes; in the Academy collections, they are always designated by red ribbons [above] and are the second-most important.

Yet another frequently asked question is, “do you have to kill the specimens?”  The answer lies in the fact that not all characters (similarities and differences) are observable from the outside.  With animal groups like frogs, one has to look deeper, and this is impossible with living specimens. Below is a collage of some of the sorts of characters I had to examine in determining the relationships between members of African tree frogs of the family Hyperoliidae— found in Africa, the Seychelles and Madagascar.

All RCD phots.

Notice that the x-ray in the lower right hand corner revealed to us that the two geckos mentioned above not only lack thumb nails, they lack the entire terminal bone of the thumb! (the new species, Hemidactylus principensis, is on the left). So far as we know, they are the only two members of the genus Hemidactylus, (90+ species) that exhibit this characteristic.  This might suggest they are each other’s closest relatives, but we are in the process of determining that by further DNA analysis that includes other closely related species.

During GG VI we did another kind of collecting:

Rayna Bell, Cornell University. A. Stanbridge phot – GG VI

Notice that in her left hand, Rayna Bell is holding an adult Sao Tome giant treefrog (Hyperolius thomensis), while in her right she has a cotton swab.  She swabbed the skin of each frog she collected a number of times in a number of places in order to detect the presence of chytrid fungus. The swab will also detect the actual infection load if the fungus is present.  This is the first attempt at detecting the fungus on the islands of Sâo Tomé and Príncipe, and we do not yet have results. It is certainly present in other areas of Africa. Batrachochytridium dendrobatidis (Bd for short) is a fungus that has been implicated in the mass die-off of populations of frogs in many parts of the world.  Frog skin is a living membrane through which gasses and water can freely pass; while the mechanism is not well-known, the fungus seems to totally disrupt these functions causing the demise of the infected frog.

 

Cross section of Bd infected frog skin.  (A) are sporangia with zoospores visible. (B) tube through which zoospores are released to the environment. Phot courtesy of A. Pessier, U. Illinois

Another real value to collections is the fact that past history can be discovered through our specimens. It turns out that Bd can also be detected by swabbing alcohol preserved specimens regardless of age, although the resulting data are not quite so informative as samples from living material.  Below is Dr. Dave Blackburn’s “chytrid crew” (mostly undergrad and graduate students) swabbing specimens collected from the Impenetrable Forest of Uganda many years ago.  Dave is our new curator in herpetology and a real expert on Bd.

Dave Blackburn’s “chytrid crew”.  D. Blackburn phot.

Every trip to these small amazing islands yields new discoveries. We are planning our next expedition for 2013 and excited at the prospect of the new stuff we will find.

Here’s the parting shot.

Autonomy Day in Principe, 2012 A. Stanbridge phot, GG VI

PARTNERS

We gratefully acknowledge the support of the G. Lindsay Field Research Fund, Hagey Research Venture Fund of the California Academy of Sciences, (GG I, II), the Société de Conservation et Développement (SCD) and Africa’s Eden for logistics, ground transportation and lodging (GG III-V), STePUP of Sao Tome http://www.stepup.st/, Arlindo de Ceita Carvalho, Director General, and Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for permission to collect and export specimens for study. Special thanks for the generosity of private individuals who made the GG III-V 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 HBD of Bom Bom 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. Abell, Prof. & Mrs. Evan C. Evans III, John and Judy Sears, John S. Livermore and Elton Welke.

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

 

 

The Race: Gulf of Guinea VI Part II (Sharing the Wealth)

eddie-herbst-principe

Principe Island from the northeast. This island is at least 31 million years old [Phot. Eddie Herbst]

Some years ago the Gulf of Guinea Project “morphed” from a pure multidisciplinary research focus to include an additional and parallel effort to share our science with the local people and non-scientists everywhere. My first couple of visits to São Tomé and Príncipe followed over thirty years of fieldwork on the African mainland, essentially doing science that is read and used by other scientists; this had been wonderfully exciting, rewarding and fun (sometimes scary).But my exposure to the islands of São Tomé and Príncipe changed my outlook and to some degree, the direction of my work in a fundamental way.

Here is a tiny two-island nation absolutely unique and rich biologically, yet still poorly known to the world of science. At the same time there is a looming threat to the environment with the recent discovery of off-shore oil, and the real danger that the world might lose this biological richness before it is even discovered and described! Moreover, the delightful citizens of these islands have, by and large, no idea how rich and special their biodiversity heritage is. Perhaps if we could make the citizens aware of what they have that is unique, found nowhere else in the world, they might be in a better position to make informed decisions as change occurs in the future.

Here, I realized, is an opportunity to help an entire nation prepare for change through awareness of the unique nature of their environment.But how?

blog-image

The sharing of knowledge is fundamental to my discipline, and I have always brought or sent copies of our published scientific results to the governments and appropriate institutions of the African countries in which I have worked, but it was not until several years into our islands work that it occurred to me that we should be including a Portuguese abstract in each of our publications (above left). The abstracts at least make our work understandable to Portuguese scholars who read scientific journals,but on the islands, it was only the various ministries and specialists who even received these articles, and they are technical in nature (we have published 18 so far).In 2008, I began to write this monthly blog (above right). While it is written for a popular audience and hopefully helps bring world attention to the biological uniqueness of the islands, it is still only available to English speakers with access to the worldwide web.

Visitors to the California Academy of Sciences are aware of our work in the islands as we occasionally have small semi-permanent exhibits on our island work on the public floor (see below)

ex

I should mention that the Academy is nearly 160 years old, and we receive over one and a half million visitors per year.We have frequent after-hour public and fundraising events, and whenever possible we have a Gulf of Guinea Islands display which give those of us who are involved an opportunity to describe our research to our public in person.

exped-nite

Velma Schnoll at Discovery Evening.March 2, 2012 (Phot. RCD)

While these events help make our local visitors aware of our island work, they obviously have only indirect impact to our island friends.

During expeditions in the past few years we have been interviewed by local media (radio and television) and have been asked to give lectures on biodiversity at a number of schools and institutions, especially at the Instituto Superior Politecnico, thanks to Dra. Alizira Rodriguez, and also at a biodiversity conference under the auspices of Regional President Tosé Cassandra of Príncipe.

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Lecture at Instituto Politecnico Superior in Sao Tome (Phot. A. Stanbridge, GG V)

school-lecture

Local high school lecture (Phot A. Stanbridge, GG V)

These have been excellent opportunities to communicate directly to advanced students and conservation workers (through a translator), but the information still does not get down to the fishermen, the kids, or the people in the market places.

Sometime in 2010, before GG IV, it occurred to me that we might be able to reach the local populations visually.One of the things we have that nobody else has is outstanding images, not just of pretty beaches and Câo Grande, but of the unique living plants and animals themselves!And we know what they are, and sometimes where they came from.So, I put together a series of powerpoint mockups of colorful biodiversity posters on my laptop, and during GG IV Ishowed them to everyone who would look; they all seemed to like the colorful montages.

robertas

Roberta Ayres (CAS) and Roberta dos Santos (STeP UP). [GG IV, from V. Schnoll presentation, 2011]

During GG IV, one of the expedition members was Roberta Ayres, MSc., an Academy educator who runs our nature center (see earlier blogs).Roberta’s mission was to assess the level of knowledge of biodiversity in the islands’ school system, assess the likely impact of the posters, and to discover what else we might do to raise biodiversity awareness through the schools.I have written a number of blogs about how the posters were ultimately produced, thanks largely to the efforts of Velma Schnoll, Docent Coordinator and Jim Boyer of our CAS Docent Council (see below) and with funding from STeP UP.

posters-only

Posters, GG V. from V. Schnoll presentation, 2011.

Gulf of Guinea Expedition V was a largely educational mission dedicated to the distribution on both islands (see earlier blogs) of the 200 posters we produced, and this was accomplished by Mrs Schnoll, Andrew Stanbridge (our photographer), and I, along with a host of local friends, including Marnie Saidi of Príncipe and Antonio Fernando of São Tomé. A very central figure in all of our endeavors, both scientific and educational since the very first expedition in 2001 has been Quintino Quade of STeP UP.Readers will know that he appears in virtually every blog since the first one.

dist-collage

Poster distribution. [all photos A. Stanbridge-GG V; from V. Schnoll presentation, 2011]

At the same, we continued to pursue information on our ultimate goal which is the creation of a Gulf of Guinea Biodiversity Center, a place on the islands where the citizens can access all of the information being gathered about the environment, and which can serve as a clearing house for all science and natural history research on the islands. Over the years, we have shared this idea with many citizens and foreign researchers on São Tomé and Príncipe.

In Part I of this blog, I described the scientific goals of GG VI (which begins next month) and introduced the investigators who will be on the expedition.The education component (Part II) is meant to build upon the efforts of GG V, and two educators will be coming along as well: Roberta Ayres (GG IV) and Velma Schnoll (GG V).

Like the overall project, our biodiversity education efforts have morphed into a team with Mrs. Velma Schnoll as Biodiversity Education Project Manager.After much debate (including the possibility of an animated cartoon), the team decided to produce 2,000 coloring books for young elementary school students, featuring the same endemic species that appear on the posters of last year. We have selected four primary schools on São Tomé and, of course, the one in Santo Antonio, Príncipe as our trial sites.

cb-team-2

The Biodiversity Education Team (l-r): T. Daniel (science advisor), V. Schnoll (project manager), J. Boyer (chief illustrator and production), C. Schneider and S. V. Edgerton (fine art), R. Ayers (text and translation) and M. Murakami (graphics and production). Absent : L. & C. Rocha (translation) and E. McElhinny (cartography). [Phot RCD]

cb-team

The team at work [Phot V. Schnoll]

v-proj-mgr

Velma Schnoll, project manager, with initial page layout [phot RCD]

The coloring books are being printed as I write;

solstice

Thanks to Mike Murakami’s friend, Richard Engle, proprietor of Solstice Press, Oakland California, we got a very favorable discount on the printing costs. So, here is what they will look like:

stp-front-cover-faded1

The front cover artwork is by Sean Vidal-Edgerton. Sean and Corlis Schneider (back cover art) were both in a biological illustration program at the University of California at Santa Cruz, and California State University, Monterey Bay. In 2011, this group produced a wonderful on-line account of our São Tomé and Príncipe biodiversity research: http://sciencenotes.ucsc.edu/2011/pages/eden/eden.html We were extremely fortunate that they joined us.

ibc

The inside back and front covers have color images of the same living plants and animals that are illustrated in the cartoons.

contents-overview

This is a spread of the full contents of the book; except for the first two pages, the cartoons will not necessarily be in this order. The two game pages will be in the middle of the book.

Here are a couple of Jim Boyer’s fabulous cartoons as they will appear as full pages:

sele

The Giant Begonia and Newton’s sunbird, both endemic to Sao Tome.

puddle-frog

The Principe puddle frog found only on that island.


Back cover Sao Tome e Principe coloring book

Again, the back cover artwork is by the talented artist, Corlis Schneider.As the logos indicate, much of the production costs of this project have been provided by a Goldman Fund donation to STeP UP, one of our main partners on the islands.Africa’s Eden is already well known to readers of this blog; the rest of us are volunteers.

So, Part II of Gulf of Guinea Islands Expedition VI is the distribution of the coloring books. Moreover, Roberta Ayres and Velma Schnoll have produced a teacher’s guide incorporating both the books and the posters with island evolutionary principles, and they hope to conduct a workshop for teachers in São Tomé and later on Príncipe.

Now, all we have to do is get 2,000 of these books to the islands by hand, and somehow procure enough colored pencils (crayons do not work well on the Equator) over there for the kids.

As usual I will post from the islands.

Here’s the parting shot:

atopochlis-exerata1


Atopochlis exerata, one of the many unique snails on the islands. Photo byM Morais, courtesy of A. Gascoigne]

PARTNERS

We gratefully acknowledge the support of the G. Lindsay Field Research Fund Hagey Research Venture Fund of the California Academy of Sciences, (GG I, II), the Société de Conservation et Développement (SCD) and Africa’s Eden for logistics, ground transportation and lodging (GG III-V), STePUP of Sao Tome http://www.stepup.st/, Arlindo de Ceita Carvalho, Director General, and Victor Bomfim, and Salvador Sousa Pontes of the Ministry of Environment, Republic of São Tomé and Príncipe for permission to collectexport specimens for study. Special thanks for the generosity of private individuals who have made the GG III-V 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 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. Abell, John and Judy Sears, John S. Livermore and Elton Welke.

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

The Race: News from the Moss People

This month I had hoped to describe part of our island educational project but things have been too busy here at the Academy; our Fall calendars are always full as it is the beginning of the academic year in the US. We have also hit some snags on our planned São Tomé- Príncipe biodiversity posters. We have the funding (thanks to donors to STeP UP) and the imagery from our expeditions; in fact we have everything except a designer. In the March blog, I included two of the mockups which were among those we took to the islands; everyone we showed them to liked them very much. It turns out though, that producing 200 high resolution posters of large size is not an easy undertaking, but we are working on it. Below is one of my favorites:


“Only In Sao Tome!!” D. Lin phot – GGI

There is much to relate on many fronts including news of the millipedes which are currently with an expert in Belgium, the identity of the Príncipe shrew, Lisette’s work on cobra jita, etc., all which I must write about later.

If you follow this account, you have already met Jim Shevock, one of the foremost bryophyte workers anywhere, and you will recall that during GG IV, he made a huge collection of liverworts, hornworts and mosses.


[Jim Shevock with Afrocarpus mannii, Morro Provaz. T. Daniel phot – GG IV

Jim’s island collection represents an enormous amount of work and will take a long time to fully analyze, but here is a progress report from him:

“Of the 700 bryophyte collections made during GGIV, nearly 275 of them represent liverwort and hornwort specimens. The specimens obtained from these two taxonomic groups have now been named to species by two world experts residing in Dresden and Budapest who specialize in African liverworts. The results are very impressive. Prior to GGIV, the published liverwort and hornwort checklist contained 85 species for São Tomé and 33 species for Príncipe (20 species are shared between the two islands). We have now added 26 liverworts as “new” for São Tomé and 19 species as “new” for Príncipe.


Orthostichella sp. (hanging). Lagoa Amelia. J. Shevock phot – GG IV.

The next phase is to complete the identification process for the mosses collected during GGIV. We anticipate the data on the mosses will be similarly impressive. The currently published moss checklist for São Tomé contains 76 species and only 14 mosses are reported in the literature for Príncipe. We anticipate the moss species list to expand markedly. We already have a moss family to report as new not only to the Gulf of Guinea but to West Africa (the Symphyodontaceae); other African records are known only from Malawi, Mozambique, Madagascar and Reunion Id. We have at least one species of moss new to science , and there will probably be more. All of the material obtained within the moss families Fissidentaceae and Neckeraceae are now identified to species, and in both cases, new taxa are documented for the country and from both islands. Some species will also be reported for the first time as occurring within West Africa. Based on the data obtained so far, the actual bryoflora of São Tomé and Príncipe is much richer than initially projected. Conducting these expeditions is but the first step toward the discovery of the biota of São Tomé and Príncipe.

Moss Fissidens ovatus and Begonia annobonensis, SW Príncipe. RCD phot – GG IV.

I posted a similar photo in the March blog when we thought the tiny flowering plant above might be an undescribed species of Begonia; we subsequently learned that this species is already known to science, but it is still possible that this southwestern Príncipe population may represent the smallest Begonia plant in the world. And, now we know that the rest of the rock is covered with the widespread tropical moss species, Fissidens ovatus.

The parting shot:


Santa Catarina buddies. 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) 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 and Velma and Michael Schnoll 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: http://www.nadiplochilo.com/milli.html 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]

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 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 – Return to Paradise

Having just returned from Ethiopia, I am now “gearing up” for GG IV.  If all goes well, we will be returning to the islands on the 19th of February for a month, thanks especially to continuing logistical support from our partner, Africa’s Eden (SCD), and the generosity of friends (see Partners below).

GG IV will be one of the smaller expeditions (four of us), due in part to financial constraints, but also because I have a particular, non-exploration focus in mind.  But first the GG IV players:

Dr. Tom Daniel, is returning with us; as I mentioned in the last blog, he has just published a major paper on the island shrimp plants (his specialty) and along with more botanical exploring will be doing some technical pollination studies this time.

Dr. Tom Daniel on Sao Tome. RCD phot, GG III

Among our plans is a survey of the top of Pico do São Tomé which is at about 2000 m.  None our previous expedition members has ever sampled the Pico so everything will be of interest, but our one of our special goals is to collect examples of Afrocarpus mannii, which is endemic to this mountain.

Afrocarpus mannii  WWW. photo

This tree is a member of the yellow wood family (Podocarpaceae), and it is thought that all of its nearest relatives are found thousands of kilometers away in the East African highlands.

Distribution of Afrocarpus relatives. RCD construct.

This strange distribution pattern is showing up rather frequently in the various sorts of organisms we study (for instance, my frogs and reptiles) so we are always interested in testing these relationships using DNA technology; i.e., if these species are really closest relatives, what are they doing thousands of kilometers apart?

Dr. Shevock in Yunnan, China. Phot. D. Long – 2007

This is Dr. Jim Shevock, who recently joined the Academy faculty.  Jim is one of the world’s foremost authorities on mosses. His latest book came out only a few weeks ago.

California Mosses. 2009. Micro-optics, New Zealand

Jim will be conducting the first comprehensive moss survey of São Tomé and Príncipe.

Dr. Shevock drying moss specimens.  Phot. A. Colwell, 2009

Recall that when our expeditions began back in 2001, there were only four species of mushrooms known from the islands; as a result of GG II and III, Drs. Desjardin and Perry have identified some 225 species, including new ones.  Phallus drewesii, an endemic to São Tomé, was just described in August.  I have a strong suspicion that Jim Shevock is going to come up with similar surprises.

The other critter work will include hooking up with Jose Lima to obtain more shrew specimens and to find and identify the mysterious Charroco, the fish we missed on earlier expeditions and which is thus absent from the islands  list.  Jose is doing the research for his PhD with the University of Lancaster. Jose “rediscovered” the supposedly rare, possibly endemic São Tomé shrew, Crocidura thomensis.  It is certainly not rare; as so often is the case, one just has to know where to look. Ricardo does.  We have permission to collect a few and test their tissues to see if they are in fact true endemics, or whether they were brought to the islands via human activity.

“Cobra bobo” endemic to São Tomé.  Phot. J. Juste

You will recognize this as the flamboyant caecilian, Schistometopum thomense or “cobra bobo,” known only from São Tomé.  The photo was taken many years ago by my friend and colleague, Dr. Javier Juste of the Doñana Institute in Seville in Spain. Javier thinks he may actually have taken this photo on São Tomé, but he is sure that he has seen caecilians on Príncipe Id!  This would be most exciting, and of course we will be looking for it.

And now that we have found a myriapod (millipede) expert, Dr. Rowland Shelley of the North Carolina State Museum (see November post), we will be collecting these critters as well, and I have no doubt that this group will turn out to be as poorly known as the others…. More surprises in store.

A myriapod (millipede). www phot.

Now for the special focus.  I have long thought that the citizens of the Republic of São Tomé and Príncipe need to be aware how absolutely unique and special their islands really are.  My groups of scientists and I can continue to explore and conduct research and make neat scientific discoveries.  We can continue to publish scientific papers, and we even add Portuguese abstracts.  But while this is great for “Science”, the majority of Tomeans will never see these papers. This popular blog has been an attempt to “spread the word”, but the vast majority of the people there do not read English, and most certainly do not have computers.  What good does all of our work and discoveries do if the citizens who live there are remain unaware of how special their islands are?  For example, Martim Melo, an outstanding ornithologist and expert on the birds fauna of the islands has just established the fact that the islands of São Tomé and Príncipe, together, have the highest concentration of endemic bird species in the world!  I doubt if anyone on the islands knows this fact, and think of what such a statement might mean to tourism!  The people should know and be proud of the unique nature of their nation, especially because they will have hard decisions to make in the future, if and when the oil revenues come… that is why this blog is called the Island Biodiversity Race (go back to the first two postings, if you need to) – there is a real urgency to what we are doing.

So above and beyond our usual critter searches, I am going to spend a significant part of our time during GG IV meeting with various people who are involved in appropriate government ministries, education, tourism and the environment,  in order to come up with ideas for a multi-level educational program.  We hope to learn what the citizens want and need in this regard.  This is where the fourth member of GG IV comes in: Mrs. Roberta Ayres.

Roberta Ayres (left) in the Naturalist Center, CAS.  RCD phot. 2010

Roberta is Manager and Senior Educator of the Naturalist Center, which is a major part of the Koret-Taub Education Center of the Academy.  Roberta has a Master’s degree in science education and, having been born in Brazil, speaks fluent Portuguese.  Together, Roberta and I hope to learn how we can raise biodiversity awareness on the islands through our meetings and interviews with its citizens.

The California Academy of Sciences Naturalist Center. RCD phot. 2010

Barring technical or other problems, I plan to keep blogging from the islands.

The parting shot:

“Island Tranquility”- Laguna Azul, on Sao Tome. D. Lin phot. GGII

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 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.

The Race: Taming of the Shrew (and updates)

Things have been very busy.

Our flower people, Dr. Tom Daniel and Rebecca Wenk have been very active. Rebecca successfully completed her M.Sc . degree at San Francisco State University, based in part on plants she collected in the islands during GG III (A). She then published her research, with Tom, in the latest Proceedings of the California Academy of Sciences; the publication includes a special treatment of the genus ElytrariaE. mariginata is the little flower that Rebecca finally found high up on Pico Papagaio on Príncipe that was so exciting and which we reported last May. (See “News from the Flower people”).

Rebecca among the giant Begonias. Lagoa Amelia. Wenk camera: GG III.

Tom is nearing completion of his monograph on the acanthus (shrimp) flowers of São Tomé and Pr<!–[if gte msEquation 12]>í<![endif]–>ncipe. He is now collaborating with Estrela Figueiredo, a Portuguese botanist who has been on the islands many times and has added much to our knowledge of the botany of the Gulf of Guinea as a whole There are several species of the Acanthaceae found only on São Tomé and Príncipe and one (Heteradelphia paulowilhelmia) which may be endemic at the genus-level. We found this beautiful flower in the middle of Lagoa Amelia at 1400 m. during GG III.

Heteradelphia paulowilhelmia. Lagoa Amelia. Weckerphoto: GG III

Wes Eckerman (photographer) Rebecca and Tom at Lagoa Amelia. RCD phot: GGIII

Another species in Tom’s group that is of particular interest to me personally is an acanth called Justicia thomeensis. This flower is known only from São Tomé and has not been collected since the late 19th Century – the original collector did not provide detailed locality data, and we have not found it on the islands yet. We hope it is not extinct, or that it was not actually collected somewhere else. Tom has examined the original dried specimens in Coimbra; it is perfectly valid, and he is re-describing the species. Another botanist named Hedrén examined this material, and in a 1989 study found that J. thomeensis is more closely related to a group of species in East Africa than to any in nearby West Africa. I am finding the same strange disparate distribution patterns among my island frogs.

Justicia relationships. RCD

On the crustacean front, Alex Kim, our freshwater shrimp colleague (see January and March blogs) has decided to attend Harvard for his undergraduate studies, and this summer he is doing fieldwork on prawns in Puerto Rico. He tells us that the São Tomé specimens collected for him earlier this year by the marine group “..represent at least two species which, based on morphology, have clear affinities with New World forms. DNA analysis is still ongoing, but the geological youth of these prawns leads me to suspect that we will soon have genetic evidence of trans-Atlantic larval dispersal.” Pretty impressive stuff for an incoming freshman.

A New World prawn.  Alex Kim phot.

A few months ago I had an opportunity to lecture on some of the scientific results of our Gulf of Guinea Island expeditions to a group of biologists at international meetings in Sardinia; afterward in Spain, I met one of the foremost experts on bats, Javier Juste, of Institute Doñana in Seville who has also worked on the islands and has discovered some of the same strange evolutionary relationships that we have. We are about to send him bat tissues from our GGI collection for DNA extraction.

7th Congresso Nazionale, Societas Herpetologica Italica. Sardinia. C. Corti phot.

In earlier blogs, we discussed the fact that mammals make poor dispersers over saltwater barriers; except for some bats, mammals simply cannot survive long enough to colonize oceanic islands. This is because we mammals have to eat regularly in order to maintain constant body temperatures –without “stoking the furnace” by eating regularly, mammals quickly die of exposure. This is specially the case with shrews; because of the ratio of their tiny body masses to their surface area, shrews lose heat faster than any other mammal, and a shrew has eat almost constantly, or it dies of hypothermia. It is perhaps a testimony to the great geological age of São Tomé and Príncipe that shrews appear to have, nevertheless, successfully colonized both islands! If the two species are indeed valid and occur naturally (not brought to the islands accidentally through human agency), then these may be the only oceanic island shrews in the world.

The Sao Tome shrew (Crocidura thomensis). R. Lima phot. 2009

To our knowledge, the photographs above and below are the first ever published of the supposedly endemic São Tomé shrew, Crocidura thomensis. The photographs were taken by Ricardo Lima, a doctoral candidate at the University of Lancaster who is studying the environmental effects of various agro-forestry techniques on São Tomé. Ricardo tells us that the shrews are not at all rare; in spite of this, we don’t know much about this little critter, nor the one on Príncipe. The São Tomé species was first discovered in 1886 by the great Portuguese explorer, Francisco Newton and described in 1887.

C. thomensis. R. Lima phot. 2009.

The relationships of these small island insectivores were not assessed until nearly 100 years later by Heim de Balsac and Hutterer in 1982. These authors concluded (on morphological evidence) that the São Tomé shrew was a full endemic species, and that the Príncipe form was an endemic subspecies of Fraser’s musk shrew (C. poensis) which is widespread on mainland Africa.

Ricardo Lima and friends, crossing the Rio Lemba, Sao Tome.  2009.

Now, we are in the exciting position of being able to test these assumptions using modern genetic techniques; just how closely related are the two shrews, based on DNA sequence? Did these mammals arrive naturally by rafting, as we suggest for the amphibians? Perhaps millions of years ago? Or were they hitchhikers on an old Portuguese galleon a couple of hundred of years ago? We hope to find out.

Aspergillus dykowskii and Sarophorum palmicola on monkey pod cacao; Lagoa Amelia. D Lin phot. GG II

Finally, our mycologists have a huge job. As you know before our expeditions began back in 2001, there were only four species of mushrooms known from São Tomé, and Príncipe had never been sampled. Now, after Dr. Dennis Desjardin’s work on GG II (2006) and his subsequent return with Dr. Brian Perry during GG III last year, we now have 225 species (including 75 listed for the first time on Príncipe). Many of these are probably new to science and it will be a major effort to fully analyze the entire collection. But Dennis and Brian took the opportunity to describe one of the new species separately. It has just been formally published in the journal Mycologia, and they have named it after me! It is a weird looking thing, and I will describe the whole process (including the humor sometimes involved) in the next blog.

Here’s the parting shot:

“After the Race.”  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, and Victor Bomfim, Salvador Sousa Pontes and Danilo Bardero of the Ministry of Environment, Republic of São Tomé and Príncipe 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 G. Breed, Gerry F. Ohrstrom, Timothy M. Muller, Mrs. W. H. V. Brooke and Mr. and Mrs. Michael Murakami for helping make these expeditions possible.