A very strange danio

Every well informed freshwater ichthyologist is familiar with petfrd.com forum.  Petfrd does not easily translate to Singapore Cichlid Community, which seems to be the patron of the site, but I have no clue to what petfrd could mean. The forum has several sections, and postings are enthusiastic and often with beautiful images of habitats or fresh collected fish contributed by local aquarists from all over southern Asia (and others, sure) . Ng Heok Hee moderates a section about scientific literature and usually is abreast of locating new papers about cichlids or Asian fishes. So, here is the secret about staying informed: daily visits to petfrd.com!

Betadevario ramachandrani in the aquarium. Photo (c) Beta Mahatvaraj

In January 2008, information and photos were posted of  what looked like a danio but sufficiently different not to be ignored as ‘one more…’. ‘Beta’, with a sad smiley reported that his fish were in the freezer. Fang immediately got in contact with ‘Beta’, whose name is actually Beta Mahatvaraj, enthusiastic Indian aquarist. His five frozen fish were preserved in ethanol and formalin and shipped to us. They weren’t really fresh, and the colour somewhat faded, but still all anatomy and DNA could be extracted from hem. Fang started writing up a description and gave it a manuscript name, ‘Betadevario longibarbis’.

It wasn’t a Danio as some of the first reports had it. It was more like a Devario, with those longitudinal depressions above each eye filled with sensory organs, characterizing Devario and Chela. But still not looking like a Devario, above all because of the long barbels, but also with regard to the coloration, which is unique among danios, with a light band along the middle of the side, and dark abdominal sides. Devario typically have very short barbels or they are missing completely, whereas most Danio have long barbels.

Actually, Devario is a speciose genus with more than 50 species, many still undescribed, and there is considerable variation in shape and colour pattern among them. They never have long barbels, though.

Since everything takes long time for us (the Cichla revision published 2006 took 17 years, and other papers not published were written in the 1980s, to give you a hint), it wasn’t unexpected to learn in 2009 that an Indian team was also working with the same species, and also with very few specimens. It took till December 2009 before we established contact and things were arranged for a collaborative effort to get the fish described. In the meantime much more material had become available, and indeed there were already numerous presentations of the fish on the web, e.g., in PFK  and in several Indian news sites.  Collaborating meant we could provide a fairly complete review of the new species with habitat data and image, live colour photo, and molecular and morphological phylogenetic analysis, better than individual papers would have been.  Betadevario got the species epithet ramachandrani for A. Ramachandran. Our molecular and morphological analyses differ with regard to its placement in the phylogenetic tree, but it is certainly a very basal taxon among Devario like danios, and we have more confidence in the molecular data which places it as sister group to other Devario.

Why not make it a Devario then? A good question for any genus, and will never have a good answer for all who come up with it. In this case we reasoned that the molecular analysis provided better clarity. The morphological dataset was good for distinguishing Devario and Danio, but not for resolving relationships within Devario. We have to work a lot more on that. And we are doing it, with both morphology and molecules. Betadevario presents a unique colour pattern and long barbels to distinguish it from all Devario (including Inlecypris),  providing a morphological justification for the genus.

Betadevario ramachandrani immediately after capture. Photo (c) P.K. Pramod.

The largest specimens measured about 60 mm. There are no characters to distinguish males and females, except the tubercles on the pectoral fin in males, a secondary sex indicator shared with most Danio and Devario. Information on live colours are not in concert. The live fish on Beta Mahatvaraj’s image are golden and brown/blackish, whereas the illustration by P.K. Pramod shows a a fish with a blue stripe along the side and lemon yellow fins. It will be interesting to see Betadevario alive sometime. It does not look like it will be a big aquarium fish, but who can say. It is so far known only from a small mountain stream in the Western Ghats, with relatively cool water, and may be expected to be sensitive to transportation.

Betadevario ramachandrani is known only from a very small area in the Western Ghats, the mountain range along the western coast of India, where it lives in fast running clear waters in the forest.

You don’t need a longer story here. The full description of Betadevario ramachandrani with habitat image, trees, and other details is available free to download from the Zootaxa website.


Pramod, P.K., F. Fang, K. Rema Devi, T.-Y. Liao, T.J. Indra, K.S. Jameela Beevi & S. O. Kullander. 2010. Betadevario ramachandrani, a new danionine genus and species from the Western Ghats of India (Teleostei: Cyprinidae: Danioninae). Zootaxa, 2519: 31-47.


Thanks to Beta Mahatvaraj and P.K. Pramod for making the images of live fish available.

In Memoriam: Alf Johnels

My former superviser, Alf G. Johnels, passed away on 24 May, 2010, at age 93.

Alf has a long and interesting professional record, crowned by his professorship at the Department of Vertebrate Zoology at the Swedish Museum of Natural History (1958-1982), and his presidency of the Royal Swedish Academy 1981-1983. He was also a key person in the discovery of mercury poisoning in Sweden in the 1960s, which laid the foundation for advanced ecotoxicology. By analyzing bird feathers in museum collections, Alf and collaborators could show that mercury poisoning in birds coincided with the introduction of methyl mercury as a pesticide. This led to the establishment of a biobank at the Swedish Museum of Natural History in which samples are preserved under sterile conditions, and which will enable the study of time series of toxic substances in the environment.

More important for us, however, is Alf’s ichthyological career. It started in early childhood, and led him to become a fisheries biologist first — to be certain of a fallback — but he also studied lamprey anatomy at the Zootomical Institute (now Department of Zoology) of the Stockholm University where he obtained his PhD in 1948. Those were the days when anatomy was important, and every biologist knew the name of every cell and every structure inside an animal, and the Zootomical Institute had several world renowned researchers of the kind. Alf and others in Stockholm were very much interested in the early development of fishes, as a way to understand the formation of organs in adult fish. The one major field trip Alf made had as objective to sample ontogenetic series of various “primitive” fishes, principally lungfish and bichirs.

Together with Ragnar Olsson and Björn Afzelius he travelled to the Gambia in 1950, staying there from May till November. This was long before the Gambia became a tourist spot. The trio settled in Bansang, in the middle of a swampy area full of lungfish and bichirs. They searched out nests of these species, and took out and preserved some eggs, later larvae, about every day. They also collected other fish, of course, and together with the material from an earlier expedition in 1932, the University built up a significant collection of Gambian fishes. The collection is now in the Swedish Museum of Natural History. The team also took back quite a number of living fishes, and some of the smaller species entered the aquarium trade in Stockholm for a while.

In 1954 Alf produced a descriptive catalog of the collections made at Bansang, in which he described three new species, Pellonula afzeliusi, Syngnathus olssoni, and Barbus svenssoni. Unfortunately, they are all synonyms. Chrysichthys johnelsi, named after himself by Daget, is however still a valid species. He also produced a small paper about the scales of Gambian fishes, showing that in the low water/dry period, the scales are eroded at the margin.

The Swedish Expedition to the Gambia, 1950.
From left to right Björn Afzelius, Alf Johnels, and Ragnar Olsson. Photo Alf Johnels.

After retirement, Alf had time to devote to his main passion, salmon fishing, going fishing ever so often to Norway and returning empty-handed. One of his last works was a descriptive catalog of the salmon rivers of southern Norway. Nevertheless, he kept coming to the museum every day till just a few years ago when his health conditions became preventive.

Alf was a very special person, always neatly dressed, always a bowtie on, always well articulated on any subject of science. His mind wasn’t as conventional as his dress, and he described himself as more of an engineer than a scientist. It is well remembered when he helped us fishing stone loaches — dressed in suit, and using his umbrella as fishing gear.

Johnels, A.G. 1954. Notes on fishes from the Gambia River. Arkiv för Zoologi, 6: 327-411.
Berg, W., A. Johnels, B. Sjöstrand & T. Westermark. 1966. Mercury content in feathers of Swedish birds from the past 100 years. Oikos, 17: 71-83.

In Memoriam: Fang Fang

Fang Fang in lab

Fang in the lab, May 2002. Photo (c) Staffan Waerndt.

Little has appeared in this blog lately, although its author is never far from the keyboard. The main reason has been the year-long, now ended suffering of my wife and research team member, Fang Fang. The last paper that she actively authored will appear the coming week, and it seems timely to write something about Fang here and now.

Fang was born in Beijing in 1962. She had an MSc in fish biology and left a position at the Institute of Zoology in Beijing when she came to Sweden in 1993 to do her PhD at Stockholm University. With time she turned Swedish, and completed her PhD with a dissertation on danios which she defended in 2001 with Richard P. Vari as opponent.

Fang remained all the time with the Swedish Museum of Natural History, upholding various positions, eventually as a curator in the Swedish FishBase team where, among other things, she was the key person behind the triannual Artedi Symposia. She was probably the greatest fan of Artedi ever. For over two years she also coordinated ECOCARP, a major collaborative project with several European and Chinese laboratories joining to search for Asian aquaculture fish candidates. Over the last years there was not much time for research. She was secretary in the European Ichthyological Society, member of the IUCN freshwater fish specialist group, and much more. Nonetheless, she was the author of 30 papers, and two are still to appear.

Fang passed away on 19 May, 2010, after a year of fight against cancer. She is buried in the St. Botvid cemetery south of Stockholm, in a peaceful setting overlooking a bay of Lake Mälaren.

Fang was an open, cheerful person that made many friends within and outside science, and in many ways made ichthyology more fun. With her passing, Ichthyology has lost much of its charm.

Fang’s research focussed on danios. Her favorite fish was one she discovered in Myanmar in 1997, a very distinctive danio, golden with dark brown spots, which she named Danio kyathit. It  soon became a much appreciated aquarium fish. Although there is a species named after her, Devario fangfangae, her name will probably be more firmly associated with Danio kyathit. Her legacy includes an additional nine species of Danio and Devario, and several other taxa.

Read more about Fang on the family’s memorial web site.

Species of danios described by Fang:

Devario maetaengensis (Fang, 1997)
Danio kyathit Fang, 1998
Devario apopyris (Fang & Kottelat, 1999)
Devario leptos (Fang & Kottelat, 1999)
Devario acrostomus (Fang & Kottelat, 1999)
Danio roseus Fang & Kottelat, 2000
Danio aesculapii Kullander & Fang, 2009
Danio quagga Kullander, Liao & Fang, 2009
Danio tinwini Kullander & Fang, 2009
Devario xyrops Fang & Kullander, 2009

Cleaning out the bugs: fruitfly name to be based on science not convention

The International Commission on Zoological Nomenclature just published an opinion on  the name of a laboratory animal known as fruit fly, and identified as Drosophila melanogaster. Yes, that standard lab creature to sort into phenotypes during genetics classes. An application was made in 2007 requesting that if phylogenetic research would show that the fruit fly does not belong in the genus Drosophila (it has always been in the subgenus Sophophora, which then is the best candidate), the generic name should nevertheless follow it by making D. melanogaster type species of Drosophila. There was quite some discussion, difficult to understand the heat of, because the fruit fly hasn’t had its name changed yet, or anything very near. Many opposed to a potential change of the generic name for melanogaster, and supported the application. There were also voices to let taxonomy have its way, change the name if need be.

In fishes, the zebrafish, tilapias, and rainbow trout – maybe a magnitude less published on, but nevertheless important species beyond ichthyological introversions — change name in a snap the day the message comes. People were equally happy with Danio rerio then Brachydanio rerio and then Danio rerio again. Tilapias are known now mostly as Oreochromis — bothers noone. And the rainbow trout is Oncorhynchus mykiss — not Salmo gardneri (doubt very few remember this “well-known” name anymore). Entomologists may be less elastic than ichthyologists, giving them more headache and pains. Ichthyologists know that fish (no scientific name known) are a paraphyletic group, nevertheless do not demand that Pisces must be used by force.

The reflection of systematics in the names of organisms and groups of organisms is one of the strengths of Zoological Nomenclature. When a species goes from a bad to a better generic allocation based on scientific evidence it gets a new generic name, and that is a very strong signal of progress. Otherwise perhaps day-to-day taxonomic landwinnings would go unnoticed. As evidenced by thousands and thousands of nameshifts over time, this is not a challenge at all to systematics or science otherwise.

Nevertheless, as in all communities there are other opinions. One drawback of nameshifts is that literature searches get more complicated, one has to seach for two instead of one name. In some databases, such as GBIF and FishBase this is taken care of because FishBase and other databases keep synonymies and serve results under the presently valid name regardless of what name was used in the references. Not magic, hard manual work over the years. But so, if you look in older literature from Artedi onward, a large part of the species descriptions are taken up by synonymies, name lists in which authors made an effort to list all mentions of a particular species. This is not done anymore because the lists would become too long and require an amount of double-checking that few modern workers can afford. Instead we look forward to the Global Names Architecture and dependable search and retrieve databases based on resources such as ZooBank. Managing names, however, will not reduce the really major problem with the knowledgebase, namely misidentifications. A solution with misidentifications has been proposed that uses species concepts (based on stated definition of a taxon by and author) instead of named species, but that approach is not very helpful because it tends to have  a different concept of the same species for every publication, and in principle makes the name independent of the species. No gain. So, we are going from Opera omnia to history-free papers, and that causes worries (about names, where the worry should be about misidentifications) and technology will need to come to rescue. And will. Someday.

Back to the fruit fly. When I was a student it was called banana fly (it is actually a vinegar fly…?), like all other small flies swarming around the fruit in the summer. What scientific name it had did not matter to teachers or students alike. And there are lots of fruit flies that are not Drosophila, D. melanogaster, or even the same family. The paraphyletic genus Drosophila as of today has 1,450 species.

The Commission has now (April 2010) issued an Opinion in response to the Application to change the type species of Drosophila to be melanogaster. The Commission says no – 23 against 4. Commissioners have different opinions, but seem to converge on the case being about taxonomy, not nomenclature, so it is not a matter for the Commission. It is also mentioned that one should not deal with hypothetical cases, as this one – actually the fruit fly is still in its old genus. You can read it all on the Commission website: ICZN Opinion on Case 3407

A Google search today finds 1,040,000 results for fruitfly (includes fruit fly); 1,570,000 for  “Drosophila melanogaster”, and an astonishing 2,880,000 for Drosophila without melanogaster, suggesting that he scientific species name combination is not that worked in rock. Nonetheless, Sophophora melanogaster has currently only 93,700. Let’s see how that has changed a year or two from now.

And the winner is:


Fruit fly busy with something else. Photo: Botaurus, Wikimedia. Public domain.

Danio year 2009: Rakhine Yoma

thermometerat minus 10 degrees

Stockholm area, Sweden, late December 2009

In the last few hours of the first decade of the 2000s, realising that my friends in the western Pacific time zones are already into the new decade, and writing this is not likely to come to an end this year. Snow and cold (-10 to -15C) outside demonstrate why tropical fish won’t do in the garden. It was night outside already at 1600h, but inside electricity glows in every corner and it is a comfortable +23C, so the fish inside are just fine.

In 1998, Ralf Britz and I travelled westward from Yangon in a battered but still comfortable Toyota minibus, with driver and guide. We did not have permission to fly to Kalaymyo to look for Badis ferrarisi (this was a Badis trip, and a considerable contribution to the Badis revision of 2002). Our substitute goal was the western slope of the Rakhine Yoma (aka Arakan Mountain Range), separated from the Ayeyarwaddy drainage and ichthyologically quite unknown. Of course Day had been there, up to Akyab (Sittwe), and Hora had reported fish from Sandoway (Thandwe), but nothing substantial was known. It took a whole day to travel the dirt roads and hitting Gwa on the Andaman Sea coast, arriving late in the evening. Of course we made a few stops on the way, but very brief.

Sampling sites in Rakhine Yoma in 1998.

The soft hills of Rakhine Yoma in a top view.

We then travelled north along the coast to Taunggok and returned to the Ayeyarwaddy at Pyay, making excursions up and down logging roads to find small streams with freshwater. The coastal stretch along the main road was either brackish water or no suitable water at all. Unfortunately, in the remaining four days, no badid came into the seine, so badids are probably absent from the area we sampled, betwen Gwa and Taunggok. That may be significant, or just due to poor sampling. We did get quite some interesting Garra, already described by me and Fang (2004), and quite some danios and other cyprinids endemic to the region. The Rakhine Yoma forests are actually a biodiversity hotspot with numerous endemic species of animals and plants, so no surprise endemic species of fish are also abundant. Endemic fish species that come to mind are Danio feegradei, Garra vittatula, G. rakhinica, G. flavatra, G. propulvinus, G. nigricollis, Puntius binduchitra, Batasio elongatus, Hara spinulus, Akysis vespertinus, and Channa pulchra.

Danio feegradei has already been described from Thandwe (then Sandoway) by Hora, who also reported Danio choprae from the same locality. The latter was first described from near Myitkyina in the upper Ayeyarwaddy drainage. The danionins we encountered, of course included D. feegradei, but also Hora’s choprae. The latter turned out to have very little in common with D. choprae and in fact to represent a new species. Along with it we were impressed by a huge danio that had two dark blotches on the side and inhabited small streams in the deep forest. This one nobody seems to have seen before, and the colour pattern was certainly not reported before.

Eventually these two new danios got their names. In 2009, 11 years and a bit more after we collected them. Why it takes so long time from discovery to description?

Danio aesculapii, preserved holotype.

The choprae-like fish was named Danio aesculapii, and with its 29 mm max SL in the wild is one tiny fish. The name was inspired by the popular name of snakeskin danio. In the living fish, not much of snake pattern shows, the most impressive in this highly agile being is a golden shine along the middle of the dorsum. In the preserved fish, however, there is a series of dark vertical bars anteriorly on the side and two rows of dark spots from above the anal-fin origin to the caudal-fin base. Unlike most Danio, D. aesculapii does not possess stripes on the caudal fin. The caudal peduncle is relatively deep, and has more scales, 12 around the middle, than most other Danio, which have 10 only. Asklepios or Aesculapius was the Ancient Greek god of medicine, typically equipped with a staff with a snake or two wrapped around it.

Devario xyrops. A male, just collected.

The large double-blotched species got named Devario xyrops with reference to the sharp ridges rising along the anterior part of the orbit (xyrops means razor-eye), and is also diagnosed by the colour pattern with a diffuse blotch formed by irregular vertical bars anteriorly on the side, and separated from a well-defined dark stripe on the posterior part of the body. It was getting dark, and we were exhausted after chasing fish up and down the stream all afternoon, but realising that this was an exceptional species, we made an effort to take a live colour photo as per above. The pink sheen ventrally probably derives from the red lid of the container on which the photo tank was placed, however. Devario xyrops is large for a danio, with the largest specimen 77 mm SL. Comparing with other Devario we found that the colour pattern is not fully unique. Also Devario browni has a separation of the lateral marking into an anterior blotch and a posterior band, but not as conspicuous as in D. xyrops. A similar species, D. anomalus, was described by Conway et al. (2009) from Bangladesh, and differs in being much more slender.

Small pool close to Thandwe, habitat of Danio aesculapii.
Habitat of Devario xyrops and Danio aesculapii north of Thandwe.
And with that pass the last day of 2009 and the first of 2010. It is still cold out there, and very dark. Not a very inspiring setting, but not all days are like this so I do dare wish you a productive 2010 full of excitement and happiness!

To learn more you can download the original descriptions of both Danio aesculapii and Devario xyrops which are available online as Open Access from the publisher’s website:

And here are other papers mentioned (none of them Open Access):
Conway, K.W., R.L. Mayden & K.L. Tang. 2009. Devario anomalus, a new species of freshwater fish from Bangladesh (Ostariophysi: Cyprinidae). Zootaxa, 2136: 49-58.
Hora, S.L. 1937. Notes on fishes in the Indian Museum. XXXI. On a small collection of fish from Sandoway, Lower  Burma.  Records of the Indian Museum, 39: 323-331.
Kullander, S.O. & R. Britz. 2002. Revision of the family Badidae (Teleostei: Perciformes), with description of a new genus and ten new species. Ichthyological Exploration of Freshwaters, 13: 295-372.
Kullander, S.O. & F. Fang. 2004. Seven new species of Garra (Cyprinidae: Cyprininae) from the Rakhine Yoma, southern Myanmar. Ichthyological Exploration of Freshwaters, 15: 257-278.
Image credits:

Map from the NRM Ichthyology Collection online database, using Google Maps

All other images: Sven O Kullander, CC-BY-NC

At last: calendar boy

Being a well-known figure in fish circles can lead to occasional highlights. Last year I delivered my first fan autograph, quite unexpectedly, and I know I didn’t manage it with the speed of a movie celebrity. And now, the next step in an ichthyological career: calendar boy. Chatting with Christer Fredriksson in his shop Akvarielagret, Christer came up with the idea to include me and my goldfish pot in the annual calendar of the Nordic Cichlid Society. And that did not work out, so the outcome is actually a shot in front of the two aquariums in our lab corridor (African lungfish in one, some other fish in the other), holding a jar of Cichla, wearing a T-shirt from the Sydney Aquarium, and a FishBase vest.

Christer is a highly motivated aquarist and innovative entrepreneur with the largest aquarium shop in Stockholm, and probably the most interesting shop in terms of fishes because he is not only focussed on selling, but also displaying and breeding quite a number of less common species. The business was recently expanded to include the Butterfly House at the northern outskirts of Stockholm (actually in Solna, the siamese twin city of Stockholm) which is now being converted to have more aquatic components. The most interesting recent move, however, was to convert supplier pricelists into a searchable online database. Customers can seach for the fish they are interested in, see the approximate retail price, and order online for pickup at shipment arrival. This saves time and money for both parties. Christer does not have to guess what customers may like, and customers can make more directed decisions.

Christer is also a persistent supporter of the Nordic Cichlid Society (Nordiska Ciklidsällskapet), which has one of the finest cichlid journals in the world, presently edited by Ola Svensson who recently graduated from Stockholm University.

You may indeed wish to join the Nordic Cichlid Society, not least for the 2010 calendar with me in it (and a number of more worthy celebrities), but also for the many well-written articles and so many beautiful photos of cichlids in the Society journal. The only bad thing is that they insist on using Metriaclima instead of the senior synonym Maylandia, and that something that really needs a blog later on.

Nordiska ciklidsällskapet

Danio year 2009: Danio quagga

In March last year, our PhD student Te Yu Liao and I were able to collect along the Myanmar border with India, in tributaries to the Chindwin River. Our intended stops were Kalaymyo and Tamu, but we also tried stopping at streams along the road. It was very dry at the time, and even large rivers reduced to small streams.

Stopping by one of these streams, in the heart of a large village, we seined a pool with some vegetation. And in there was a zebrafish! Zebrafish in Myanmar, in the Ayeyarwaddy drainage. No way. It had to be something new, and as we have learned from Meinken, all danios are striped anyway. The fish went into formalin, and the chase for the next specimen commenced.

The stream

After one more hour, still not one more specimen of this schooling species, but hundreds of Danio albolineatus, and we were called to inspection by the local authorities, so we had to move on. Indeed, we were in a hurry between Kalaymyo and Tamu to arrive at our destination before dark, not only because of likely shortage of accommodation in Tamu, but also because we were given only one day of permanence in Tamu and would need to spend time with local authorities to explain our presence. But hand on heart, you can’t leave a new species like that? Well, we moved on convinced we would find more specimens in some other place. Unfortunately it did not turn out so. We left Tamu with four more specimens from the market, where they hid in heaps of plenty of other little fish, dead to the bone, and with the total of five we eventually left Myanmar. Myanmar markets have large fish, but also large quantities of very small ones, used to prepare a special fish paste, ngapi. In the early morning there is thus plenty of fresh fish in the market and saves on collecting in the wild, with the caveat that fresh fish from the stream preserve better.

So, we were back with only formalin preserved specimens, and in the present times that is bad, because DNA sequencing is in the vogue, and generally a useful tool to check on phylogenetic relationships, and because formalin denatures DNA. Some of us can still do systematics without DNA, however, and that we did. The striped danio turned out not very similar to zebrafish at all, except in the general colour pattern. It is rather related to the spotted species, Danio kyathit from the neighbouring upper Ayeyarwaddy drainage and with one specimen recorded from the uppermost Chindwin.

The mammal: Equus quagga.

We named our striped danio Danio quagga. Quagga is the species or subspecies epithet of one of the zebras, so that within the genus of the zebrafish there should be at least one species with a name associating to zebras. We didn’t call it Danio zebra just not to confuse things for the zebrafish people (and perhaps, in the end, ourselves). (But wouldn’t it have been fun?)

The fish: Danio quagga, holotype.

See above what it looks like, now that the holotype has been dead for a while. Exciting as it was to encounter another zebrafish in the wild, I am not convinced that this will become an ornamental fish hit. Our comparisons suggest that it is most close to Danio kyathit, which hasn’t outcompeted the leopard danio in the hobby. It is also somewhat larger than a normal Danio rerio, requiring more swimming space (the holotype was the only one in 50 m of stream …). And, of course, why would we need another striped danio in our tanks. Seems we have exhausted danios as ornamental fish? But there is of course more to a fish than populating an aquarium. We are gradually building a phylogenetic history of danios, and then it is just great to find a sister-species pair, a split branch on the tree, rather than having single branches of uniques. Now we will move on to connect D. quagga and D. kyathit to the rest of the tree.
If you want to read the original description, you can download an Open Access copy, which you can share with families and friends as much as you like. Just click on the reference below. It’s magic!

Kullander, S.O., T.Y. Liao & F. Fang. 2009. Danio quagga, a new species of striped danio from western Myanmar (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 20: 193-199. Open Access PDF

Image credits:
Map from the NRM Ichthyology Collection online database, using Google Maps
Danio quagga and stream: Sven O kullander, CC-BY-NC

Equus quagga: Muhammad Mahdi Karim, from Wikimedia Commons, GNU FDL 1.2

Danio year 2009: Danio tinwini

The year is isn’t over, and the snow did not fall over Stockholm yet, so it may be a bit early to summarise the year. But it might be better to start early, not to end up in december 2010 summarizing 2009, and then it has to be piece by piece. Writing this, I am reminded of one of the most famous aquarium books, Aquarienfische in Wort und Bild, which was issued over a period of 30 years in instalments of a smaller number of sheets, and the authors were indeed writing continuously on it between 1932 and 1962. In the 1950s there were similar initiatives elsewhere in Europe, e.g., the Belgian Vissentlas, but I have not kept track of them. Also TFH publications tried, much later, to publish a loose-leaf edition of Exotic Tropical Fishes, and the German cichlid society sent out species descriptions with their journal supposed to be collected by the subscribers. Of course, this is in most cases an awkward way of publishing where the customer is the big loser. The following scenarios are possible:
  • Publication terminates after a smaller number of pages. You end up with a quarter or half of a book.
  • Publication goes on indefinitely, with or without revisions of outdated pages. You end up with a book containing almost duplicate pages, and a span of many years between the earliest and the latest information (could be blurred b/w photos mixed with excellent digital colour photos. Half the book will be outdated.
  • You stop subscription and end up with a half book
  • Everything completes but in the meantime you already had to buy some complete books, just not to stay ignorant about everything from H to Z.

Die Aquarienfische in Wort und Bild (there is also the corresponding Aquarienpflanzen) is an exception among the bad examples because it (1) terminated a long time ago, so all you can buy now is the complete book (€70-150 on eBay), and (2) it was actually quite complete in coverage, and especially the species descriptions were more detailed than anything in other contemporary aquarium books. I am actually not quite sure when it was published, but it seems it started in 1932, and went on till 1962, with Hermann Meinken (1896-1976) as the sole author after co-authors Maximilian Holly (*-*) and Arthur Rachow (1884-1960) long before had retired from writing. Die Aquarienfische in Wort und Bild was a highly influential work, the mother of all other European aquarium books.

That it wasn’t published after 1962 seems likely because the description of Brachydanio frankei, described by Meinken in 1963, is missing. Every other fish imported to Germany up till the late 1950s/early 1960s are there, and Meinken had a particular interest in danios.

Now we are getting to the point: Brachydanio frankei was, at that time, known only from aquarium specimens. It is still not known from the wild, as already pointed out in an earlier post about fish from nowhere. Meinken lists the following species of danios:

Danio devario
Danio malabaricus
Brachydanio kerri
Brachydanio albolineatus
Brachydanio nigrofasciatus
Brachydanio  rerio

That is a very short list. (And by the way, Brachydanio is presently considered a junior synonym of Danio, and Meinken’s Danio are now in Devario; but they are all danios.) A much larger number of species of Rasbora are described in the same book. At present it looks to me that there are more danios than rasboras in the shops, but I may may be biased.

Danio rerio, aquarel by Curt Bessiger, used in Die Aquarienfische in Wort und Bild.

The most characteristic about Danio frankei is the colour pattern, which is golden with small brown or black spots. At the time that was quite remarkable because all known danios (listed above) were striped (except that the stripe in Devario devario is not so prominent, and Danio nigrofasciatus has spots on the abdomen. Over the time, no wild specimens of D. frankei appeared, and it has turned out rather certain that it is a colour mutation of the zebrafish, Danio rerio.

In the original description of Danio frankei, Meinken wrote (my translation from the German):

Shortly before the end of the year 1962, through my old friend Heinrich Grauel from “Vivarium-Bremerhaven”, my friend Diplom-Biologe Hanns-Joachim Franke in Gera/Thüringen, gave me  six captive bred specimens of a very handsome Brachydanio novelty with the request that I study the specimens, if possible identify them and give them a name. According to the information from Mr Grauel, at that time Mr Franke already had bred several hundred specimens of this species in his breeding tanks. Unfortunately the novelty was lacking a name.

Upon my immediate checkback concerning details about the importation and my notice that — in case the animals would represent a scientifically still unknown and undescribed novelty — a determination based on captive bred specimens would be unreliable because that could lead to mistakes, Mr Franke then most friendly told me that he had obtained the import specimens from the familiar and active Diplom-Biologe Stanislav Frank in Prague. Unfortunately, the place of capture was unknown also to Mr Frank. That the animals came from India was only a very weak consolation, because first all Danio and Brachydanio species come from India and the Indo-Malayan area, and second India is large, about several times larger than the previously undivided Germany. Mr. Franke, however, made the great sacrifice in the interest of science and nomenclature, killing four of his import specimens and sent them to me for examination.

Meinken then went on with a very detailed description, a lamentation of the lack of locality data, and finally, a long discussion about relationships. According to Meinken, D. frankei would be most closely related to B. tweediei (from Malaysia), a nominal species currently synonymized with D. albolineatus.

Leopard danio. Preserved aquarium specimen from Taiwan.

That did not stop other leopard danios from coming in. The first was Danio kyathit, described by Fang in 1998. The second was just described by me and Fang based on a large series collected by U Tin Win in the same region as D. kyathit, and we named it Danio tinwini.

Danio tinwini, female paratype.

Danio tinwini is spotted and the spot pattern is quite similar to that of D. kyathit, but not so much to that of the leopard danio. It is a much smaller species than either D. kyathit or the leopard danio, however, and external morphological characters suggest that it is more related to D. nigrofasciatus or D. aesculapii. It is indeed a remarkably small species. The largest wild male examined was 21.7 mm in standard length, the largest female 25.6 mm. That is almost as small as the smallest Danio species (D. margaritatus, 21.3 mm).  What provokes most thought, however, is the fact that the only two spotted danios, D. kyathit and D. tinwini occur in the same general area (but not syntopic), and are not closely related. Would we be surprised if D. frankei were found in northern Myanmar one day as well?

Danio tinwini lives together with a much more famous fish, Danionella dracula, the Vampire or Dracula fish. Much more we do not know about the natural habitat och accompanying species. You can read all the technical details about Danio tinwini in the Open Access original description.

Holly, M., H. Meinken & A. Rachow. 1932-1962. Die Aquarienfische in Wort und Bild. Kernen, Stuttgart, Loose-leaf publication, 1328 pp.

Kullander, S.O. & F. Fang. 2009. Danio tinwini, a new species of spotted danio from northern Myanmar (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 20: 223-228. [errata:  p. 227, col 2, para 2, line 10: “1” should be “4”; p. 228, col. 1, line 3: “19.3” should be “21.2”]

Meinken, H. 1963. Mitteilungen der Fischbestimmungsstelle des VDA XLII: Brachydanio frankei spec. nov., der Leopard-Danio. Monatsschrift für Ornithologie und Vivarienkunde. Ausgabe B. Aquarien und Terrarien, 10:75-79.

*-* I cannot find any information about Maximilian Holly. I don’t even know where to start…?

Photo credits:
Top image: living Danio tinwini,Sven O Kullander, CC-BY-NC
Leopard danio and Danio tinwini, Sven O Kullander, CC-BY-NC
Danio rerio aquarel, public domain.

Penis fish

My friend Ralf Britz (Dracula fish), just released the description of a species related to the Dracula fish (Danionella dracula), viz., the Penis fish (Danionella priapus), which is number four in the genus (D. mirifica, and D. translucida were described earlier). This was a bit of of concern to me. Is the evolution of Danionella Nature’s idea of a freak show (or a Japanese game show…)? Or perhaps the fishy Fantastic Four (Bloodsucker, Flasher, Beauty, and Invisible). By the way, there is a Marvel character called Priapus.

Danionella priapus, the male holotype. Photo (c) Ralf Britz, reproduced with kind permission.

The penis, a worm-shaped intromittent reproductive organ characterizing mammals, some reptiles and very few birds, has fascinated male ichthyologists for a long time. The typical penis, in placental mammals, contains some spongy tissue and ducts for urine and sperm. Many species of fish with various elongate reproductive structures have been named with reference to the penis, or after the Greek minor deity commonly associated with an oversize penis (list below). The only group of fish that had escaped until 2008 were the Chondrichthyans, equipped with the most penis-like intromittent organ among fishes, known as mixipterygia and a synapomorphy of the group. Perhaps because in Chondrichthyans, the male reproductive organ, a modifiction of the pelvic fin, is paired. Yes, they have two. Theoretically a male shark could mate with two females simultaneously, but there seems to be no observation of such act and perhaps for reason that such pairing must be very complicated to coordinate, no other vertebrates invested in double external reproductive organ. But we all do maintain a paired internal set, except that in many fishes there is only one ovary. Marsupials, it must be added, are an exception here as in so many other respects. Males usually have a bifurcate penis unfit for peeing with and consequently used only for copulation. Females have two vaginas (with common opening), each with its own uterus, and a third for parutition.

In actinopterygian fish, sperm-transmitting elongate structures, intromittent organs, are technically known as gonopodium (live bearing poeciliids), or andropodium (Hemiramphidae; Goodeidae) in livebearers and are formed by modifications of anal-fin rays. Egglaying fish are commonly equipped with a genital papilla through which sperm or eggs are released. The genital papilla of females is sometimes referred to as the egg-laying tube or ovipositor. It is very impressive in bitterlings in which it is used for depositing eggs deep inside bivalves. None of these terms were considered suitable for a fish name, however.

What about naming fish after mammal female external reproductive organs? There is the south Asian Trypauchen vagina Schneider, 1801, a goby, for which Schneider did not explicitly explain the etymology but gave the local name, obviously based on information from the collector/correspondent in Danish India (Tranquebar), Christoph Samuel John (1747-1813), “Sewwöli, i.e., vagina cultri rubra, Tamulice”, what would mean: “Sewwöli, that is, red knife slide, of the Tamils”, i.e., nothing to do with humans. It is a slender, caudally tapering fish with a predominantly red colour. Schneider gave the same name also, spelled Sewwöti, for Cepola coecula (junior synonym of another goby, Taenioides anguillaris), another slim, red fish. There are also Thai cyprinids Probarbus labeamajor and P. labeaminor, named by Tyson Roberts in 1992, but for the thick and thin lips, respectively, of the mouth.

Trypauchen vagina

Priapos was an ancient Greek god of fertility, usually depicted with an oversize, erect penis, and according to Lemprière a rather deformed figure, not worthy of residing in the Olympics, and on the whole more of a rapist and rascal than anything else, requiring sacrifice in the form of of asses (Equus africanus, that is). This sad figure seems eventually to have converted into a garden gnome in Western garden culture. In Rome, Mutunus was the approximate equivalent. There is no fish named for Mutunus.

Interestingly, a very large number of fish possess a well developed genital papilla. In cichlids, for instance, it is long and pointed in males, but short and swollen, with at least a transverse fold in females, although there is variation, and in some genera it is hard to tell males and females apart on the papilla alone. Cottids are also well known for the long male genital papilla, as exemplified by Phallocottus, and it is also prominent in gobioids, but I could not find any relevant name in that group. The group that have most of the penile names are the livebearing poeciliids, in which the anal fin is modified to an elongate intromittent organ.

Back to Fantastic Four. As Ralf points out, these miniature fishes, and other miniatures as well, are remarkable not only for their small size. They also have conspicuous novel characters which gain significance particularly in a small body. The elongate fangs of Danionella dracula are the only oral tooth structures known from cypriniforms, and may very well be the beginning of a radically different evolutionary lineage. Males of Paedocypris (another miniature), and D. priapus have developed accessory sexual organs not known from other cyprinids, and may be actively evolving in organs that larger, more long-lived fish with longer generation times cannot afford to invest in.

Small size apparently calls for radical solutions, and those radical solutions may be the matter of evolutionary leaps. We are accustomed to think, at least I am, of evolution among fishes being based on something mid-sized rather normal fusiform, which then strikes off in different feeding niches, becoming smaller or larger depending on energy householding. But, that does not seem to lead to much real diversity (after all, all cichlids and cyprinids look very much the same). Radical morphological change as within Danionella may be one of the clues to evolutionary stepping. Most striking about species of Danionella is that they are so radically different from each other. Had they been a few centimeters longer, they would probably have been placed in different genera, but now they are considered as specialized close relatives. So, there is something important to be learned from the penis fish and friends.

Not all small fish present novel characters or behaviour, but the phenomenon seems to be particularly evident in progenetic species. This is then seen clearly in the larvae of many marine fishes in particular. The amazing spinature of many pelagic acanthopterygians, and the strange leptocephali of eels, demonstrate this. In this case, the “normal” adult provides for a backup morphology, should juvenile experimentation fail, but if these fishes turn progenetic and skip the adult stage, they may blast off in a different morphological evolutionary trajectory.

Origin of a new phylum? Larva of an unidentified species of the family Gempylidae. Photo NOAA.

Can I then go so far as to postulate that larval specializations must precede progenesis? Or does necessarily specializations have to come along with or after progenesis? Or am I making it unnecessarily difficult?

As you can now see, this post is all about evolution. But, you are right: I am also fascinated by biological nomenclature as a graffitti wall…

Anyway, here is the list of the penis fishes, based on active search in Catalog of Fishes. Many of them do not have a common name, so here is a goldmine for those constructing common names based on translations of the scientific ones … Of course all names are masculine, and all the authors are male… And, by the way, penis means tail, but is never used in that meaning, and the word phallos wasn’t intended for other than the drawings and sculptures, and again is not used in that meaning.

Phalloceros Eigenmann, 1907. Poeciliidae: Poeciliinae.

Allophallus Hubbs, 1936. Synonym of Carlhubbsia Whitley, 1951. Poeciliidae: Poeciliinae.

Arthrophallus Hubbs, 1926. Synonym of Gambusia Poey, 1854. Hubbs, 1926. Poeciliidae: Poeciliinae.

Aulophallus Hubbs, 1926. Synonym of Poeciliopsis Regan, 1913. Poeciliidae: Poeciliinae.

Curtipenis Rivas & Myers, 1950. Synonym of Poecilia Schneider, 1801. Poeciliidae: Poeciliinae.

Dicerophallus Alvárez, 1952. Synonym of Gambusia Poey, 1854. Poeciliidae: Poeciliinae.

Flexipenis Hubbs in Rivas, 1963. Synonym of Gambusia Poey, 1854. Poeciliidae: Poeciliinae.

Furcipenis Hubbs, 1931. Synonym of Alfaro Meek, 1912. Poeciliidae: Poeciliinae.

Heterophallina Hubbs, 1926. Synonym of Gambusia Poey, 1854. Poeciliidae: Poeciliinae.

Heterophallus Regan 1914. Synonym of Gambusia Poey, 1854. Poeciliidae: Poeciliinae.

Rivas 1963. Synonym of Gambusia Poey, 1854. Poeciliidae: Poeciliinae.

Phalloptychus Eigenmann, 1907. Poeciliidae: Poeciliinae.

Henn, 1916. Poeciliidae: Poeciliinae.

Plectrophallus Fowler, 1932. Synonym of Brachyrhaphis Regan, 1913. Poeciliidae: Poeciliinae.

Regan, 1913. Poeciliidae: Poeciliinae.

Priapichthys Regan, 1913. Poeciliidae: Poeciliinae.

Trigonophallus Hubbs, 1926. Synonym of Brachyrhaphis Regan, 1913. Poeciliidae: Poeciliinae.

Ctenophallus Herre, 1939. Synonym of Neostethus Regan, 1916. Phallostethidae.

Herre, 1925. Phallostethidae.

Mirophallus Herre, 1926. Synonym of Gulaphallus Herre, 1925. Phallostethidae.

Penicelinus Bolin, 1936. Synonym of Icelinus Jordan, 1885. Cottidae.

s Hubbs, 1924. Poeciliidae: Poeciliinae.

Phallobrycon Menezes, Ferreira & Netto-Ferreira, 2009. Characidae.

Phalloceros Eigenmann, 1907. Poeciliidae: Poeciliinae.

Phallocottus Schultz, 1938. Cottidae.

Phalloptychus Eigenmann, 1907. Poeciliidae: Poeciliinae.

Regan, 1913. Phallostethidae.

Solenophallus Herre (ex Aurich), 1953. Synonym of Neostethus Regan, 1916. Phallostethidae.

Galeus priapus Séret & Last, 2008. Scyliorhinidae.

Online reference:
Britz, R. 2009. Danionella priapus, a new species of miniature cyprinid fish from West Bengal, India (Teleostei: Cypriniformes: Cyprinidae). Zootaxa 2277: 53-60. (Abstract only; not Open Access)

Book references:
Lemprière, J. 1984. Lempriere’s classical dictionary. Bracken books, London, xv+734 pp. Original: A classical dictionary; containing a full account of all the proper names mentioned in Ancient authors. Reading, 1788. [Second edition, 1792] [11th edition, 1820]

Image credits:
Cover of Fantastic Four from Wikipedia, copyright, but used in fair sense, to illustrate that Nature and Fiction sometimes go hand in hand, you just need the imagination.
Trypauchen vagina from Wikimedia Commons
Gempylid larva from Wikimedia Commons.
Danionella priapus courtesy Ralf Britz

The New Nominomania

Roger Hyam’s blog post Calling time on biological nomenclature and the comments it received, also on Taxacom, makes me wonder if not biodiversity informatics is the enemy rather than the servant of science. What some of my colleagues argue for are empty name lists, including also artificial constructs like barcode species. Then erecting the haplotype as the focal point of taxonomy is apparently to be expected lying in ambush.

For taxonomists, names are abstractions of scientific knowledge, and cannot, consequently, be managed in a formalised top-down system. To call for science to be published in only certain journals, to advocate that certain kinds of “species” should be the only ones permitted, are not friendly proposals to rationalise information flows, but denials of the process of free information gathering. It is plain denying that taxonomic papers are primarily contributions to science in the first place, and name machines only secondarily. Taxonomy must remain a scientific exercise, and cannot be a mechanical process.

The idol project brought forth is the International Code of Nomenclature of Bacteria, where there is a Committee to decide, a single place to register names, and — most importantly, forgotten by the supporters — less than 10 000 diagnosable units are included. Since bacteria are so different from other organisms, and the named units so few (at least that have been admitted by this Committee …) the ICNB is simply not possible to use as a model for the several million species of multicellular organisms, most of which have not been named yet.

Whereas I am a friend of registration of names, and advocating that scientific names as defined in the Botanical and Zoological Codes are as good markers as can be (human-friendly they are) of scientific processes of elucidating the characteristics, whereabouts, and history of pieces of biodiversity, I cannot be positive to registration replacing the scientific procedure of testing hypotheses of phylogenetic distinctness labelled with scientific names. No committee should certainly be involved here. And whereas barcodes can probably be an interesting tool for the food industry and similar, I don’t see much use for it in taxonomy where we have species concepts based on evolutionary theory, type specimens, and diagnoses that are compatible with scientific theory and hypotheses. In taxonomy, contrast to the barcode shop, we also have flexible systems to classify biological units other than “species”.

Whereas taxonomists must be more collaborative with biodiversity informatics in, e.g., voluntary registration in ZooBank, and show more effort to make their work and naming visible, it is the task of biodiversity informatics to find the methods to discover, assemble, and present the objects of biodiversity. We must not adapt science to fit the index.

The concerted effort of GBIF and Encyclopedia of Life to build a Global Names Architecture (GNA), providing a Global Names Index (GNI), seems to me to be a way out of the dilemma that biodiversity informatics is entangled in: information about biodiversity cannot be extracted because there are too many names (with misspellings, synonyms, homonyms, etc.) out there and the approximate (can never be exact) meaning of a name may vary from one mention to another. Certain related efforts, such as transparently tagging names with identifiers, as is being done in Zootaxa and ZooKeys, are bridging the gap between computerified and human-mediated names. Thus the technology is there, it is evolving, and taxonomy should be able to continue as a science.

The real difference between the mega-name-consumers and taxonomy is that mega-name-consumers wish to have all in one place, which is probably of zero interest to taxonomy. They are also not interested in metadata such as diagnosis, type specimens, etc., and they do not want taxon concepts to change, which they inevitably must do in science. In taxonomy, only small sets of taxa (and names) are handled at any given time, and of these, all have a definite function in the particular study, may be a revision, a field guide, a phylogeny, or a classification. In such contexts, the name domain is self-contained, and all named units are related to each other by the hypothesis or scope of the study. Everything else is of zero interest. For a study of cichlid fishes, it is of no interest whatsoever if New Zealand Lepidoptera exist. Enter mega-name-consumers, who will need both in the same list because those lists are not based on any scientific criterion and it is absolutely not known what the list is for. If consumers could define their precise needs from study to study, it might be easier to design the tools to extract the names and concepts actually needed. To maintain lists of millions of names, even in a database, for no specific purpose does not make much sense. Indeed, most checklists of smaller scale as well, especially when produced by non-specialists are equally meaningless anachronisms of apparently undefeatable listmania.

So, we must ask from biodiversity informatics:

  1. Proper specification of what their taxonomic units (text-names or LSIDs) are going to be used for. Map species occurrences, make phylogenetic hypotheses, sort out homonyms, …?
  2. Design systems that can effectively detect, maintain, and trace name usage and relevant metadata, compatible with taxonomic objectives and procedures.
  3. Provide voluntary registration systems, and other tools facilitating the exchange of names and metadata between taxonomists and consumers.

Whereas 2 and 3 may be underway, I am beginning to doubt that anyone can give a good answer to 1…

For those who cannot embrace taxonomy fully, I recommend stamp collecting. It has all the flavors of registration, codes, hybridisation, phylogeography, central committees, misidentifications, rare haplotypes, identical reissues, fakes, top-down standards, and stasis. It is a totally unscientific enterprise with no limits to organisational options suitable for old frustrated men obsessed with control. Ooops, does it sound like DNA barcoding …?

Image: Wikimedia Commons, public domain