New look

This blog was built on the Theme Suffusion, probably the best ever WordPress theme. Sadly, the maintenance of the theme was discontinued, and as WordPress is now introducing a new visual editor, it seems the time has come to try something different. Since the rest of the blogging community is focused on maximum design and minimum text, and this blog heads in the opposite direction, it will at least be interesting to see how it ends, if it ends. Who can know. One good change is obvious, however, it is responsive.

Enthusiasm for nominomania

LundbergTitle2There is a little book – a dissertation actually – that lists every Swedish publication on fishes. Published in 1872 it of course had some advantage over any similar project to be raised today, but nevertheless it is a commendable work. It was presented as a doctoral dissertation at Uppsala University by Fredrik Lundberg, and  comprises 18 pages of introduction and 56 pages of bibliography. The author, Lundberg, vanished in the shadows of time, at least this dissertation is the only evidence I can find of the person. Both Fredrik (currently first name of 95962 men and 2 women in Sweden) and Lundberg (currently last names of 21123 persons, first name of 3 men and one woman in Sweden) are common names in Sweden.  Well, even if people may be interesting, it is a person’s work that counts, so I am basically content. Lundberg’s dissertation is important for tracking the history of ichthyology in Sweden, and for me it was the key to finding a rare publication that practically every other ichthyologist in Sweden refused to cite.

On page 29 Lundberg cites an article “Om Ichthyologien och Beskrifning öfver några nya Fiskarter af Samkäksslägtet Syngnathus. Af G. I. Billberg, (Linn. Samf. Handl. 1832, p. 47-55 m. 1 col. pl. Sthlm 1833).” The article was evidently in a journal with the name encrypted. It was somehow resolved as Linnéska Samfundets Handlingar (Proceedings ot the Linnéan Society). Decryption of journal name abbreviations is not for the impatient and weakhearted; luckily this tradition has been abolished in favor of very short names easy to mix up or very long names difficult to remember. As I could not find any further mention of pipefish species named by Billberg in other Swedish fish literature, or elsewhere – they were not incorporated into the Catalog of Fishes until in February 2016 – it was too good bait to resist.

This was in 2004 and although libraries were already restricting access to their older publications, online antiquariats were few. A copy of the particular journal issue could be found, however, in a Real Life antiquariat in downtown Stockholm for a considerable price. A second copy was lent to me by Professor Bertil Nordenstam, then at the Phanerogamic Botany department of the Swedish Museum of Natural History. The author, it turned out, was mainly a botanist or horticulturist, and the publication contains images and descriptions of plants

Image used in various printed and online sources, probably public domain

“Om Ichthyologien …”, indeed, the whole issue of the Linnéska Samfundets Handlingar (the first and only), and not least the curious author, were found to be extraordinary in many ways, good and bad. It was a discovery of a forgotten milestone in Swedish natural science that certainly needed attention. Billberg, a lawyer and judge,  botanist and natural historian by devotion, and funder of of the Linnéska Samfundet, attempted to present a new classification of fishes, and also, a man of classical education more than biological, had a lot to say about other people’s scientific names on fishes. The publication is sprinkled with new names on all kinds of fishes, family names, generic names, species names, but practically all of them needed to be evaluated in relation to the International Code of Zoological Nomenclature, and most of the fragmentary literature references pointed to sources not so easy to find in 2004 as they are now. So it wasn’t just the exciting discovery of three overlooked pipefishes. It was a true Pandora’s box, or can of worms, can of names.

Billberg proposed five new family names, only one of which survives as it is anolder homonym (Diodontidae). He mentions 61 genera of fishes, 41 of them listed only by name; out of  20 “new” generic names, none is valid. He he lists 31 species of fishes.; out of 28 “new” species names, one is potentially valid and a species inquirenda. Hardly anything in the taxonomy is justified by anything oyher than imprecise references. It turns out that Billberg probably based the whole paper on only one or two earlier works, by La Cepède (1798), and Cuvier (1817), with the outstanding exception of the description of three new pipefish species. The pipefish descriptions were based evidently only on three drawings made by Johan Wilhelm Palmstruch in 1806, probably from living specimens. So Billberg could have written his paper having examined zero fish, read two already long outdated books, and counted fin rays on three drawings. Of couse, the three new pipefish species are also junior synonyms.

Plate in Billberg 1833 showing new pipefish species 1, Syngnathus pustulatus (male 2, Syngnathus typhle), Syngnathus virens (female Syngnathus typhle), and 3) Syngnathus palmstruchii (Entelurus aequoreus)

Plate in Billberg (1833) showing new pipefish species 1, Syngnathus pustulatus (male Syngnathus typhle), 2 Syngnathus virens (female Syngnathus typhle), and 3 Syngnathus palmstruchii (Entelurus aequoreus)

What man had set his footprint so deep in the mud that it could not be retracted? In short, Gustaf Johan Billberg was born Karlskrona in Blekinge, southern Sweden in 1772. He studied law in Lund University and got a position as auditor in Stockholm in 1793. He took a similar position in Visby on the island of Gotland in 1798, but returned to Stockholm in 1808 and held various administrative and juridical positions there, mainly as a judge, until 1840. He became a member of the Royal Swedish Academy of Sciences in 1817, and corresponded with Linnaeus’s successor in Uppsala, Carl Per Thunberg, but he never had a formal education in natural sciences. He was a collector, with large entomological collections, and took particular interest in botany and economic botany. If he had not been caught in some controversy between the Academy and Uppsala University, perhaps he could have developed a career as a botanist. Instead he devoted his fortune and time to publishing more or less unfinished works that along with other events drove him to bancrupcy. Some of these publications are significant, like his two issues of the work Ekonomisk botanik (Economic Botany) and a few  parts of the book series Svensk botanik (Swedish Botany) and Svensk zoologi (Swedish Zoology), the latter in particular a pioneering work with descriptive text and hand coloured plates of Swedish animals. The society that he initiated, Linnéska Samfundet, was equally commendable, but quickly dissipated. The society produced just one issue of its proceedings, all articles in it written by Billberg, and apparently biologists showed no strong interest in the society. Billberg did make a lasting contribution, however, in developing one of the green areas in the heart of Stockholm, Humlegården. There he organised a Linnaeus Park, including a hilly flowerbed area still present today and known as Flora’s hill, named for his daughter Flora Mildehjert. Boethius (1924) wrote a detailed biography of Billberg.

Floras Hill

Flora’s Hill, May 2014. Photo Sven Kullander, CC BY-NC

Billberg’s enthusiam for natural sciences, particularly plants and animals, carried him high up among the clouds, and let him fall hard. When he died in the winter of 1844 he was broke and ill. By contrast, his brother Johan, without interest in natural history was ennobled af Billbergh in 1826. On the other hand Gustaf Johan brought up 9 children and one of them, Alfred, a medical doctor, became a well renowned pioneer in psychiatric medicine.

Years passed, however, as they tend to. “Om Ichthyologien…” remained a resting treasure as many other projects called for attention. The idea remained, however, to present an analysis of Billberg’s paper, and particularly to call attention to the existence of three forgotten species description contained in it. I started, stopped, and started, compiling names and checking literature sources. At first I thought that a tabular presentation would be enough, but no, too much needed to be said about this work. Eventually, after a senseless, sleepless final effort in early 2015 could I deliver a manuscript for submission. But it should take long time to see it in print. The main problem was obviously finding a reviewer. At last things could be resolved and in October 2015 there was an accepted manuscript. I will spare you all the details why its publication (Kullander, 2016) was then delayed till January 2016.

As you can read the whole analysis of Billberg’s fish names here, thanks to Open Access and somebody paying for that, this is not the place for reiterating detail that is already there. If you want a different context you can also find much of the information in the Catalog of Fishes.

Billberg’s many publications drew considerable criticism already during his lifetime, especially his unsuccessful habit of reforming the Swedish names on animals and plants. Billberg’s fish paper was ignored by all Swedish ichthyologists first probably because he was not accepted by the contemporary academics, and later because he simply fell out of memory. Several large volumes on Scandinavian fishes were published in the period 1836-1893.

Billberg has been called enthusiast, dilettante, and many other things, but on the positive side he was really an educator at heart, and it is difficult to criticize a person following a vocation to investigate things and try to make the world a better place, no matter how awkward the result then can be. The history of science is full of worse people. The worst that Billberg did was to put newly constructed names on plants and animals. That is something that many of us do …. Perhaps the review of his fish names can contribute to make him remembered more for his good aspirations than his formal failures. And serve to remind one always to be very careful when playing with names.

References

Billberg, G.J. 1833. Om ichthyologien och beskrifning öfver några nya fiskarter af samkäksslägtet Syngnathus. Linnéska
samfundets handlingar, 1: 47–55. [at Internet Archive]

Boethius, B. 1924. Gustaf Johan Billberg. Svenskt biografiskt lexikon, 4, urn:sbl:18212.

Cuvier, [G.] 1816. Le Règne animal distribué d’apres son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Tome II. Déterville, Paris, xviij + 532 pp.

Kullander, S. O. 2016. G. J. Billberg’s (1833) ‘On the Ichthyology, and description of some new fish species of the pipefish genus Syngnathus. Zootaxa, 3066:101–124.[at Zootaxa]

La Cepède, [B.G.] 1798. Histoire naturelle des poissons. Tome premier. Plassan, Paris, cxlvij + 532 pp.

Lundberg, F. 1872. Bidrag till öfversigt af Sveriges Ichthyologiska literatur. Akademisk afhandling med vidtberömda filosofiska fakultetens i Upsala tillstånd för Filosofiska Gradens erhållande till offentlig granskning framställes af Fredrik Lundberg Filos. kand. af Westmanl. Dala Landskap, å Zoologiska lärosalen, Lördagen den 25 Maj 1872, p.v. t. f. m. Stockholm Sigfrid Flodins boktryckeri. xviii+52 pp.

 

The office … ichthyology version

Natural History Museum

The Natural History Museum, London. Dome of sacred saloons of science

Marvel at those majestic buildings harbouring the biological heritage of nations. Natural history museums. Other research collections. The millions by the thousands of corpses and skins, leaves and stems, rocks and gems. Wondering about the shadows cast from time to time across the occasional window lit night after night. Dinosaur or Man, ghost or guard? Strolling through the galleries, what’s behind all those doors that remain locked? More treasures or just the junk? It is not so straightforward to explain what goes on in research collections, and difficult to imagine up. But for sure, there are collections safe in store rooms. And there are the scientists and the collection care staff. And, of course, some administrators. There is always something you can see of collections, and there are the reports, the scientific and not so scientific papers and web presentations, so nothing is really hidden.

Swedish Museum of Natural History. Magnificiently shelled science shelter

But have you ever had a glimpse of a professional abode therein? Is it roomy or squeezy, white-walled or padded with trophy heads? Is there always a Larson in lieu of less intelligible art? Coke or coffee? White coats or tees? When I was so much younger than today I had an idea but not all the imagination. Then, indeed a long, long time ago, I was led by Gordon Howes through a maze of corridors and through the one locked door after the other to arrive at the magical heart of the fish division of the British Museum (Natural History) – now known by a lesser name – and there it was, the air, perfused by alcohol fumes, the books, the microscope steady on the bench, and the uncmfrtbl (so they pronounce it) chair, the tall windows and the big men, books, books, and reprints, reprints, greasy jars, soft dead fish with their autographs helped to them by the wisest of ichthyologists, Günther, Boulenger, Regan. Enchanted, it was a revelation of what life ahead was to be like. And now it is all gone, all the smell, the patina, the deoxygenated atmosphere, the dirty windows, and the kind of yellow Wild microscopes. Everything is new and shining and the nervousness – or was there any – about that spark that would send the alcohol to flames and the building to a cloud of dust, it is no longer there, fire regulations everywhere. And all the other museums are going modern as well. Actually I like the new style too – the facelift is an expression of value and respect for the scientists and their working material. As I much later came to my museum in Stockholm, it was just like that, dirty, dark, dull, and a bit dumb. Now it is full of fresh fish, gas driven chair seats, top-end computers, motorized microscopes, and all the papers are becoming pdfs. As I started on this essay, it was because I were soon to move to new quarters, smaller, newer, and the present, acquired coziness would be part of history. I have been in this office since … 1980? and it was upgraded only once with a little paint and new floor. Since nothing of the classical ichthyological laboratories of the overladen, all-inclusive kind was saved elsewhere (or ?… challenge me!) I decided to photo-freeze a bit of ichthyological history, speaking for many a demolished scientist’s office as new times have moved in. End of the commercials, take a seat and enjoy my research workshop, something like six by three meters and the ceiling truly up in the sky almost. If you ever wondered what a classical fish researcher lab looked like a late afternoon in the winter of 2012, here are all the details (well, a good part of them). If you came upon this text my daily practice unbeknownst, you may wish a confirmation that I am a fish systematist – there it is.

What you see to left and right, front and back, upp the walls and on cabinets and desks – are books and reprints. A sine qua non for life and science alike. The books and reprints in my office are those that are required for ongoing projects and such that are needed for various office tasks such as identifying fish for the public, colleagues and whoever calls. Books that are needed for finding information fast. Highlights are of course the Scott Liddell Greek lexicon, and Erik Wikén’s fabulous Latin for botanists and zoologists (Latin för zoologer och botaniker), in Swedish. I certainly need that copy of Artedi all the time, and right now I am trying to speed learn about Tanganyika cichlids from Günther and Boulenger to Poll and a massmess of molecular writings. If the hand library fails, topping it is the department library which has one of the best collections of fish books, journals, and reprints. Would it not suffice, there are of course AnimalBase and the Biodiversity Heritage Library.

Reprints used to be the blood running through the veins of ichthyology, and here they are still running across the walls.  Authors always procured a hundred or two of each of their publications and distributed them for free.  The point with reprints was that one would not need to subscribe to a journal, and one would not need to page through a whole volume of articles to find tiny scraps of information. Consequently, every footnote and misspelling was elegantly served by reprints, and so taxonomist could indulge in nitpicking ad libitum. It is only in taxonomy, in all of scientific publishing, that people comment on each other’s misprints, yes? The pdf option and Open Access have now killed the reprint collecting efficiently, but that’s all right as long as the printing errors remain …

It takes two (actually three) computers to keep things running, and all data are saved in paper format in three filing cabinets, a dossier for every species and dossiers for other data. Two microscopes may seem overkill, but one is dedicated for photography and one for fish examinations. So one can work in parallel. You will not find many jars in the study, because of fire regulations. Only lesser amounts of ethanol are permitted in offices those days, and the jars needed for the time being are assembled in trays on a cart for convenient transport back to the store rooms.

Stop the presses! Miraculously, this installation still remains. By a lucky chance the move was cancelled at the very last minute, and consequently change is not going to play with my order. Not now. I am happy. What about you? And, what’s your office like?

 

My office

Looking in from the corridor

Northeastern corner, journal shelves

lab bench

Midroom bench, all at hand

Lexicons

Reference books

Within arm’s reach

book shelf

Standing, leaning, lying, a diversity of books

File cabinets

File cabinet

A file for every species …

Microscope

Microscope for imaging

Imaging in action

Canned fish in trays

microscope staring at you

(Is this microscope staring at you?)

Photos: Sven Kullander, CC-BY-NC

Northern glowlights

My kitchen window

My kitchen window

Winter in Sweden, and that means darkness, cold, and snow covering all and everything. No wonder every window is lit, day and night, with glowing stars, moons, snowflakes, menorahs, or the new fashion little reindeer or bears lit by led from inside. Without all those warming winter lights darkness would bend our backs, and we would get swept away by depression. Or maybe not.

Nevertheless, I would like to take this window view to remind myself of another northern glowlight, recently named Dano flagrans. It is a little fish from warmer waters. From where it hails, however,  you can actually view the snow of the eastern outcrops of the Himalayas. It is certainly the most septentrional of the Myanmar Danio, but rivalled, apparently defeated in northerliness by Danio dangila which occurs in the Brahmaputra basin in India up to the Dibru River. No other species of Danio reaches so far north.

Looking northeast from Putao

Looking norwest from Putao

The scientific history of Danio flagrans begins in 1988, when I, in the company of Ralf Britz and our guide Thein Win arrived in Putao, the northernmost major city in Myanmar. Putao is close to the Chinese and Indian borders, on hills forming the headwaters of the Mali Hka, major tributary of the Ayeyarwaddy River which then runs through Myanmar as one big muddy aorta. Up in the Mali Hka, however, the water is clear, at least in the dry season, not very deep, and the river beds paved with stones and rocks. The fish fauna of northern Myanmar mountain streams is little known. Transportation in the area is relatively complicated, and a lot remains to be done up there in terms of ichthyological exploration.

Mali Hka in Putao

Mali Hka in Putao

Back to our story, our little team was quartered in the military camp and we immediately set out to fish, having only two full days at disposal. The Mali Hka itself was too big for fishing, although alright for sightseeing, but around the regiment there were several small streams with low water and convenient for seine and handnets. The streams were shallow, the water was clear, rather cool, and fish were plenty. Here we found Badis pyema which was promptly described already in 2002, and Puntius tiantian in 2005, but other fish have lasted longer to be worked up. Walking along one of the streams, we switched direction to follow a tickle of water, almost no water, coming down the left bank hill, and in there were little skittish fish, almost invisible against the beige earth and seen only as moving shadows. A number of them, certainly Danio choprae – such was the field identification – came into formalin and one made it to a tube of alcohol. Neither Ralf nor I was into danios at the time, so the fish we just hoped would be useful for Fang, and  we went on happily, catching Badis pyema and similar fish that had more of our attention those days.

Fresh collected northern glowlight danio, Putao 1998

Fresh collected northern glowlight danio, Putao 1998

One of the Putao danios was photographed but this was in times of film photography, with no immediate quality check, and much is to be regretted by the quality of the shot. Publishable it is not, but here it can be showed off as the first image ever taken of a Danio flagrans. The alcohol specimen was sequenced and appeared in a phylogenetic tree as Danio choprae (Fang et al., 2009), and by that time noone had looked at it closely (we had other specimens, true D. choprae for the morphological data). Time passed on. This was one species that Fang never worked on, but which obviously was somewhat different from the other samples of D. choprae, and I decided to give it a go in the Spring of 2012. The manuscript was already in hand as I again met Ralf in Belgium and we spoke about past achievements and plans for the future. As he had more of the danios from Putao from a later trip, and more D. choprae, he insisted that I include this material, and so it was. The paper had to be done from almost scratch but Ralf’s material certainly improved a lot on the description and conclusions. The description of Danio flagrans, the northern glowlight danio, eventually appeared in late 2012, 14 years after its discovery (Kullander, 2012). Incidentally, it is my first own danio paper, and it was fun to do. It was enjoyable in particular, because Danio flagrans and its sister species Danio choprae do not differ only in colour (in fact they are very similar in colours), but also present some very solid morphometric and meristic differences. I am otherwise much too used to cichlid species that differ by just some pigment spot. Danio flagrans has a shorter anal fin, with less fin-rays, and longer caudal peduncle compared to Danio choprae. Perhaps this relates to their environment. Danio choprae lives more to the south, near Myitkyina, and in warmer habitats; Danio flagrans in cool hillstreams. Beware that these species may not be correctly identified in the shops. Danio choprae, the glowlight danio may appear in the market as northern glowlights, a more expensive fish. I know, three of the false northerns are swimming in a tank in my garage. These changelings are beautiful fish decorated with orange stripes. Unfortunately, they never stay still, but are constantly on the move, and they move fast, so a good view of them remains an illusion of expectation. This brings me, by association, to the conclusion of this post: Besides lights in the windows, there is one more resource to overcome winter gloom. An aquarium with beautiful fishes (all fish are beautiful). Always something to see, to learn, to enjoy.

aquarium800

References

Fang, F., M. Norén, T.Y. Liao, M. Källersjö & S.O. Kullander. 2009. Molecular phylogenetic interrelationships of the South Asian cyprinid genera Danio, Devario and Microrasbora (Teleostei, Cyprinidae, Danioninae). Zoologica Scripta, 38: 237-256.

Kullander, S.O. 2012. Description of Danio flagrans, and redescription of D. choprae, two closely related species from the Ayeyarwaddy River drainage in northern Myanmar (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters 23: 245-262. Open Access PDF from Verlag Dr. Friedrich Pfeil.

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. 2005. Two new species of Puntius from northern Myanmar (Teleostei: Cyprinidae). Copeia, 2005: 290-302. Open Access PDF.

Photos: Sven O Kullander, CC-BY-NC

The snail, the snake, the frog, the toad, and now the fish

Yesterday’s discoverers are forgotten, faded to oblivion, erased from their maps. As I ask the students, do you know Rolf Blomberg’s books? They stare bufoed, but that’s not an imitation of the gaze of the giant toad discovered by Rolf Blomberg, Bufo blombergi. It is the gaze of the blankness of mind. Too much information around, and too much gets lost. How small our world is, that of travelling biologists and likes, traversing the world in pursuit of dreamed discoveries of new exciting animals or plants, new lands full of things to know and name. It has come to almost nothing and all the thorny paths of the past are paved. Why remember that transatlantic flights were unthinkable just two generations past.

Rolf Blomberg was born into a family residing in Stocksund, just a short bike-ride from the Swedish Museum of Natural History.  It was in 1912, 11 November, in times of relative peace and a relatively orderly Swedish society . The new building of the Museum, at the northern end of the experimental field, was up and running, although not complete until 1916, and young Blomberg became a frequent visitor. Crowding up with loads of stuffed skins, dried bones and spirited fish, however, wasn’t on the agenda for the future. Only 17 he took a job as ship hand, and only 22 he was on his life’s endless journey landing him first in the Galápagos, and eventually taking him around the world  in the quest for the undiscovered, for the great adventure, in a time when everything was already discovered. Although familiar with Africa and Asia, he always returned to the rain forests of the Amazon and the trails of the treasure maps. Blomberg eventually settled in Quito, Ecuador, to become an old man never ceasing to dream of another adventure or the gold of El Dorado. He died in 1996 in Quito. Elderly Swedes, less and less of them, will mostly remember him for his jungle books and films, spiced with exoticism and anacondas, but yet important documentaries from now lost worlds. Others for his engagement in human rights, born out of his observations of the miserable social conditions and political alienation in which he encountered ethnic groups during his travels, particularly in the Amazon, but also extended to protesting the Viet Nam war in the 1960s. In Ecuador his name lives on. There is a good website at Archivo Blomberg with many of his photographs. The English Wikipedia has basic information, also carried by the German, but the Swedish almost zero.  But, after all, he is not quite overboard in Sweden either: Not a little dose of nostalgia and substantial admiration for the explorer was manifested recently in a comprehensive biography by journalist Walter Repo (Repo, 2011), who  also keeps a blog featuring blombergiana of all sorts, rolfblomberg.se. In Swedish. Let’s hope the book gets translated for the rest of the world.

Photo of NRM21169 Chelonoidis nigra

Galápagos giant tortoise Chelonoides nigra, collected by Rolf Blomberg (NRM 21169). Photo Sven Kullander, CC-BY-NC.

Blomberg collaborated with  several museums and systematists. The museums in Gothenburg and Stockholm possess numerous specimens preserved in ethanol, and particularly noteworthy there are some outstanding mounted specimens of Galápagos tortoises and iguanas.

His collecting resulted in four species being named after him. The most spectacular must have been the giant frog Bufo blombergi Myers & Funkhouser, 1951, now often seen as Rhaebo blombergi. Phyllomedusa blombergi Funkhouser, 1957, is a synonym of Phyllomedusa vaillantii Boulenger, 1882, a handsome little tree frog, dubbed white lined leaf frog in spaced English. Bulimulus blombergi Odhner, 1951, now Naesiotus blombergi, is one of so many land snails in Ecuador. Most colourful may be Boa annulata blombergi Rendahl & Vestergren, 1941, now Corallus annulatus or – for us who shun trinomina – Corallus blombergi, which despite its associative name is not a coral snake but a small non-venomous boid snake.

Now, 100 years after Rolf Blomberg was born, it seems pertinent to add another name to the list, because he also collected fish and the fish collections distributed in the museums of Gothenburg and Stockholm have rested magically untouched for much too long. The species Andinoacara blombergi Wijkmark, Kullander & Barriga (2012), is a handsome fish which is known for sure only from the Esmeraldas drainage, the river of emeralds, on the Pacific versant of Ecuador. Some old specimens collected by Manuel Olalla are labeled with a locality in the more northern río Santiago, where it has not been found again, and some that Blomberg got from Ramón Olalla have the locality río Pucayacu, in Amazonian Ecuador. The latter locality is most certainly in error. Mistakes happen. Specimens collected by Blomberg in the río Blanco, one of the main sources of the Esmeraldas, are, however, included in the type series.

Andinoacara blombergi, the holotype, MEPN 11180. Photo by Nicklas Wijkmark, CC-BY-NC.

Andinoacara blombergi is very similar to A. rivulata, and has been confused with it for all of the existence of the latter, but it is more slim and with higher meristics.  Andinoacara rivulata is a common species in the Guayas and Túmbes drainages in southern Ecuador and adjacent Peru. Everything taxonomic about Andinoacara blombergi is available by open access, so it might be a better idea to read there than to search for the same information here.

The description of A. blombergi is based on the work of Nicklas Wijkmark as a Masters student under my supervision, presented in 2007. Seven years ago. Things take time. Nicklas actually made a revision of the whole genus Andinoacara, and more papers are in the tow. Nicklas has since attended to other career opportunities. One of his talents is photography, in which he excels in images of life in wild waters, close-ups of little things, and panoramas of the open landscape. Just sit down with a cup of something and cklick slowly through the marvellous photos at Wijkmark Photography.

Rolf Blomberg lived for travelling and by publishing. He wrote numerous articles fror magazines and newspapers, Swedish and international, mainly about his travels. He made numerous public presentations, and produced alone or together with Torgny Anderberg several documentary or semidocumentaty films for television or cinema. His intellectual legacy is embodied mainly by his books, many of them translated to several other languages, the first in 1936, the last exactly 40 years later:

  • Blomberg, R. 1936. Underliga människor och underliga djur. Hugo Gebers Förlag, Stockholm
  • Blomberg, R. 1938. Högkvarter hos huvudjägare. Hugo Gebers Förlag, Stockholm
  • Blomberg, R. 1940. Underliga människor och underliga djur. Gebers Förlag, Stockholm

    Cover of Blomberg's book Underliga mäniskor och underliga djur

    Front over of Blomberg’s book Underliga mäniskor och underliga djur, 1953 edition

  • Blomberg, R. 1947. Sydvart. Gebers Förlag, Stockholm
  • Blomberg, R. 1948. Nya Smålands upptäckt. Gebers Förlag, Stockholm
  • Blomberg, R. 1949. Vildar.Gebers Förlag, Stockholm
  • Blomberg, R. 1951. Såna djur finns. Gebers Förlag, Stockholm
  • Blomberg, R. 1952. Ecuador. Gebers Förlag, Stockholm
  • Blomberg, R. 1956. Guld att hämta. Gebers Förlag, Stockholm
  • Blomberg, R. 1958. Xavante. Gebers Förlag, Stockholm
  • Blomberg, R. 1959. Jätteormar och skräcködlor. Gebers Förlag, Stockholm
  • Blomberg, R. 1960. Latitud 0°. Almqvist & Wiksell/Gebers Förlag, Stockholm
  • Blomberg, R. 1962. Äventyr i djungeln. Folket i Bilds Förlag, Stockholm
  • Blomberg, R. 1964. Människor i djungeln. Gebers Förlag, Stockholm
  • Blomberg, R. 1965. Mina tropiska öar. Gebers Förlag, Stockholm
  • Blomberg, R. 1966. Rio Amazonas.  Gebers Förlag, Stockholm
  • Blomberg, R. 1967. Imbabura – bergsindianernas land. Gebers Förlag, Stockholm
  • Blomberg, R. 1973. Bufo blombergi. Iskry, Warzawa
  • Blomberg, R. & A. Lundkvist. 1973. Träd. Bokförlaget Bra Böcker, Höganäs
  • Blomberg, R. 1976. Tropisk utsikt. Bokförlaget Bra Böcker, Höganäs

 

References
Repo, W. 2011. Folkhemmets äventyrare. En biografi om forskningsluffaren Rolf Blomberg. Atlas, Stockholm, 335 pp. ISBN 978-91-7389-380-0
Wijkmark, N., S. O. Kullander & R. Barriga S. 2012. Andinoacara blombergi, a new species from the río Esmeraldas basin in Ecuador and a review of A. rivulatus (Teleostei: Cichlidae). Ichthyological Exploration of Freshwaters, 23: 117-137. Open Access PDF from Verlag Dr. Friedrich Pfeil.

The artist with the golden guppy

Tor Otto Fredlin photograph

Tor Otto Fredlin. Photo published in Akvariet, 1932. Unknown artist. Public domain.

In my mildly energetic aspirations to acquire a full set of the Swedish aquarium magazine Akvariet (1927-1988), I was fortunate to add recently the 1932 volume, which turned out to contain  several highlights. This volume, the October issue contains a rare description of a new fish by Wilhelm Schreitmüller, naming it after the Swedish painter Tor Otto Fredlin. It is not cited in the Catalog of Fishes and Google doesn’t know about it yet, so it may be worth commenting on.

Tor Otto Fredlin was born in north Sweden, in the town of Härnösand, 16 June 1890. After school he tested different professions and among other things assisted a a taxidermist. His inclination seems to have been toward painted more than real animals, however. He studied painting at Althin’s School of Painting (in Swedish: Althins målarskola) in Stockholm 1910-1912, and his first exhibition was in Härnösand in 1915. He moved to Lund in southern Sweden in 1920, and stayed there for the rest of his life. Fredlin never married, and he made only one trip abroad, to study art in France in 1924.  As an artist he mainly painted nature settings with animals in focus, mostly birds, but also colourful landscapes. Most paintings seem to have been aquarels and oil paintings of small size.

Akvariefiskar, from Djurens värld 1939, 1948. Artist T.O. Fredlin

Fredlin’s colour plate Akvariefiskar I in Wallengren & Hanström, 1939, 1948, Djurens värld, Fiskar. Actually a composite copied from other illustrations not in copyright.

On the fish side, he made the colour illustrations for the encyclopedic Djurens värld (Wallengren & Hanström, 1937-1940, reprinted 1948), at least one of them depicting fishes he might have had in his own aquarium. Yes, he was also a fish keeper and fish breeder, an aquarist. I cannot find much about Fredlin as an aquarist (or even as an artist), but an idolizing article by the signature Bej in Akvariet, 1932, may be cited in part:

!… one of our country’s greatest aquarium friends. … It is Tor Otto Fredlin, the artist, “the nearsighted nature observer” as an art critic once so characterisingly expressed it. Because our friend Fredlin is a nature observer as few others! He sees everything i nature. Some things even that are secluded from us normal people. … He has the true ability to decorate his containers as cosy and attractive. It is always a pleasure to visit him and hear him speak about and describe his animals. … He shared generously from his rich knowledge. When this was so distributed in the timid way that is so particular for Fredlin, one felt his greatness and was impressed. Already in 1926 he had in addition to many egglayers a remarkably beautiful stock of mirrored golden platys, cultivated by himself. It was lost during a trip up to Norrland. But he did not despond. A lot was still in the wait. He observed our commonest fish, the guppy. Wasn’t there something to do? Couldn’t one get something exceptional out of it? And thus the thought of the yellow guppy that is now a reality had run up to his brain. Who else among us would have had the patience then to spend 7 whole years with this “inconspicuous” fish? It had to be a Fredlin! [The translation attempts to preserve some of the spirit of the original.]

Breathtaking praise. Fredlin must have been a star among aquarists. More sober, but still generously appreciative was Edvin Brorsson’s (1956) obituary. Obviously Fredlin ended his days in the autumn of 1955 as an aquarist in action, among his aquaria in his home in Lund: “He was in the middle of work with cleaning a container as he sat down to rest on a chair, and from here fell dead to the floor.”

His artistic career apparently was not rocketing at any time. His paintings still sell, at modest prices, but let us remember him primarily for painting a grey fish golden, as close to an alchemical GMO one can come. One would expect Fredlin to have painted his golden guppies also in oil, but I am unaware of a colour illustration. Brorsson (1942) reproduces a drawing in monochrome halftone, presumably of a colour painting. It is a typical Fredlin scene, with half the painting empty, and the subject hovering in front of vegetation. Like Fredlin’s fish illustrations in Djurens värld the drawing lacks artistic quality. Many or all of his fish drawings seem to be exact or crude reproductions of photographs. In the colour plate shown here, the Rivulus urophthalmus (yellow top fish) and Hemigrammus ocellifer (group of greys to the left) are exactly as on photographs in Innes (1935, pp. 112, 243), even in number and position, even the background has been copied. Presumably living aquarium fish weren’t within the artistic nearsightedness of the master.

I am ignorant enough about guppies to not know if the golden guppies in the aquarium shops today are descendants of the Scanian breed. It seems likely that pigment mutations of later days may have taken the place of Fredlin’s creation. Nevertheless, at the time, this golden guppy was regarded as something very special. Edvin Brorsson, publisher of Akvariet, sent a batch of living goldens to Wilhelm Schreitmüller in Frankfurt, editor of the Wochenschrift für Aquarien- und Terrarienkunde, and author of several books on aquarium fishes. Schreitmüller responded with an article describing the goldens as a new “variety”, naming it for Fredlin, which in its entirety reads like this when translated from Swedish to English:

Lebistes reticulatus (Peters) var. Fredlini (Schreitm.)
By Vilhelm Schreitmüller, Frankfurt am Main

A short time ago, Mr Brorsson, Malmö, sent me a fish can with some golden yellow Lebistes reticulatus (Peters), millionfishes or Guppi), along with a message that these were produced in culture from the typical and natural form by a Swedish aquarium friend, Mr T. O. Fredlin, over a period of 8 years, and that the fishes nowadays leave a stable offspring. In shape those animals are typcal Lebistes reticulatus (Peters). The males have a length of 2-2.5 cm., the females 3-3.5 cm., all including caudal fins.

On the back, the males are dark yellow, while hindbody and caudal pedcuncle are light yellow, the latter with 2 vermilion round spots, in front of which are placed a dark and a copper red spot. One of the males sent along has a dark, white-margined dot at the base of the caudal fin. The scales on the anterior par tof the back have a faint dark margin.

The females are orange yellow on the back and on the entire hindbody with caudal peduncle. The pregnancy spot is pink red. The scales on the anterior part of the back are finely dark margined. The abdomen is yellow white.

We thus have to deal with heare typical xanthoristic specimens, whose fins are transparent and seem to be equipped with a yellow cast. Pectoral and pelvic fins’s base is reddish.

In recognition of the breeder and to differentiate these animals from the typical Lebistes reticulatus (Peters), I call this fish:“Lebistes reticulatus (Peters) var. Fredlini (Schreitm.)”
even if xanthoristic forms “actually” should not have a distinct name (usually!) – But that even the professional zoologists make exceptions in this regard is apparent best from the names below:

“Carrasius auratus (L.)” = goldfish,
“Tinca aurata (Cuv.)” = guldsutare,
“Cyprinus auratus (Mats.)” = gold carp,
“Platypoecilus immaculatus (Myers)” = gold platy,
and many others, which all of them also are only xanthoristic forms of the species in question.

In the same manner certain melanotic forms of lizards, e.g., “Lacerta lilfordi maluqueurorum (Mertens)” and several others receive their particular variety names.

So – why should one not then be able to give the xanthoristic form of Lebistes reticulatus (Peters) a proper variety name? “Was dem Einen recht ist, ist dem Anderen billig”!

Food, care and water temperature is for the new guppy the same as for the original species.
(Frankfurt am Main 10/10  [Italics as in the original, parentheses as in the original. PDF of original]

A similar text, in German, was published in the Wochenschrift in 1933 (Schreitmüller, 1933). Schreitmüller’s  description does not make the name fredlini available for purposes of zoological nomenclature. On the one hand,  the International Code of Zoological Nomenclature covers names of domestic animals (e.g., Canis familiaris for dogs), but is also expressly excludes names proposed for varieties after 1960. For names of varieties proposed before 1961, Articles 45.6.1 and 45.6.4 apply, and they exclude names given to what obviously is not a subspecies except if a later citation exists, before 1985, that uses the name for a valid species or subspecies. This may be worth looking into, but for now, fredlini is not going to lengthen our lists of guppy synonyms, as the name was applied explicitly on a colour form.

Fredlin’s golden guppy is not the only one, and not the only one named by Schreitmüller. Already in 1934 Schreitmüller  reported on specimens of “Goldguppy” sent alive to him and originating from German breeders in Czechoslovakia (Schreitmüller, 1934). He compared them with the Fredlin guppy, noting that the Czechoslovakian guppy lacked spots on the side and fins, and that the Fredlin guppy was more intensely golden yellow. Schreitmüller named the new form Lebistes reticulatus aurata, noting that: “Dieser name soll nur als Unterscheidungsmerkmal dienen.” [This name must only serve as distinguishing character.] One may interpret this as meaning that Lebistes reticulatus aurata was intented as just a category stamp and not as a scientific name. Seven specimens are preserved in the Zoological Museum in Berlin, and were listed as syntypes of of Lebistes reticulatus aurata by Paepke & Seegers (1986), so obviously these authors considered Schreitmüller’s name available, although they identify the specimens as Poecilia reticulata.

In a later issue of the Wochenschrift, Franz Melecky, from Kremsier (Kroměříž, in the present Czech Republic), described how he discovered golden guppies in his plant breeding compound, propagated them and started distributing Goldguppy as Lebistes reticulatus aurata from 1925 onward. Interestingly, Melecky uses the name Lebistes reticulatus fredlini at the end of his story, speculating if a cross with the Fredlin guppy could improve on the size of his own Goldguppy, as they didn’t reach the sizes of the original stock. Does this make fredlini available? Or is this the end of the story?

References

  • Bej. 1932. Känd akvarieentusiast. Akvariet, 6: 39–40.
  • Brorsson, E. 1942. Den stora akvarieboken. Andra, reviderade upplagan. Sundqvist & Emond, Lund, 300 pp.
  • Brorsson, E. 1956. T. O. Fredlin. Akvariet, 30: 47.
  • Innes, W.T. 1935. Exotic aquarium fishes. Innes Publishing Co., Philadelphia, 464 pp.
  • Melecky, F. 1934. Goldguppy (Lebistes reticulatus aurata). Wochenschrift für Aquarien- und Terrarienkunde, 31: 427.
  • Nyman, T. 1944. Fredlin, Tor Otto. P. 592 i Bohman, N. (ed.), Svenska män och kvinnor 2. Albert Bonniers Förlag, Stockholm.
  • Paepke, H.-J.  & L. Seegers  1986 Kritischer Katalog der Typen und Typoide der Fischsammlung des Zoologischen Museums Berlin. Teil 1: Atheriniformes. Mitteilungen aus dem Zoologischen Museum in Berlin, 62: 135–186.
  • Schreitmüller, V. 1932. Lebistes reticulatus (Peters) var. Fredlini (Schreitm.).  Akvariet, 6: 118–119.
  • Schreitmüller, W. 1933. Neuimporte und anderes. Hyphessobrycon bifasciatus Ellis, Cichlosoma cutteri Fowl., Colossoma species, Hyphessobrycon species I und II und Lebistes reticulatus (Pet.) var. fredlini (Schreitm.). Wochenschrift für Aquarien- und Terrarienkunde, 30: 145–149.
  • Schreitmuller, W. 1934. Der “Goldguppy” und ein Totalalbino von Xiphophorus hellerii Heckel. Wochenschrift für Aquarien- und Terrarienkunde, 31: 242–243.
  • Wallengren, H. & B. Hanström (eds.). 1939. Djurens värld. En populärvetenskaplig framställning av djurens liv på grundval av Brehm’s Tierleben utarbetad av Ingvald Lieberkind. Fiskar Band II. Svensk uppslagsbok, Malmö, 439 pp.
  • Wallengren, H. & B. Hanström (eds.). 1948. Djurens värld. En populärvetenskaplig framställning av djurens liv på grundval av Brehm’s Tierleben utarbetad av Ingvald Lieberkind. Fiskar Band II. Förlagshuset Norden, Malmö, 439 pp.

That Africke evermore bringeth forth some new and strange thing or other

Caius Plinius secundus. Image from Wikipedia, public domain

Ex Africa semper aliquid novi, “From Africa always something new”, is a well-known expression, applied whenever something is discovered in Africa. It is usually attributed to Caius Plinius secundus (23-79), in his Natural History (Naturalis Historiæ).

In my copy of Plinius (1536-1538), the proverb is in Book VIII, chapter XVI, about lions, where he remarks on the diversity of animals in Africa: “Unde etia vulgare Græciæ dictum, Semper aliquid novi Africam affirre.” (Approximately: “Hence there is a common proverb in Greece, that there is certainly Always something new of Africa”). There are many printed editions of Plinius, and apparently variations in the divisions of the text and the exact wording. In the index of my copy one finds: Africa semper aliquid novi affert, i.e. Africa always brings something new. Personally, I prefer this variant above the others,  both in Latin and English.  In another edition Plinius is cited as: “unde etiam vulgare Græciæ dictum semper aliquid novi Africam adferre“. In an early translation the proverb is cited as “That Africke evermore bringeth forth some new and strange thing or other” (Holland, 1601). In a later translation (Bostock & Riley, 1855), Plinius is cited “Hence arose the saying, which was common in Greece even, that ‘Africa is always producing something new.'”

Since Africa wasn’t known by that name to the ancient Greek, and Plinius drew heavily from earlier authors, not least Aristoteles Stagirites (384-322 BCE), it may not be surprising that the proverb seems to come straight from the Greek and from Aristoteles’s Historia Animalium ( Περὶ Τὰ Ζῷα Ἱστορίαι), Book 8, part 28: Ἀɛὶ Λίβύη ϕέρɛί ṯί καίνόν. Thompson (1907) translates the relevant text as: “As a general rule, wild animals are at their wildest in Asia, at their boldest in Europe, and most diverse in form in Libya; in fact, there is an old saying, ‘Always something fresh in Libya.'” Aristoteles did not mean the present-day country of Libya, but Libya at the time should here be understood as northern Africa excluding Egypt.

When it comes to biodiversity we know better now, as tropical Asia and even more tropical South America are better places for searching animal diversity, but the world of Aristoteles and Plinius was essentially restricted to the Mediterranean and immediate neighbourhood. True, though, is that the assortment of large-sized vertebrates is much more conspicuous in Africa. And, certainly, new organisms are still being discovered also in Africa.

When I first became interested in fishes, African cichlids dominated the aquarium world. This was in the late 60s early 70s and I lived in a small town far away in the north of Sweden. With some other fanatics I imported Malawi cichlids from Germany and made occasional visits to Stockholm, capital of Sweden, to watch more expensive Tanganyika cichlids. Those are memories for some other time, but the consequence of the devotion was that I spent considerable time copying and deciphering the French in the monumental work of Poll (1956), Exploration Hydrobiologique du lac Tanganika. Poissons Cichlidae; and published some articles on African cichlids in Buntbarsche Bulletin as well as in the journal of the Nordic Cichlid Association (Nordiska ciklidsällskapet).Eventially, I shifted attention to South American cichlids, but in 1990 there was a golden opportunity to make a collection trip all over Zambia, targeting tilapias, but also preserving everything else with fins. Together with my friend Erkki Schwank, it was a long trip by car across the country, to Lakes Mweru, Mweru wantipa, Bangweulu, and Tanganyika. We didn’t stay long on the shores of Lake Tanganyika, but in a couple of days at Nsumbu we obtained a sizable collection of shore and deep water cichlids. Along with collections from an earlier trip by Gunnar Berglund to Kigoma, the Swedish Museum of Natural History now had a useful representation of Tanganyika cichlids, which was further added to by Tyson Roberts and Erkki, who stayed in Zambia.

Much later I was happy to meet underwater photographers and exporters Mikael and Magnus Karlsson who lived by and in the lake for many years and we teamed up to publish their observations on some of the lesser known species in Lake Tanganyika. The first paper is a description of a Lepidiolamprologus from the coast of Tanzania. Lepidiolamprologus includes a core number of species that are relatively elongate and marked by dark stripes or rows of blotches. Strong canine teeth emerging out of the lips and the somewhat oval pupil give them the look of a fierce predator. And predatory they are, and some definitely prey on babies of other cichlid species. These core species are L. elongatus, L. mimicus, L. kendalli, L. profundicola, and maybe L. nkambae. Lepidiolamprologus elongatus is distributed along the entire lake coastline, and L. profundicola may also be widespread. The remainder, and our new species, L. kamambae, are from the south of the lake. The status of L. nkambae and L. kendalli is uncertain, as no distinguishing characters are known. Lepidiolamprologus kendalli was described first, by Max Poll and Donald Stewart (1977), based on two specimens. They reported scales on the cheek and illustrated the holotype with a line drawing. Lepidiolamprologus nkambae was described by Wolfgang Staeck the following year, based on a single specimen from close to the type locality of L. kendalli. It was also illustraded with a line drawing. Staeck compared with the description of L. kendalli and concluded that they were different. Among distinguishing characters L. nkambae was said not have any scales on the cheek. Both names have since appeared in the aquarium literature but neither aquarists nor scientists have really been able to conclude about the validity of L. nkambae. Synonym of L. kendalli or distinct species? Only after our description of L. kamambae was I able to examine the holotype of L. nkambae and the paratype of L. kendalli. The L. nkambae specimen is very well preserved, and agrees with the description. The paratype of L. kendalli, on the other hand, is in a very poor state of preservation, something that was not mentioned in the original description. Scales are lost and the colour is washed out. I will not tell you my decision here on the validity of L. nkambae here, it will be something for a forthcoming paper, but I agree on one of the two alternative conclusions already presented ….

The material of L. kamambae that Mikael and Magnus had, is excellently preserved. Colours are fine, all scales are in place, the body is straight and the fins naturally spread, with no signs of decomposition anywhere, some mouths are a bit open but that makes counting teeth convenient. I like. Lepidiolamprologus kamambae shares most of its features with L. kendalli, in particular the beautiful contrasting light and dark markings on the top of the head, distinguishing the two from all other lamprologins. The body coloration, however, is very different. Lepidiolamprologus kamambae is more similar there to L. elongatus and L. mimicus, with rows of blotches along the back and flanks where L. kendalli instead has broad bands. One may guess that the blotch pattern is the plesiomorphic version in this group of fishes. There is probably not much more to say about this fish here, because the description is available as Open Access from Zootaxa.

Reflections may be in place, however. Lake Tanganyika is the oldest of the three Great Lakes of East Africa.  More than 200 endemic cichlid species have been described from the lake, and estimates suggest a total of 300. Unlike in Lakes Malawi and Victoria, dominated almost exclusively by mouthbrooders of the ‘haplochromine’ group, the distantly related lamprologins, of several genera, all substrate brooders, are a large unit in Lake Tanganyika, with over 70 valid species known so far. Several recent studies have investigated their phylogenetic relationships, and relations to the half dozen species of Lamprologus that only occur in the fluviatile environment of the Congo River (e.g., Schelly et al. 2006).  The latter are obviously a group on their own, but within-lake relationships still offer much to be investigated. I am curious how things will develop, and it is interesting to work with fishes offering this kind of perspective. When it comes to African cichlids, in particular those of the Great Lakes, systematics has investigated them from many different angles, but still is way off from a coherent and credible evolutionary history. There are papers on dried out lakes, explosive radiation, lake level fluctuations, depth segregation, sympatric speciation, not to mention hybridisation. Academic analyses are enough to fill volumes, and it is engaging and important that this unique fauna, evolutionary hotspots with dense concentrations of phylogenetically close, but morphologically often far apart species, will continue to receive attention. Even so, and connecting back to Plinius and friends, the following passage, from the chapter of the famous proverb seem like they may be pertinent, and of course inspired by Aristoteles, in light of some of the ideas about African cichlid evolution, or perhaps not ideas but frustration, read cichlid for lion:

The noble appearance of the lion is more especially to be seen in that species which has the neck and shoulders covered with a mane, which is always acquired at the proper age by those produced from a lion; while, on the other hand, those that are the offspring of the pard, are always without this distinction. The female also has no mane. The sexual passions of these animals are very violent, and render the male quite furious. This is especially the case in Africa, where, in consequence of the great scarcity of water, the wild beasts assemble in great numbers on the banks of a few rivers. This is also the reason why so many curious varieties of animals are produced there, the males and females of various species coupling promiscuously with each other.3 Hence arose the saying, which was common in Greece even, that “Africa is always producing something new.” (Plinius, Naturalis Historiae; translation by Bostock & Riley, 1855)

From this we obviously learn that at any point in time we shall know less than at a later time (especially if we don’t investigate the facts ourselves), and that everything has been thought of before.

References

Bostock, J., & H. T. Riley (eds.) 1855, Pliny the Elder, The Natural History. Taylor & Francis, London.

Holland, P. (translator), 1601. The Historie of the world. Commonly called, The Naturall Historie of C. Plinius secundus. London.

Kullander, S. O., M. Karlsson & M. Karlsson. 2012. Lepidiolamprologus kamambae, a new species of cichlid fish (Teleostei: Cichlidae) from Lake Tanganyika. Zootaxa, 3492: 30-48. Open Access PDF from Zootaxa

Kullander, S. O. & T. R. Roberts. 2011. Out of Lake Tanganyika: endemic lake fishes inhabit rapids of the Lukuga River. Ichthyological Exploration of Freshwaters, 22: 355-376. Open Access PDF

Plinius Secundus, C. 1536-1538. Naturalis Historiae. Aldus, Venetia.

Poll, M. 1956. Poissons Cichlidae . Exploration Hydrobiologique du Lac Tanganika (1946-1947). Résultats scientifiques, III (5B): 1-619.

Poll, M. & D. J. Stewart. 1977. Un nouveau Lamprologus du sud du Lac Tanganika (Zambia). Revue de Zoologie africaine, 91: 1047-1056.

Schelly, R., W. Salzburger, S. Koblmüller, N. Duftner and C. Sturmbauer. 2006. Phylogenetic relationships of the lamprologine cichlid genus Lepidiolamprologus (Teleostei: Perciformes) based on mitochondrial and nuclear sequences, suggesting introgressive hybridization. Molecular Phylogenetics and Evolution, 38: 426-438.

Staeck, W. 1978. Ein neuer Cichlideaus dem südlichen Tanganjikasee: Lamprologus nkambae n. sp. (Pisces, Cichlidae). Revue de Zoologie africaine, 92: 436-441.

Thompson, D. W (translator). 1907. The history of animals. John Bell, London.

Photo of Holotype of Lepidiolamprologus kamambae

Holotype of Lepidiolamprologus kamambae. Photo Sven Kullander

Swedish Fishes

cover of Chordata: ActinopterygiiI was comfortably seated in the sofa,  in a conversation about books bad or good, literature that is. My eyes crawled for cues along the backs of books in the sunlit bookshelf across the room, eventually steadying on a 15 volumes work from the early 1960s that presents the animal kingdom in a phylogenetic sequence and providing some food for thought. It is in Swedish and intentionally popular but probably too advanced for readers in the 21st Century. Departing from the traditional Brehm, Djurens värld edited by Bertil Hanström, (1891-1969), professor in Zoology at Lund University, treats every major group in some detail, with illustrations and facts mostly taken from scientific literature. In some respects a lay version of the monumental work of comparative morphological zoology, the Traité de Zoologie. The first 14 volumes were systematic reviews written by expert biologists, the 15th, treating animal preservation techniques, by Yngve Lövegren (1899-1901), an outstanding specialist in animal preservation and the history of Swedish museum collections. Hanström had help from Alf Johnels (1916-2010) with the two fish volumes, of which the first covers a significant part of vertebrate anatomy and physiology. These volumes descend from a period when there was no Internet, not even fax machines. A time when communication means were by print, letters (aka snail mail), and telephone. Libraries, personal or institutional, were the main source of information and copies were made by photostats or photographs, but largely by studying in the library and taking notes. Comparing with today’s mountains of readily accessible information, printed, online, or xerographed, and the ease by which intelligeble text can be produced just by a little copying and pasting, one has to be respectful of the writings of the not so distant past. So, Hanström and his peers need a special honorable mention here, before we come to the point of this text, which is more specifically about monographs or faunal reviews of Swedish fishes. The two fish volumes of Djurens värld played a significant role mostly as an inspiration for becoming fish taxonomists, and the inspiration to write this report, possibly shamelessly promotional and an exercise in name dropping. But I can’t just write “hey there, there’s a new fish book, and it is really nice…” Or maybe that would have been enough…

The earliest post-Linnaean attempt to fully cover the Swedish fish fauna in a single volume was probably by Bengt Anders Euphrasén, but his manuscript was never published. It was not until 1836 that the first portions of the intended first complete, illustrated monograph on Swedish fishes appeared, Skandinaviens fiskar. It was authored by curators at the Swedish Musuem of Natural History, Bengt Fredrik Fries (1739-1839) and Carl Ulric Ekström (1881-1858), with illustrations made by Wilhelm von Wright. Fries died in 1839 and Carl Jakob Sundevall (1801-1875) stepped in as a contributor. The work was meticulous, but slow, and the last few pages of this terminally unfinished work were published in 1857.

In 1892, however, it came to life again with the first volume of two of a much reworked version with the same title, but considerably improved and exmpanded by Fredrik Adam Smitt (1839-1904), professor at the Swedish Musem of Natural History. It is a magnificent, heavy volume of 1239 pages, accompanied by colour litographs. Volume 2 came in 1895 and the plate volume was complete also in 1895. An English translation, Scandinavian fishes, also  appeared in 1895. In the meantime Sven Nilsson (1787-1883) in 1855, then professor at Lund University, presented a full descriptive catalogue in the fourth volume of Sveriges Fauna, but without illustrations. Between 1881 and 1891 Wilhelm Lilljeborg (1816-1908) published three volumes of descriptions of Scandinavian fishes, Sveriges och Norges fiskar  (Sweden and Norway formed a union, The United Kingdoms of Sweden and Norway from 1814 till 1905), but none figured. Just as Smitt’s and Lilljeborg’s books were out,  Anton Stuxberg (1849-1902), curator at the Museum of Natural History in Gothenburg published a faunal compliation covering the Scandinavian fish fauna, Sveriges och Norges fiskar (1894-1895), a very basic work, which profited from illustrations reproduced in grayscale from Smitt.

Especially Smitt’s work is something of a masterpiece, putting together all information available at the time about all fish species occurring in Scandinavia, as a rule with descriptions and figures from specimens in the collection of the Swedish Museum of Natural History, and with a considerable amount of original drawings and descriptive data. von Wrights’s figures were used also for Smitt’s edition of Scandinavian fishes,  but several artists were involved in the production of illustrations of species not covered in the first edition.

An amazing work of later days, published mid-war, 1942, is the treatise of Nordic fishery by Karl Andreas Andersson (1875-1968), MP and director of the Swedish Fisheries Agency. It incorporates descriptions of all Swedish fishes written mostly by Orvar Nybelin, curator and eventually director of the Natural History Museum in Gothenburg. Most species were illustrated, some of them in charming full page color photographs of carefully arranged freshly caught fish specimens. A second edition appeared in 1954, and a third in 1964. Nybelin was well prepared for this task. His pocket guide to Swedish fishes, Våra fiskar, was published in two volumes, one for freshwater and brackish water fishes published in 1933, and one for marine fishes published in 1937. They were followed by improved editions up till 1956, and one usually finds a freshwater and marine book bound to a single volume. In a very compact format this book provided both illustrations (colour and greyscale, mostly from Smitt), and determination keys to all Swedish fishes.

Recent heroes of popular Swedish fish faunal monographs include Kai Curry-Lindahl (1917-1990), zoologist with various positions, who maintained a steady series of books, based to some extent on Danish or Norwegian templates, but with a unique body of information on specificially Swedish conditions. His Fiskarna i färg (1953-1979) was unimpressing, but Våra fiskar (1985) is a classical book that will never completely go away. Unfortunately becoming rare in the used book market, as one would hesitate to part from it. Both Fiskarna i färg and Våra fiskar have been continued by mostly translated works on the Nordic fauna. An important Danish work in several editions, Havfisk og fiskeri i Nordvesteuropa and Europas ferskvandsfisk by Bent J. Muus and Preben Dahlström, have been an important field guides in their Swedish translations from 1965.

So, considering the excellence of the historical works, it is with considerable humbleness that one presents now the second fish volume in the Encyclopedia of the Swedish Flora and Fauna, published 24 September 2012: Chordata: Actinopterygii.  The first volume was mainly about tunicates, but treated also hagfish, lampreys and chondrichthyans. The second volume treatsalmost  all actinopterygians encountered in Swedish waters, and since all are illustrated in colour is is quite a lot more colourful tome than the first volume. The production of the second book has gone on for almost seven years in parallel with the first volume,  and the quality and usefulness of it is largely due to the skills of the illustrators, Linda Nyman and Karl Jilg.

The book covers 216 species, from Acipenser baerii (an introduction) to Mola mola (sporadic). Two species have not been reported before from Swedish waters, namely Lebetus guilleti and Ciliata septentrionalis, but are illustrated here based on Swedish specimens. A third novelty is Carassius gibelio, but it is well known as a specimen game fish since the early 1990s. Incidentally, two new species for Sweden were reported too late to be included in the Encyclopedia, viz. Reinhardtius hippoglossoides and Beryx splendens, encountered in 2011 and reported by Leif Jonsson. A few species, such as the single record of Canthidermis maculata in 1857, did not make it into the book as singularities that have not been observed in Sweden after the 19th Century were outside the concept of the product.

The book on Swedish actinopterygians is contained in a series called, in English, The Encyclopedia of the Swedish Flora and Fauna. The Encyclopedia is a project started at the Swedish Species Information Centre in Uppsala in 2001  and aims to produce a series of identification handbooks with keys in Swedish and English to the Swedish plant, fungi and animal species. It is a long-term project, aimed at covering the 30 000-40 000 species which can be identified without highly advanced equipment. They will be described in detail, including information on distribution and biology. For most of them, distribution maps as well as illustrations will also be provided.

Back to the beginning of this blog, which is the best of all books? I don’t know, but probably not a fish book. It is probably one of the many novels, and maybe none in particular. There is a different book for every moment and every task in life. As long as there are books, there is life.

With this conclusion, it is alarming that a government-appointed  reviewer of the Encyclopedia and other activities at the Species Information Centre, has suggested termination of the Encyclopedia in its printed form. It is not over yet, though, and the Swedish biological community is up to fight for knowledge.

 

References, in chronological order to major faunal field guides and monographs of Swedish (and sometimes Norwegian) fishes

Fries, B. Fr., C. U. Ekström & C. J. Sundewall. 1836-1857. Skandinaviens Fiskar. P. A. Norstedt & Söner, Stockholm, IV+222 pp. Appendices 1-44, 1-140, pls. 1-60. Fascicle 2-3 (1837),  4 (1840)  5, 1839 (p. 111 dated 22 October 1839, ) 6+pls 31-36, Latin text 57-72 (1840), 7+pls 37-42, Latin text 73-92 (1842).

Nilsson, S. 1855. Skandinavisk fauna. Fjerde delen: fiskarna. Bokhandlaren C.W.K. Gleerups Förlag, Lund XXXIV+768 pp. [ Online in Biodiversity Heritage Library.]

Lilljeborg, W.. 1881-1891. Sveriges och Norges fiskar [Also as Sveriges och Norges Fauna, fiskarne]. W. Schultz, Upsala. In fascicles: 1 (pp. 1-208) 1881;  2 (pp. 209-496) 1884; 3 (pp. 497-782) 1884; Andre delen, 4 (pp. 1-416) 1886;, 5  (pp. 417-788) 1888; Tredje delen, 6 (pp. 1-336) 1889; 7 (pp. 337-672) 1890; 8 (pp. 673-830+I-XXI+title pages) 1891. [Online in Biodiversity Heritage Library.]

Smitt, F.A. 1892. Skandinaviens fiskar målade af W. von Wright beskrifna av B. Fries, C.U. Ekström och C. Sundevall. Andra upplagan. Bearbetning och fortsättning. Text. Förra delen. P.A. Norstedt & Söners Förlag, Stockholm, pp. 1-566+I-VIII+2 pp.

Stuxberg, A. 1894-1895. Sveriges och Norges fiskar jämte inledning till fiskarnes naturalhistoria. Wettergren & Kerber, Göteborg, 678 pp. [Issued in four fascicles.]

Smitt, F.A. 1895. Skandinaviens fiskar målade af W. von Wright beskrifna av B. Fries, C.U. Ekström och C. Sundevall. Andra upplagan. Bearbetning och fortsättning.Text. Senare delen. P.A. Norstedt & Söners Förlag, Stockholm, pp 567-1239+1 p.

Smitt, F.A. 1895. Skandinaviens fiskar målade af W. von Wright beskrifna av B. Fries, C.U. Ekström och C. Sundevall. Andra upplagan. Bearbetning och fortsättning. Taflor. P.A. Norstedt & Söners Förlag, Stockholm, pls I-LIII, pp. I-III. [Plates from English edition available at Biodiversity Heritage Library.]

Nybelin, O. 1937. Våra fiskar och hur man känner igen dem. Illustrerad fickbok med korta beskrivningar av deras utseende och levnadssätt, förekomst, fångst och användning. Del I. Fiskar i sött och bräckt vatten. Andra upplagan. Albert Bonniers Förlag, 59 pp.[1st edition 1933, 3rd edition, 1943, 4th edition 1948, 5th edition 1951, 6th edition 1956.]

Nybelin, O. 1937. Våra fiskar och hur man känner igen dem. Illustrerad fickbok med beskrivningar av deras utseende, levnadssätt och förekomst. Del II. Havsfiskar. Albert Bonniers Förlag, 83 pp.[2nd edition 1945, 3rd edition 1951, 4th edition 1956.]

Andersson, K. A. (red.). 1942. Fiskar och fiske i Norden. Band I Fiskar och fiske i havet. Band II Fiskar och fiske i sjöar och floder. Natur och Kultur, Stockholm, XXIV+1016 pp. [Additional editions 1954, 1964.]

Curry-Lindahl, K. 1953. Fiskarna i färg. Tredje upplagan, Almqvist & Wiksell, Stockholm, XVIII+189 pp. [1st edition 1953, 2nd 1954, 3rd 1957, 4th 1961, 5th 1964, 6th 1966, 7th 1969, 8th 1975, 9th 1979.]

Curry-Lindahl, K. 1985. Våra fiskar. Havs- och sötvattensfiskar i Norden och övriga Europa. P.A. Norstedt & Söners Förlag, Stockholm.

Kullander, S.O., T. Stach, H.G. Hansson, B. Delling & H. Blom. 2011. Nationalnyckeln till Sveriges flora och fauna. Ryggsträngsdjur: lansettfiskar-broskfiskar. Chodrata: Branchiostomatidae-Chondrichthyes. ArtDatabanken, Uppsala. 327 pp.

Kullander, S.O., L. Nyman, K. Jilg & B. Delling. 2012. Nationalnyckeln till Sveriges flora och fauna. Ryggsträngsdjur: Strålfeniga fiskar. ArtDatabanken, Uppsala, 517 p.

Honorary listing:

Hanström, B. (ed.), 1960-1963. Djurens värld, en populärvetenskaplig framställning av djurens liv. Andra omarbetade upplagan. Norden, Malmö, 15 volumes: 1, Dahl, E, 1960, Ryggradslösa djur 1, 500 pp.; 2, Brinck, P., Dahl, E. & Hanström, B., 1961 Ryggradslösa djur 2, 536 pp.; 3, Brinck, P., Ryggradslösa djur 3, 1963, 562 pp.; 4, Dahl, E. & Hanström, B., 1963, Ryggradslösa djur 4, 692 pp.; 5, Johnels, A. G. & Hanström, B., 1961, Fiskar 1, 422 pp.; 6, Hanström, B. & Johnels, A. G., 1962, Fiskar 2, 611 pp.; 7, Kauri, H., 1962, Grod- och kräldjur, 606 pp.; 8, Ulfstrand, S., 1960, Fåglar 1, 496 pp.; 9, Ulfstrand, S., 1961, Fåglar 2, 571 pp.; 10, Ulfstrand, S., 1960, Fåglar 3, 607 pp.; 11, Hanström, B., 1960, Däggdjur 1, 571 pp.; 12, Hanström, B., 1960, Däggdjur 2, 528 pp.; 13, Hanström, B., 1960, Däggdjur 1, 415 pp.; 14, Hanström, B., 1960, Däggdjur 1, 462 pp.; 15, Löwegren, Y., 1961, Zoologisk museiteknik, 551 pp.

 

Ciliata septentrionalis

Ciliata septentrionalis, NRM 46235, Gullmarn, 1909

 

 

 

Sirens of Liège

This is the first day of silence after coming back to Stockholm from the XIV European Ichthyological Congress in Liège, Belgium (3-8July).

Aquarium and Museum Liège front

Entrance to the museum and aquarium in Liège, venue of the XIV European Congress of Ichthyology, July 2012

Whereas the meeting, attended by a round 260 delegates from all over Europe and occasional from farther away, was well organised and quite informative, heat hit us and noise. Coming from a cold Swedish summer, the rainiest June in 200 years, Liègian temperatures were an amiable mid-20s and only one shower wet us. But it took buying a few more T-shirts (from Swedish H&M) to feel comfortably neat around the clock. I loved the heat, actually. But the noise… Constructed as an intricate labyrinth of narrow streets between stone and brick buildings, sounds from everything sounding cascaded through the streets of Liège and there was no escape even in the hotel room. Most remarkable, however, were the frequent ambulances and police cars continuously racing in and out of streets and alleys, with a 2000 db (exaggerating, but well …) high pitch sirens resounding all over the municipality. Quite remarkable. Liègians seem to love noise more than anything else.

 

 

Photo of stuffed mermaid in Liège museum

Sirena anthroposelacia pilifera in the exhibit of the Aquarium in Liège

 

 

 

Fleeing the street sirens I had a brief encounter with a real one in the lower floor of the combined public aquarium and zoological museum (Liège Aquarium-Museum) , which was also the venue for the Congress. This little lady had a glass cage all for herself, in swimming pose, and is indeed the first real stuffed mermaid I have come close to in real life.

 

Fish exhibit of the zoological museum in Liège

A part of the systematically ordered exhibit at the zoological Museum in Liège

 

The rest of the aquarium is maybe mainly for the local audience. The Museum, however, two floors up, was neat and had a number of interesting objects, making it well worth a visit. Displaying real animals in systematic order it was ideal and should be a lead star for other natural history exhibits.

 

 

 

 

 

 

Photo of notebook with fish drawings

Pages in notebook by Francis de Castelnau in the exhibition of the Zoological Museum in Liège

One table display featured a charming set of castelnauiana that was news to me, including drawing equipment, notebooks, and sketches from the later days of Francis de Castelnau (1810-1880). Apparently Castelnau donated specimens to the museum along with the notebooks, and there is now a plan to repaint the stuffed fishes based on the drawings.

The Congress? Well, I probably sat more than the usual number of sessions. Belgium has a big advantage in having several institutions providing training and research opportunities in morphology and anatomy, making it an interesting breeding ground for evolutionary and functional analyses of fish physionomies and behaviour, and has also a very strong research in African freshwater fishes. This balanced very well a large number of more or less conclusive or inconclusive molecular presentations. I travelled with NRM FishBase staff Michael Norén and Bodil Kajrup who presented a talk on our ongoing hagfish research, and a poster of a mapping of Swedish fish type localities, respectively. I have been too occupied with other things recently, and did not feel like giving a talk. Some other time. Maybe on mermaids. This is not the last mermaid post. It didn’t turn out a meeting report, at least.

By the way: The European Ichthyological Congresses are organised locally (this one mainly by the University of Liège) but are actually a significant part of activities of the European Ichthyological Society. The first congress was organised in Sarajevo in 1972. I was one of the organisers of the Congress in Stockholm in 1985. Membership in the society is open for all, and there is a website (http://artedi.nrm.se/eis/) providing the details how to join.

 

All photos © Sven O Kullander, CC-NC-BY

Sharks, skates, and Swedish seas

Today is the offcial release day of the 13th volume of the The Encyclopedia of the Swedish Flora and Fauna, dedicated to lower chordates, ie., lancelets, tunicates, hagfish, lampreys and chondrichthyans. It also comes with an introduction to chordates and to craniates, the latter sprawling with colorful dino drawings. Although I am first author, most of this tome is about tunicates with fantastic images from within and without that makes it a particulatly worthwhile reading (and buying, come on it is only SEK 345 and you get the sharks for free!) Thus, tunicate expert Thomas Stach, Freie Universität Berlin, and the late Hans G. Hansson, Tjärnö Marine Biological Laboratory, provided most of the species in this volume. But it also has one hagfish, three lampreys, and 29 chondrichthyans.

What you will like most about this volume is probably the graphics. For the craniates major contributing artists Linda Nyman and Karl Jilg, guided in the intricate details by Bo Delling,  have excelled in creating live-to-touch impressions of fish and the like that few of us have actually seen alive and healthy.

Given that there already many shark books, not least the excellent compilations by Leonard Compagno, and volumes dedicated to hagfish and lampreys, and of course there is FishBase, one might as author feel like facing a table already laid with easily digested goodies. Especially in a species-poor country like Sweden with an ocean part that on a world map looks like you can jump over it to land dry. This is not so. Thousands of fisheries and fish biology papers appear every year, and still nobody seems to know what marine fish eat, how big they get, how they reproduce, how old they get, or even where they occur or what they look like. Nobody even knows which one is one of the biggest skates in Europe, Dipturus batis. Honestly, von Bertalanffy curves carry no meaningful biological information.

Nevertheless, it was indeed possible to provide details on all the Swedish species of hagfish, lampreys and chondrichthyans.  It took six years to complete this volume, but rewardingly for all involved it feels like one has now turned pages entering into a new era of fish information in Sweden with the first real updated national fauna since 1895 in Skandinaviens fiskar (Fries et al., 1836-1857; Smitt, 1892-1895), and a worthy replacement to the popular standard Våra fiskar (Curry-Lindahl, 1985). It summarizes current knowledge and it provides a new platform for ecological and taxonomic research. And yes, of course the ray-finned fishes were not forgotten. They have been worked out in parallel and will be published in a separate fish-only volume to appear in the autumn of 2012.

The Encyclopedia is a project started at the Swedish Species Information Centre in Uppsala in 2001  and aims to produce a series of identification handbooks with keys in Swedish and English to the Swedish plant, fungi and animal species. It is a long-term project, aimed at covering the 30 000-40 000 species which can be identified without highly advanced equipment. They will be described in detail, including information on distribution and biology. For most of them, distribution maps as well as illustrations will also be provided.

With the present volume, there is now a newly published checklist of Swedish lancelets, cyclostomes and chondrichthyans. It is not long, so here it comes before it gets outdated. Species known only from occasional records are annotated. For those interested in Nordic exotisms, you also get the Swedish name.

Branchiostoma lanceolatum (Pallas, 1774) lansettfisk

Myxine glutinosa Linnaeus, 1758 pirål

Petromyzon marinus Linnaeus, 1758 havsnejonöga

Lampetra fluviatilis (Linnaeus, 1758) flodnejonöga

Lampetra planeri (Bloch, 1784) bäcknejonöga

Chimaera monstrosa Linnaeus, 1758 havsmus

Lamna nasus (Bonnaterre, 1888) håbrand

Cetorhinus maximus (Gunnerus, 1765) brugd

Alopias vulpinus (Bonnaterre, 1888) rävhaj

Galeorhinus galeus (Linnaeus, 1758) gråhaj

Mustelus asterias Cloquet, 1821 nordlig hundhaj

Carcharhinus longimanus (Poey, 1861) årfenhaj (single record)

Prionace glauca (Linnaeus, 1758) blåhaj

Galeus melastomus Rafinesque, 1810 hågäl

Scyliorhinus canicula (Linnaeus, 1758) småfläckig rödhaj

Scyliorhinus stellaris (Linnaeus, 1758) storfläckig rödhaj (two records)

Hexanchus griseus (Bonnaterre, 1788) sexbågig kamtandhaj (single record)

Somniosus microcephalus (Schneider, 1801) håkäring

Etmopterus spinax (Linnaeus, 1758) blåkäxa

Squalus acanthias Linnaeus, 1758 pigghaj

Oxynotus centrina (Linnaeus, 1758) trekantshaj (single record, actually from Danish Skagerrak)

Squatina squatina (Linnaeus, 1758) havsängel (single record)

Torpedo nobiliana Bonaparte, 1835 darrocka (two records)

Dipturus batis (Linnaeus, 1758) slätrocka (apparently two species involved)

Dipturus linteus (Fries, 1838) vitrocka

Dipturus nidarosiensis (Storm, 1881) svartbuksrocka (single record)

Dipturus oxyrinchus (Linnaeus, 1758) plogjärnsrocka

Leucoraja fullonica (Linnaeus, 1758) näbbrocka (two records)

Amblyraja radiata (Donovan, 1808) klorocka

Raja clavata Linnaeus, 1758 knaggrocka

Rajella fyllae (Lütken, 1887) rundrocka (single record)

Dasyatis pastinaca (Linnaeus, 1758) spjutrocka

Myliobatis aquila (Linnaeus, 1758) (single record)

 

References

Curry-Lindahl, K. 1985. Våra fiskar. Havs- och sötvattensfiskar i Norden och övriga Europa. P.A. Norstedt & Söners Förlag, Stockholm

Fries, B. Fr., C. U. Ekström & C. J. Sundewall. 1836 -1857. Skandinaviens Fiskar. P. A. Norstedt & Söner, Stockholm, IV+222 pp. Appendices 1-44, 1-140, pls. 1-60. Fascicle 2-3 (1837),  4 (1840)  5, 1839 (p. 111 dated 22 October 1839, ) 6+pls 31-36, Latin text 57-72 (1840), 7+pls 37-42, Latin text 73-92 (1842)

Kullander, S.O., T. Stach, H.G. Hansson, B. Delling, H. Blom. 2011. Nationalnyckeln till Sveriges flora och fauna. Ryggsträngsdjur: lansettfiskar-broskfiskar. Chodrata: Branchiostomatidae-Chondrichthyes. ArtDatabanken, Uppsala. 327 pp.

Smitt, F.A. 1892. Skandinaviens fiskar målade af W. von Wright beskrifna av B. Fries, C.U. Ekström och C. Sundevall. Andra upplagan. Bearbetning och fortsättning. Text. Förra delen. P.A. Norstedt & Söners Förlag, Stockholm, pp. 1-566+I-VIII+2 pp.

Smitt, F.A. 1895. Skandinaviens fiskar målade af W. von Wright beskrifna av B. Fries, C.U. Ekström och C. Sundevall. Andra upplagan. Bearbetning och fortsättning.Text. Senare delen.. P.A. Norstedt & Söners Förlag, Stockholm, pp 567-1239+1 p.

Smitt, F.A. 1895. Skandinaviens fiskar målade af W. von Wright beskrifna av B. Fries, C.U. Ekström och C. Sundevall. Andra upplagan. Bearbetning och fortsättning. Taflor.  P.A. Norstedt & Söners Förlag, Stockholm, pls I-LIII, pp. I-III.

Amblyraja radiata image

Amblyraja radiata, from Fries et al.