Scientific questions and answers of general interest (of course, not just frequently asked questions : our potential community consists of about 5000 persons around the world and not all ask questions).
Questions may be sent anonymously (but if so, it will only be published if no personal quarrel is involved).
Answers are supplied by a renown selected expert on the subject.
If you want to ask a personal scientific question, click on QUESTION.

Questions and answers may be freely published in Killifish Association magazines, provided that both authors (if any) of the given question and of its answer have formally agreed to (and provided that the date of updating is mentioned).

To go directly to the relevant FAQ answer, please select and click the appropriate link below.

  • Your NEXT QUESTION ? (48)
  • MOLECULAR results and species : isn't it already the end of the game ? (47) [December 2016]
  • MOEMA KENWOODI a new name in an unexpected genus : why ? (46) [March 2016]
  • COMPLETE MESS IN KILLIFISH NAMES in associations' contest lists of fish : any help ? (45) [June 2015]
  • CONTRADICTORY OSTEOLOGICAL RESULTS for Cyprinodon martae : any problem ? (44) [May 2015]
  • ANNUALISM IN NORTH AMERICA : any killifish species reported ? (43) [February 2015]
  • NOTHOBRANCHIUS NEUMANNI : change in species identification of aquarium strains along description of cryptic sagittae, serengetiensis, usanguensis (42) [January 2015]
  • RACHOVIA-MOEMA : new publication by Costa bringing new synonymies of Austrofundulus and Aphyolebias, not taken into account by Killi-Data (41) [November 2014]
  • HYPSOLEBIAS : new publication not yet quoted or quoted with delay by Killi-Data (40) [September 2013, January 2014, October 2014]
  • RIVULUS : one genus or several genera (39) [October 2012]
  • KRYPTOLEBIAS : ocellatus or hermaphroditus (38) [March 2012, October 2012]
  • RIVULUS MAHDIAENSIS : in the subgenus Laimosemion or Owiyeye (37) [April 2011, February 2012]
  • PSEUDEPIPLATYS : follow or not Collier and co-workers who synonymize Pseudepiplatys into Epiplatys (36) [September 2010]
  • DIAPAUSES : the embryonic diapauses in Nothobranchius, summary of knowledge (35) [January, August 2009]
  • USAGE OF NEW SUBGENERA NAMES : the cases of Simpsonichthys and others (34) [February 2008, December 2010, October 2012, July 2016]
  • CORRECT YEAR OF PUBLICATION : the cases of Nothobranchius cardinalis and others (33) [February 2008]
  • APLOCHEILUS MELASTIGMUS or Oryzias melastigma (32) [April 2007]
  • PLATAPLOCHILUS/PROCATOPUS (31) [January 2006, November 2009]
  • FLUVIPHYLAX OBSCURUM or obscurus (30) [May 2005]
  • APHYOSEMION PRIMIGENIUM GBN 88/10 population, variability or distinctive species (29) [March 2005]
  • Peruvian RIVULUS DISTRIBUTION patterns (28) [November 2004, January 2012]
  • Killifish of ORINOCO basin in Venezuela, non annuals, regional aquarium (27) [October 2004, May 2011]
  • EPIPLATYS LOKOENSIS, distinct valid species or a junior synonym (26) [October 2004]
  • CHROMAPHYOSEMION, distinct valid genus or still subgenus of Aphyosemion (25) [September 2004]
  • The physico-chemical characteristics of waters for TERRANATOS DOLICHOPTERUS (24) [June 2004]
  • The complete list of DNA PUBLICATIONS on Killifish (23) [February 2004]
  • What is your view about GRESENSI and other mirabilis, moensis, traudeae, intermittens  : species or subspecies ? (22) [December 2003, updated November 2010]
  • What is valid for Killi-Data, the case of Callopanchax HUWALDI (21) [December 2003]
  • Valid name for Rivulus ORNATUS and Rivulus obscurus (20) [September 2003, November 2004]
  • Availability of the new name Nothobranchius NUBAENSIS, without any type designation (19) [July 2003, December 2009, October 2012]
  • Distribution of Procatopus NOTOTAENIA and similis (18) [March 2003]
  • Vicariance or dispersalism for Killifish in Madagascar and MOLECULAR CLOCK (17) [February 2003]
  • Identity as Plataplochilus MIMUS for the Gabunese population of Tchibanga (16) [February 2003]
  • Fundulopanchax DELTAENSIS separate valid species or not, versus gularis (15) [January 2003]
  • Megalebias ROBUSTUS valid species or not (14) [December 2002, January 2003]
  • Austrolebias CARVALHOI 's original describer: Myers or Costa?  (13) [December 2002, July 2006]
  • Collection in COSTA RICA: best legal proceedings (12) [September 2002]
  • EPIPLATYS species from lake Fwa in Zaïre (11) [July 2002]
  • Isolated Rivulus GEAYI populations outside its normal range (10) [May 2002]
  • Rivulus HOLMIAE: type locality recollection (9) [May 2002, November 2011]
  • BUALANUM or elberti, following Lazara's in Kathetys special issue of 2001 (8) [April 2002, July 2002]
  • CODES for population names (7) [December 2001, July 2003]
  • MEASUREMENTS of geographical coordinates, egg diameter, stages of life (6) [April 2001]
  • ABBREVIATIONS of generic names (5), [March 2001, February 2009]
  • DIAPTERON identification (4) [September 2000]
  • COMPARISON with Lazara's KMI list of 2000 (3) [August 2000, June 2002]
  • PROCESSING Killi-Data (2) [July 2000, January 2004]
  • Ideal Expert Killi-Hobbyist LIBRARY (1) [July 2000, September 2004, April 2006, January 2007, December 2009]



Q(47): I am interested in killifish research with all its novelties and surprises. And I want to know your analysis on tools and techniques using genetics and killifish at the species-level, notably those so easy to understand horizontal trees (I know that you discussed many times the same topic at the genus-level). Isn't it already the end of the game  (47)? (Anonymous, Brasil, December 2016)

A: Your question is a frequent item in direct messages (mails sent to the editor) that are herein received and usually the answer is general and balanced (and not published herein), because the question is too complex to be dealt with seriously in a few lines and because the question is too general (wide) to have an approriate universal answer (only for killifish the situation is so complicated from the very-very little we know with some comfort that it would be better to pass the question to the next generation of researchers, say in 30 years) ; therefore, the answer to your interesting question could be easily finished at this point… but it happens by hazard that 2 new simultaneously distributed publications with comparative data allow to answer your question with more solid and exemplary arguments than in the past ; still, let's start by some basic evidence : yes, molecular data (a topic much narrower than simply genetics) have revolutionized killifish research output since the last 24 years (first paper in 1992 … and remember that, before, electrophoretic studies were the golden standard, now outdated) and yes that molecular evolution is great (even if morphological and osteological techniques of the old times are still valid in modern research notably when they use computerized softwares), and herein, in Killi-Data (see LISTS), it is so much an evidence and a priority that a molecular list of all studied so far species-populations has been lately created (to overcome the limits of GenBank) and is continuously updated, and, that a molecular bank of ethanol-fixed specimens available freely to researchers has been built, in order to boost research in that field (many, many species have not yet been studied with molecular techniques, and some species have been studied only by using 1 or 2 segments of the genome… and since we know for sure that multigene studies, i.e. studies with longer and multiple sequences of genome, with the ultimate target of the entire genome, mitochondrial and nuclear, give much better results, you see… it is not the end of the molecular studies with killifish, and researchers can be easily compared to Sisyphus !) ; obviously some -rare- species have been studied using distinct specimens and populations by distinct researchers and they have given sometimes different results pushing to a reserved attitude (and also to consider that the molecular techniques require high skills from the concerned researcher in order to align sequences properly and get correct results) ; anyhow now with the many collections of live killifish and with the costs of molecular techniques that have fallen sharply, molecular techniques are available to any student with access to a PCR machine (also much cheaper) and this is no surprise to see that in your own country (Brasil) several new teams have recently started to molecularly study killifish… and this brings me to the 2 new publications that change the perspective of your question ; those 2 publications of 2016 are led by a single author Wilson Costa ([ref. Costa, W.J.E.M, P.F. Amorim & R.C. Rizzieri. 2016. Molecular Phylogeny and Biogeography of the South American Savanna killifish genus Melanorivulus (Teleostei: Aplocheilidae). Vertebrate Zoology (VZ), 66 (3): 267-273, 2 figs.] and [ref. Costa, W.J.E.M. 2016c. Comparative Morphology, phylogenetic Relationships, and taxonomic Revision of South American killifishes of the Melanorivulus zygonectes species group (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters (IEF), 27 (2) (November): 107-152, 29 figs., 4 tabs.]), both studies are exactly using the same genetic segments [ND2, 16S rRNA, S7], are linked to an homogeneous timing (both distributed with less than 3 weeks gap) and are using mainly the same specimens (21), consisting of related species of the same group Melanorivulus, with the same outgroup (benchmark species), then results should be the same… but actually they are not, and by far ; the difference lies only in the slightly distinct sample of species corresponding to 2 executions of the computer programmes analysed by Costa's team ; precisely, in VZ, there are more studied species (and 1 less, wallacei) so that 4 newcomers [Rivulus (Melanorivulus) atlanticus, dapazi, egens, rutilicaudus] are added ; and indeed it reshuffles the obtained results, i.e. the 2 obtained trees are very different from each other, with (1) a distinctive topography, the tree in IEF having 9 branches, while the tree in VZ has 4-5 branches, with (2) species violaceus placed directly below the primitive schuncki in IEF and down the tree closely related to pindorama in VZ (itself isolated in IEF), with (3) species jalapensis placed directly below punctatus (huge geographical gap) in IEF and further down the tree closely related to atlanticus in VZ (small geographical gap, but still 2 distinct regions, coast and plateau), and so on ; in conclusion, yes the molecular technique is a real asset in the tools available to killifish research (in this case, species relationships), but clearly, no, any technique, even with those today's genetic data (erroneously tagged as ultimate by some people !), cannot be the end of the game, and there are no risk to predict that future trees, either in that group (both publications use only half of the total of over 46+ species, described for that group, nearly all by Costa) or elsewhere in killifish, with more sequences and more species in a given group will be different from present ones… research is moving progressively… there is no end of the game, even with molecular data (not to speak of other techniques we simply do not have any idea of, today) (Jean Huber, finalized December 2016)
legend    legend



Q(46): I am confused and lost. A few years ago I have received from a friend aquarist who got it through WildPeru (Brian Perkins) an annual fish that was labeled Aphyolebias sp., or Aphyolebias aff. rubrocaudatus. Now I see that it has a new name (if it is the same fish, I am not sure) as Moema kenwoodi, a fish named after Perkins, precisely. Why [the hell] that fish had an Aphyolebias label say 2-3 years ago and even less, and now it is named in Moema, a quite distinct fish I think from what I know of a fish I had a long time ago, now as Moema piriana from a trade import (NSC)  (46)? (Anonymous, USA, March 2016)

A: First, thank you for this interesting question which tackles the hot issue of generic naming but with a different angle, i.e. with the angle of the scientific magazine where a name is published (and specifically here you speak of the Journal of A.K.A. where Moema kenwoodi was described lately) ; you are absolutely right, that is puzzling, and on the other hand, the describer of the new name, the editor of the journal and the reviewers of the manuscript are also right ; that may look paradoxical but everybody is right, and before giving an explanation, let's tell you that for the specific case of Moema and Aphyolebias there is already a discussion in this FAQ page (question 41) and that there is for you a practical solution (to the paradox) also discussed in this FAQ page (question 45) ; you are right because still very recently (maybe also now) in the websphere (including in group discussions or in Facebook public or closed pages) those annual killifish collected by Brian Perkins around his company base are still named as Aphyolebias ; however, the describer (and the editor, and the reviewers) could not escape from naming the fish in the Moema genus because there is an unwritten but prevalent rule for any genus naming in scientific publications that is to use the latest published evidence, i.e. the last published systematic research results on the subject ; and specifically for your fish, they had to use a recent work by Costa (2014) where he synonymizes the genus Aphyolebias (he had created himself) into another genus, Moema (he also had created himself but previously) based on the results of his newest study ; that rule is neutral or a sort of gentleman's sound and fair agreement which imposes to take into account the latest evidence-based research on a given subject and that is a good rule ; here, in the present case, it is even simple to apply because nothing had been published to discuss systematics on Moema and Aphyolebias (maybe some doubts here and there, but nothing based on evidence) since their descriptions over 15 years ago until Costa's move 2 years ago ; besides (but that is another story because it does not apply in this case) the rule has a subrule which says that if several -not only one- evidence-based works are available and with alternative or conflicting results on a given subject and published in about the same period of time, the describer must use the latest evidence, must present the various alternative results, and if he does not use the latest published work, he must provide with an argumentation against it (or the editor or the reviewers must require that from him) ; actually the describer of Moema kenwoodi could have argued against the genus placement in Moema and select Aphyolebias instead, or say Trigonectes to escape from that duality, but he would have had then to provide new evidence (not only opinion) or objective weaknesses in Costa's work, in favor of his choice… and most descriptions of new species these days do not tackle the genus issue, they just follow the basic rule (sometimes adding that the genus discussion is beyond the scope of their work, in order to be ethically and prospectively on the safe side) ; on the other hand, if you read in details Costa's involved publication you will observe that the synonymization appears as a direct (automatic) consequence of the resulting tree organization of its sub-branches and that he is not afraid to change systematics following new results on a group he incrementally studies every 2-3 years like for Rachovia and Austrofundulus and back again (some of the reasons why Killi-Data prefered to take time and did not, yet, follow the move, notably waiting for molecular results) ; besides Costa's proposal to synonymize Aphyolebias into Moema is not that extravagant (just need time and external support) even if the 2 fish groups are quite dissimilar in morphology (as you say) and sympatric and cannibalistic one for the other (Moema adults feed on Aphyolebias !) : we have several similar examples (just to speak of Nothobranchius, a single genus, with up to 5 species of quite distinct morphology being sympatric and one, ocellatus "generously" feeding on smaller species… originally described in the full genus Paranothobranchius) ; then, yes, as a final word you are right and they are right, and all that is just automatic consequences of unwritten but useful rules, but we can live with that paradoxical complexity, can't we? (Jean Huber, finalized April 2016)



Q(45): I have been attending to several congresses at EU Killifish associations since last Summer and names are in a complete mess because of generic fluctuations : can you help please, at last  (45)? (Anonymous, Europe, June 2015)

A: This is a recurrent and regular, understandable, question although this time it is dealt from a different angle (listings in congresses of killifish associations) ; I shall tell you in the end what I personally believe is today the single respectful, still not absolute solution, but first I prefer to explain, hoping that you will understand better the pros and cons of that complicated issue ; let's recall that in the old times (say, from the 1950ies to the 1980ies), in the early days of killifish associations, there were many quarrels between experts (and their followers who were sometimes even more strict and ultimate) and these quarrels were opinion based ; the solution was to propose listings of names based on latest scientific evidence and that was the initial spur to publish Killi-Data (or K.M.I in the US, or,much later, Costa's own listings of 2008) and along incremental editions these lists tended to converge with fewer and fewer differences (but still some, due to varying respectable readings-interpretations of the ICZN code) ; that scientific evidence was based on new findings published in peer reviewed magazines (this means that the manuscript is criticized and improved by other scientific experts on the targeted theme before it can be published), but, frankly along time, the peer review process has become more or less a no-man's-land of neutral attitudes from colleagues to other, reputation-wise, supposed-equal colleagues (i.e., an article is left substantially untouched, except details, implying that the next reciprocal review will end up the same way… here I am exaggerating a «little» bit to make things easier to catch) ; as a consequence, along with other parallel considerations, new names have been inflating in a speedy way (still, each is not based on opinions… it is based on new molecular data, or on new micro-morpho-osteological data, or on combined molecular and morpho-osteological data, but everybody knows that with killies almost all populations are genetically isolated, then potentially there are thousands of new names to be described if that route is maintained in the future) ; besides, at the generic level (not at the species level, although the example could work too for the species level and a group of populations) if a researcher is studying a large group of species and his data show that a subgroup is nested within a larger subgroup, then (s)he is tempted, because a publication will be career-wise more valued if it includes a new name than not, to create one to several new generic names ; this a fact of life and only time will tell if the move and new name(s) will still be stable when new techniques will rise (e.g., complete genome vs. up to lately meaningful small portions of that genome) and, seriously, there is nothing you can do about it as an editor or as an expert or as a simple spectator (like you), except, (1) to develop (beforehand) double blind review processes of your magazine as an editor in order to push for more critical -in a positive way- reviews (it means that the reviewers as peers do not know who is the author or authors of the manuscript that they review and are not influenced by his -or her- notoriety/reciprocal reactivity) and this is sometimes undertaken in Killi-Data-Series, but this is of limited value because, for the usually narrow subject of a manuscript, there are not more than 4-5 potential reviewers then the secret is easily evaporated (vanished), and except (2) to analyse (afterwards) in depth an actual publication where a new name is created, e.g. benchmark the scaled length of the branches (the longer the better, but with the risk of bias), or, scrutiny the bootstrap value (the near to 100 value, as the maximum, the better) of the sub-branch, or else evaluate the solidity of the diagnoses (the new one and the old ones, re-dimensioned), of the identification key, and of the argumentation… and this is done in Killi-Data (with the assistance of other experts who easily «talk» in privacy, after publication) ; however, in case the generic change is not taken into account, there is a potential major drawback that the then freezing action is seen as being a judge, not objective, but subjective, and aiming to impose rules to everybody (and you can imagine that it is even more difficult when you are both a -say- objective judge and still a researcher, then a party, with his own publications (and his own new names !), like for Killi-Data and myself ; without playing on schizo grounds (2 personalities) you have to separate completely the 2 «jobs» or «hats» and I try to do it ; hence the new principles that rule Killi-Data since 2008 : (1) not to accept immediately and automatically any new generic change, (2) evaluate the solidity of the proposed changes (branch lengths, bootstraps values, rediagnoses) with optionally the help of other experts in private correspondence, (3) take time for the possible availability of a competing work with different results, which means in the end that there are less frequent turmoil of name changes compared to the period 1990ies to 2010s and thus more stable systematics, notably for large genera which remain un-split like Rivulus, Simpsonichthys, Aphyosemion, Nothobranchius, Fundulus) ; on the other hand you might think, and some associations do so, that the previous process (automatic acceptance of generic changes) is much better and they let things flow in the splitting way ; however it does not work any more if another author publishes with the same scientific value (and review process) a work on the same group with different or even opposite results because nobody is going to achieve to go reverse (i.e., to move back to old naming), or if so, very very slowly (e.g., the case of synonymization of Adinia or Rivulus, Simpsonichthys, Aphyosemion, Archiaphyosemion, Nothobranchius)… their process is then not flexible, it can only work one (splitting, today) way (by the way, is there any killifish association which has synonymized back (again) Austrofundulus into Rachovia or merged Aphyolebias and Moema following one of Costa's very recent works ? none, at this time, then the situation is not at all simple) ; in other words, yes, the old Killi-Data method of keeping in line with research and the same still method prevailing in killifish associations (to various and diverse degree, hence the differences you stress between each) would be acceptable today if, and only if, any new scientific peer-reviewed-at-least publication would be automatically followed, including to reverse things (you would imagine what mess it would soon become !) ; the key issue is then time if everything else is equal (if not automatically translated, how long the move will have to wait ? no answer obviously (not easy, notably because today, with the web 2.0, the scale of time for any information is in the amount of a fraction of a second… later than a second has a much deprecated value !)) ; besides, before going further, let's view the issue from a different viewpoint ; do the killifish associations want to (if they could) have a uniform listing of names ? the answer is I am not sure since today with the cooperative platforms (such as FaceBook, Twitter and forum marketplace), the issue is not, like in the 1980ies and the beginning of the Internet era, to produce evidence-based discussions, but on the contrary to accept all types of positions (no problem because it is so ephemeral, one post being replaced by another at high speed)… and even among experts (contrary to what you would imagine !) the new neutral culture is shared by many (i.e., let have several naming possibilities) and it is even enforced because no expert is willing that his naming is prevailing above all others (he or she prefers that his/her paper/contribution is actually read, valued and remembered a while which is not granted owing to the overwhelming shower of data everybody faces everyday today ; therefore, you see, things change : in the past, struggle attitudes, now tribe attitudes (and this is not only an issue of social behavior… it is also a measure of humility since it is far away the time when people thought understanding killifish phylogeny would be solved by more detailed morphology, then by osteology, then by molecular sequences, then by computerized matrices… what will happen with multi-gene techniques or full genome techniques in the near future ?) ; finally, I am going to write something that will shake you naked (beware !) ; this is not the end of the game and scientific circles where this hyper-inflation of names is harshly discussed these days, there are voices (not minorities, not minor standing) to support the disappearance of describer's names associated with taxa and even voices to support the disappearance of names at all, replaced by code numbers linked to genetic bar codes (then all discussions of the past 50 years, Killi-Data and other similar projects, even ICZN would be drown !) ; now, let's go back to ground… you have read the above difficult and technical text and hopefully you are convinced that Killi-Data is still the mirror of objective, solid and science-based evidence (with no attempt to impose names), then the practical, respectful, but limited solution to your question is that you keep away from problems of communications and use both names in case of divergences between authors, exactly and strictly the following way : first use the generic name you prefer, then second use the species name, and last between parenthesis use the alternative generic name you know, e.g. (for genera I am sure you have in mind because frequent in listings of congresses), (1) Rivulus punctatus (Melanorivulus) or Melanorivulus punctatus (Rivulus), (2) Ophthalmolebias constanciae (Simpsonichthys) or Simpsonichthys constanciae (Ophthalmolebias), but please do not use at all (1) Rivulus (Melanorivulus) punctatus, (2) Simpsonichthys (Ophthalmolebias) constanciae (parenthesis in-between), which would mean that you consider simply the subgeneric name (trivial, no interest) and please do not use at all, the following stupid nonsense (1) Melanorivulus (Rivulus) punctatus, (2) Ophthalmolebias (Simpsonichthys) constanciae (parenthesis in-between), which would mean that Rivulus is a subgenus of Melanorivulus, and (2) Simpsonichthys is a subgenus of Ophthalmolebias (that is impossible because of the law of anteriority in ICZN code) ; besides, to go back to killifish associations listings, I would suggest that they apply the same rule to avoid the non-flexibility problem I raised above in case they have to reverse their position, the number of divergent positions on genera between authors being not many, or that they wait some time to check if in scientific circles a given move with generic name changes is meeting some concensus or not (remember that even before 2008, there were attempts to split the genus Aphyosemion into several genera but generally nobody followed the moves, Killi-Data included, because no diagnoses were available for all subgenera, and note that today ironically a researcher could use my new diagnoses of 2013 to upgrade all subgenera of Aphyosemion as distinct genera !) ; besides, at some point (in time) there is absolutely no probability that everybody uses the same names on this planet (think of the old case Pseudepiplatys-Epiplatys or the recent case of Adinia-Fundulus or the brand new split of Profundulus-Tlaloc) unless a single list is used everywhere at the same time (and then people will criticize it as imperialist) ; if you are acting mainly in a country where a Killifish Association is a major player then use their names list (e.g. AKA in the US, DKG in Germany, KCF in France, etc.), if you are acting more internationally then use the above recipe with parentheses (after) ; as a final word I should say that for Killi-Data (hat n°1) I have no theoretical objections or reservations to split genera (e.g. for Rivulus or Profundulus or Archiaphyosemion, because for Simpsonichthys it is already the second or third split by Costa since Cynolebias), provided it is solidly evidenced (and diagnosed), even if as a researcher (hat n°2) I am a reasoned lumper (the contrary of a splitter) and I am reluctant to split genera (I gather units rather than partition them)… and there again, we are facing issues of philosophical strategies, Costa being a decided splitter and other researchers being strict lumpers)… you see how much your question was not as simple as it looked… and final, final comment, another researcher, Andrew Furness, has just published a new work [Furness, A.I. 2015. The Evolution of an annual Life Cycle in killifish: Adaptation to ephemeral aquatic environments through embryonic Diapause. Biological Reviews of the Cambridge Philosophical Society (Biol. Rev.)] with combined molecular data and maximum likelihood processing -not parsimony- in which he comments (and independently agrees with many including me) on the non-necessary splitting (with present data) of Rivulus, Simpsonichthys and even Archiaphyosemion and these comments have been added, as an update to the concerned newsletters, i.e. INFOWEB12 on Simpsonichthys, INFOWEB13 on Rivulus and INFOWEB11 on Archiaphyosemion ; and one final-final-final (philosophical) word, back on your question, please do not think I want all associations use the same listing of names, I too-much like freedom and diversity and I know that killifish research is too-much complicated that ever a unified list may be available and I live with my time, today neutral… a little more caution over the laisser-faire attitude (natural splitting drift) would be desirable, yes, but it is not compulsory or binding… and there is one thing I am positively sure of : there is indeed an imperious need of in-depth research for many-many recent names for which we have singly and barely (and unfortunately) the description alone with a single type locality and no other data such as distribution, life history traits, phylogenetic relationships, extended localities, variations/variability, hybridizations (etc.) ; and now truely final, let's add with some sense of humour, that in the end all that story is due (yes their fault) to computers with their non sense behaviour of "0" or "1" binary-only options and for killifish compulsorily one name labelling only, it would be much better -but unachieved yet, you will agree- that Killi-Data listings are shown according to your personal taste while your personal taste would have been deducted from your previous opinions in mails, forums, archives of your own computer, traces from your IP conversations (etc.)… and obviously (with computerized filters that automatically change the names you use) your then selected name would be translated to another when you discuss with another person with a different personal taste (also deducted, etc.)… the future ? (Jean Huber, finalized October 2015)



Q(44): I immediately noticed the contradictory osteological results for Cyprinodon martae (Cubanichthys-like) between the new Costa's work on the fish and your Yssolebias, Terranatos-like, on the same fish : aren't you personally concerned  (44)? (Anonymous, Germany, May 2015)

A: Well if your question suggests I would be upset, quarrelsome or even destroyed, you are mistaken I am sorry to say ; simply because the times when researchers were continuously debating on controversies based on opinions only is definitely over… Costa's latest publication [Costa, W.J.E.M. 2015b. Phylogenetic Position and tentative generic Placement for Cyprinodon martae Steindachner, 1875 (Teleostei: Cyprinodontiformes), a killifish from northern Colombia. Vertebrate Zoology, 65 (1): 27-30, 1 fig.] is based on hard data and respects ICZN rules, then it is perfectly acceptable and even interesting to read ; besides, research is today based on individual competition (sometimes but rarely with cooperation) and any improvement (who would claim that killifish research is simple ?) is welcome ; more specifically I do not feel concerned because the osteological finding that Cyprinodon martae is belonging to rivulins (and not to cyprinodontoids, as per Costa) is not due to me but to another researcher whom I ethically named and acknowledged for his contribution in my publication ; based on that finding I produced a computerized morphological study comparing genera related to Rachovia and demonstrated that Cyprinodon martae was very distinctive and hence described the new monotypic genus Yssolebias (Huber, J.H. 2012b. Reappraisal of the Phylogeny of Rivulus and its Allied focused on External Characters. Killi-Data Series 2012, 9-25, 3 figs., 2 tabs. | Huber, J.H. 2012c. Description of Yssolebias, nov. gen., a new monotypic fish genus for an old and phantom species of Colombia. Killi-Data Series 2012, 26-31.) ; the first osteological finding was derived from a radiograph prepared by my old friends in Vienna Museum back in the early 2000s, the new finding by Costa is based on a new radiograph of the same specimen with a new technique, and least that can be said is that the new radiograph is clear and fine (which was not at all the case of the first radiograph, as stated in my paper : "believed that it is closer to the Rivulinae because of its rather slender dentary and, although it is not absolutely clear from the radiograph, because of its anterior Dorsal fin radial and ray pattern") ; now to practical consequences… like all evidenced-based data, Costa's work has been accepted (and his finding has been confirmed by a third expert on osteology) in Killi-Data and the genus and species (Yssolebias martae) have been moved from the family-group name and tribe Rachoviina to the subfamily Cubanichthyinae (see FGN) and in the database along its next upload ; yes, surprisingly, there can also be improvements in older techniques like radiography of old types (not only in new techniques like molecular biology)… just think of the huge progresses that computer programs, like PAUP, Phylip and others have permitted to very old morphological techniques, for example ; besides, a new technique, along radiography, is emerging for preserved material, notably for old specimens, i.e., a CT scan, and in the present case, that would also be useful to dig in further and maybe Costa will publish on the single type of Cyprinodon martae with that technique ; but the best would be that the fish is recollected live… and here, yes, I feel sorry to have pushed Frans Vermeulen for 3 collecting trips in order to rediscover the fish in annual biotopes while actually it should -hopefully- be living in brackish waters of the delta of Magdalena river in northern Colombia  that refocused target of first live collection of the species is indeed a major priority because it will be much in line with the palaeo-history of Killifish along the ancestral previous Tethyan sea and the link with Cubanichthys and Chriopeoides (if valid) from Cuba-Jamaica, very far away Caribbean islands, will be tackled if Costa's hypothesis is fully confirmed of the link between Cubanichthys and Yssolebias ; finally, there is a (minor) error that I need to correct in Costa's work : he defines Killi-Data Series as my "private magazine" ; not at all, it is fully public, it respects all ICZN requirements for publication and there are other authors than me ; if Costa (or his institution on his financial account) wants to buy a printed copy, it is affordable and easy and the process is described in the PAGE in all details, just like for many scientific magazines (e.g., Ichthyol. Explor. Freshwaters, where Costa publishes frequently), and cherry on the cake, Killi-Data Series is entirely free as PDFs for members of K-D-I, the non profit association holding Killi-Data [update : Frans Vermeulen has reacted by writing "I was suspecting this fish could be a saline species from the beginning and for that reason I went collecting in various saline waters around Santa Marta as well during my surveys… without any luck"] (Jean Huber, finalized May 2015, updated May 2015)



Q(43): Dear Killi-Data, I am a former fish-keeper who is thinking of starting up again, and I am considering my options. Although I never raised killifish, I have become interested in them, so here is my question : are there any annual/seasonal killifish that are native to North America?  (43)? (Neil Barmann, on the official FACEBOOK page, February 7. 2015)

A: The answer is clearly 'yes' for one species, but before naming it, let's dig a little bit into both of your terms/words of annual/seasonal and North America ; if you define annual or seasonal as a species that lives only 1 year, then there are many in the wild in North America because many Cyprinodon sp. (and others) do not live much longer than a year or even less than a year, i.e. eggs hatch, fish grow and breed and die within a year, even if in aquarium they live 2 to 3 years (this has been proven by numerous scientific studies, for several reasons, the first being that their life-cycle is short within a cohort) ; besides several Fundulus sp. are seasonal because their biotopes are ephemeral -water dries out or moves due to tide- and they do not know well how to escape by jumping like Rivulus sp. (actually this is very part of ancestry of all killifish when, several dozens of million of years ago, they were living in their brackish very unstable original biotopes in-between sea and freshwaters (then already eggs could survive under dry or humid conditions) ; alternatively, if you mean seasonal or annual, species with eggs that can develop into dry substratum and show arrests of embryonic development (they are called 'diapauses', with 3 possible stages, I, just after egg is laid, III, just before hatching, and II in-between the two events), then also the situation is not black and white again, rather gray with intermediate species, e.g., eggs of many Fundulus sp. can undergo diapause III (also diapause I) and wait for several months before hatching (this has been proven by numerous scientific studies) ; however, if you mean by seasonal or annual, a species that compulsorily needs to undergo diapause II with eggs in the wild that compulsorily need a biotope that seasonally becomes dry each year, like most Nothobranchius, Simpsonichthys, Trigonectes sp. and others (although again this is not black and white, rather gray with intermediate species), then there is only one species in North America ; the second issue is about what is North America ; if you restrain your definition to the US and Canada, then there is no strict annual killifish species, but if you include Mexico, then here it is ; the name is Millerichthys robustus and it comes from coastal basins of rio Papaloapan, Coatzacoalcos, Chiquito, Cotaxtla, in Oaxaca and Veracruz states, eastern Mexico ; actually, it was discovered by Myron Gordon, in 1940, but only properly described by Miller & Hubbs in 1974 as Rivulus robustus, then it was NOT thought as an annual species, although authors like Huber (1992) or Costa (1995) when he moved the species to the new genus Millerichthys had doubts on the genus placement as a non-annual, and even Huber (2012) after a computerized review of all species related to Rivulus placed Millerichthys robustus not close to Rivulus, but close to true annuals from northern South America in genera Rachovia, Austrofundulus, Terranatos, Llanolebias (Colombia, Venezuela, maybe more northerly, who knows ?) ; and … very recently this has been confirmed by the first live collections and breeding, yes Millerichthys robustus is a true annual killifish [Dominguez, O.C., M.A.C. Mosqueda & S. Valdesalici. 2013. First Observations of Annualism in Millerichthys robustus (Cyprinodontiformes: Rivulidae). {Ichthyol. Explor. Freshwaters, 24 (1) (July): 15 - 20, 3 figs., 1 tab.}] and another publication by the same team is eagerly expected very soon to detail this annualism (diapause II or not, etc.) ; therefore the answer to your question is correctly 'yes'… but you are not 100% lucky, alas, because the species has not been distributed to aquarists and it may well not be ever (except at least to experts in conservation) because it is highly endangered of extinction ; sorry, you will have to move your thoughts and hopes to widely distributed annual species in aquaria, with a very large choice (several hundreds of species !)… I hope you are not too much disappointed by this frustrating answer ; alternatively you may feel to be challenged to become an expert breeder and adhere to conservationists ideas (and hobbyists' specialized groups), and maybe in some years after many efforts you will truly be so and may get the opportunity of receiving a few eggs of this unique, primitive, -but not very colorful- species to maintain it alive in our -alas- destructive world and society (Jean Huber, finalized March 2015, updated May 2015)



Q(42): Dear Dr Watters, I have read your descriptions (with Wildekamp and Shidlovskiy, in Journal of the American Killifish Association) of 3 new Nothobranchius species, sagittae, serengetiensis, usanguensis, related to neumanni (then with a new restricted definition) and seegersi, could you tell for aquarists maintaining Nothos strains the changes in species identification of past and present aquarium populations/aquarium strains derived from your publication  (42)? (Jean Huber, Paris, France, January 2015)

A: I will do my best to provide some information as below ; obviously where my own collections are involved, there is no problem and I can be quite certain what we are now dealing with ; however, with the populations collected by others, I may not have sufficient information to be able to provide a definitive interpretation ; therefore, in some cases I can only make a guess based on limited information (I do not always have access to the detailed collecting information of others) and whatever photos I have been able to view on the Internet ; besides, there may be some dispute about the identification of some of neumanni populations (in the new restricted sense) and I base my interpretation on photos (on fish & egg listings and Internet platforms of auction) and very general locality information… as with many populations of Nothos, these days, it will take genetic analysis to classify them definitively ; (1) for neumanni (in the new restricted sense), I list here only some of the more recent collections that have been offered (there are, of course, many, many more populations that have been collected over the years, notably TZ 97-17 belongs also here, collected by Seegers in 1992, southeast of Lake Manyara): Mapacha (K 2011-06), Makondo (K 2011-08), Seneki (TZ 08-5), Rubuga (K 2011-12), Mabondeni (TZ 08-17), Aghondi (K 2011-02) ; (2) for serengetiensis (I am not aware that any of these are still in the hobby at present) : Ikoma (TAN 93-1), Ikoma (TAN 93-2), Tarime River (TAN 93-5), Bunda (TAN 93-6), Ndabaka (TAN 93-7), Nyalikungu (TAN 93-9), Magu (TAN 03-17), Magu (K 2011-14), Kalemera (K 2011-15) ; (3) for sagittae (note that the Kiandere locality is the identical place to the "Tarime River" localities… the Kiandere population may be in the hobby at present) : Tarime River (TAN 93-5), Kiandere (K 2011-19), Tarime River (TAN 00-3), Ndabaka Gate (TAN 00-4) ; (4) for usanguensis (I am not aware that any of these are still in the hobby at present, but I may be wrong as I do not rigorously keep track of such things) : sp. Mbeya, Sinyanga (TZ 88-17), TZ 92-149, Halili River (TAN 03-2), Sonyanga West (TAN 03-3), Sonyanga East (TAN 03-4), Utengule (TAN 03-5), Usangu (TSTS 10-08), Usangu (TSTS 10-09), Usangu (TSTS 10-10), Usangu (K 2011-29), Ruaha (K 2011-30) ; (5) for seegersi, TZ 92-113 population (Malagarasi River) belongs here, as collected by Seegers in 1992 (the first time representatives of this species were collected), and also Ipole, Itumba (TSTS 10-4), Mabangwe (TZ 08-14), Mkola (TZ 08-12), Ngoywa (TZ 08-13) (Brian Watters, finalized January 2015)



Q(41): I have read that Dr Costa has synonymized the genus Austrofundulus into Rachovia and the genus Aphyolebias into Moema but I do not see any change in your list of valid (or correct) vs. used names (please be simple, I am an aquarist and we are several, locally, to discuss that)  (41)? (Anonymous, New York state, USA, November 2014)

A: First, as stated in newsletter INFOWEB13, Killi-Data is not following automatically generic changes any more and this is not aiming at Costa's works only ; in this case, this is the contrary as usual with that researcher's habits : here, he synonymizes taxa, he does not split taxa, but the decision by Killi-Data follows the same rules, i.e., not to disrupt names without due evidence ; actually the 2 mentioned synonymizations are different although they are included in the same publication [Costa, W.J.E.M. 2014c. Phylogeny and evolutionary Radiation in seasonal rachovine killifishes: biogeographical and taxonomical Implications. Vertebrate Zoology, 64 (2) : 177-192, 1 fig.] and within a general review (molecular and morphological) of a large group of genera related to Rachovia : actually Austrofundulus had already been fused into Rachovia in the 1990ies by Costa himself and subsequently he reversed his decision (and now he goes back to it) whereas Aphyolebias had never been fused into Moema before (but frequently the validity of both names have been questioned by several researchers, as putative synonyms of Pterolebias and/or Trigonectes) ; the in-depth analysis of Costa's new work (like for the Rivulus review a few years ago) shows that only a few species are included in the study (merging molecular and morphological data in a tree requires to have data from the 2 techniques for each species, which is easy with morphology but it is a major constraint with molecular biology needing live or fresh material) and that is a major weakness (at least temporarily, pending a future more comprehensive work by, for example, Costa himself, as usual) ; another major weakness, at least from my personal view, is to mix both types of data, molecular and morphological, in a single tree (using TNT software) without showing that the 2 individual trees are identical or similar (again that was also a problem with Costa's Rivulus review where the processing of the morphological data alone in a subsequent publication ended up with a very different tree) ; as a result, Killi-Data is not accepting these synonymizations and further evidence are needed in the future (but still Costa's point of view is not ignored and if you look in detail to the list of USED NAMES, you'll see that, for example, Moema peruensis is mentioned but as a synonym of Aphyolebias peruensis, the current name since about 20 years) ; mind you : Killi-Data is not opposed to synonymizations or splitting of genera as such (research is moving) and when the full revision of the genus Fundulus was recently published using molecular and morphological data, with separate and merged trees and high bootstrap values and solid evidence, the proposed synonymization of genus Adinia into Fundulus was right away accepted (and if you browse the Internet, you'll see that this major change is still not updated in the various online sources, like many such moves… Internet and their services providers are easily following (copy-pasting) for a new taxon, but that is another story if it is to evaluate a publication and update data without a new taxon !) ; a final word (or advice) on emotional aspects of nomenclature, because you say you are a group of aquarists (but anonymous !) : please do not take these changes too emotionally (one way or the other, splitting or synonymizing) ; with the emergence of molecular techniques, researchers (and maybe also the reviewers of their works before their publication) tend to be not very much concerned with consequences with their synonymizations implied by the position of a group (e.g., a genus) in their tree ; here, in the present case, the tree developed by Costa shows that Rachovia and Austrofundulus are each based on 2 tree branches (with Rachovia maculipinnis at the root of both branches), then Costa immediately (automatically) makes a synonymization (but that would be debatable because if that practice would be generalized, then many genera, including a lot that he himself described would be synonyms) ; and the tree developed by Costa shows that Moema and Aphyolebias are each based on 2 tree branches (with Moema staecki and Aphyolebias peruensis on the same branch as related and Moema piriana at the root of both branches, then Costa immediately (automatically) makes a synonymization (but as explained above, 3 species studied for the all species of the genera Aphyolebias and Moema, that is too limited, and besides the bootstrap value separating those and the genus Trigonectes is too low)… therefore as you see, our decisions are argumented (evidence-based) and not emotional (and in no way critisizing Costa who will, it is hoped, publish again on that group with more data and more species) (Jean Huber, finalized November 2014)



Q(40): I am a computer nut for Internet and I have been able to trace on Internet a new description by Costa, Amorim, Bragança (Hypsolebias xxxxxxxxxxus) since beginning of September from the Parnaiba River basin in Brasil, but I do not see any information in your flash news… why didn't you quote that publication or am I quicker than Killi-Data  (40)? (Anonymous, Russia, September 2013)

A: Well, you are right to be puzzled and congratulations for being so quick with Internet ! Actually we are already aware of that publication but we have refrained to mention it in order to avoid problems for its authors ; yes that description has been issued in the Journal of Zoological Systematics and Evolutionary Research, but only online (as a preview) and, IMPORTANTLY, online publication means not (ICZN) officially published ; the International Commission of Zoological Nomenclature (since 2000) requires for a new name (species or genus) in order to become available (i.e., nomenclaturally speaking valid) that it is published in print (an online publication is not enough) ; therefore do not use or quote this paper since you would create a nomen nudum until it is actually printed, i.e., in several weeks as scheduled (or wait until it is mentioned in K-D online which will mean that we have checked it is officially published) ; that new rule by ICZN has been enacted to avoid inadvertent descriptions as it used to occur before 2000 ; obviously if you do not follow this (kind) advice and publish that name in a printed magazine (e.g., a killifish association magazine) you will create a new species with you as the author of the description (describer), not Costa, Amorim & Bragança (and understandably they will not be happy) but the problem will not be major since your name will become a nomen nudum (i.e., a name without value) as the ICZN requires at the same time that the new name is linked with preserved material as holotype deposited in a scientific institution ! in conclusion, do not worry too much about the risks on Internet but do not propagate new names without ICZN availability and, maybe,… do not attempt to be quicker than Killi-Data ; addendum : it has been finally published in the February issue of 2014 of the Journal, as Hypsolebias coamazonicus ; but, but, your question appears more complicated than it looks like : it is fairly possible that an electronic publication is considered as available following the new ICZN code (only valid since 2013) and ahead of a possible printed edition (or even without a subsequent print) ; this is the case of Xenurolebias cricarensis and Xenurolebias pataxo (both KD-maintained in Simpsonichthys), published online at the end of May 2014, … but not to be printed before September 2014 ; it is impossible in a few words to delineate the case clearly as a general rule but when you read in an electronic publication both a zoobank number and the actual number of the pages (e.g., pages 371-385… not pages 1-15 !), then there is a good probability (but not 100%… it is more complicated and stricter with other detailed ICZN rules) that it is validly published with an electronic support alone (and then that will be the first date of official publication, even if a print follows) (Jean Huber, finalized September 2013, updated January 2014, July 2014, October 2014)



Q(39): Why don't you follow research anymore at the genus/subgenus level and precisely Costa's split of the genus Rivulus… a peacocks quarell  (39)? (Allegro barbaro, UK, October 2012)

A: Well, we thought we had already answered that question in INFOWEB13, but you are not alone to suspect something (several other similar questions have been raised at the same time) and just inviting to read that newsletter might not be sufficient. First, is (obviously) to say that the decision was taken in general terms and that has nothing to do with Costa (it was taken about a year before Rivulus split, i.e. along with the proposed -not followed- split of the genus Simpsonichthys by Costa, INFOWEB12)… and the proposed -not followed- split of the genus Archaphyosemion by Sonnenberg, see the discussion in INFOWEB11) ; second, is to mention that the decision was maturing since several years, not because of increased conflicts but because the arena has changed : (1) the reviewing process of scientific magazines has changed dramatically and today there is no necessity to establish that the level of distinctiveness for a generic name, genus or subgenus (i.e. that meets scientific consensus) is reached, instead it is just necessary to name to which generic concept the author(s) refer(s) and that is accepted (obviously provided that evidence data are forwarded to support the move)… and there is at least a dozen different definitions of the genus concept (and a parallel situation exists for the species concept) ; no doubt there is always one that fits with the one the author(s) is (are) comfortable with ; (2) the gap between nomenclature and systematics has also increased dramatically in recent years and that is a pity : basically nomenclature should be stable, and systematic research is more open to ups and downs (each new result being there to replace the previous one, with hopefully improvement of knowledge)… nomenclature is based on diagnosis (i.e. differentiating relative -not absolute- characters) and conservative usage, and that also has changed dramatically, not because of new laws of nomenclature -the ICZN code is still the same since 2000-, but because authors are moving their understanding of what is a genus (or a species) ; (3) the gap between a solid diagnosis (with external stable characters) and a comparative multiple diagnosis (with many minor characters overlapping for each other but with a global distinctive picture) has been dramatically enlarged (also for the species concept and here it is not surprising because authors deal more with cryptic species than in the past)… and in both directions, i.e. to describe a new genus, to split a large genus into several less numerous genera, or even to destroy a generic (or subgeneric) level (3 examples, in a few recent years : Megalebias has changed its status to full genus, mere synonym, then of a subgenus with a different definition [yes!] than the initial definition, or another example, the subgenera of Epiplatys have been destroyed because the molecular results show that the past understanding of the various existing subgenera and related genera was -according to that technique- totally erroneous, but the author did not intend to rebuild a more stable system with re-diagnoses of each or some units/subunits (that policy is respectable, but non operational for Killi-Data), the various subgenera of Simpsonichthys were split into genera by Costa, not because of new data, but because after a biogeographical re-assessment of the various sub-units, it fit better to the tree (with revalidation of Spectrolebias previously synonymized) ; do we mean that the quality of research or of researchers have weakened recently ? Not at all (on the contrary, research is much more evidence-based and less opinion-based than say 50 years ago)… research and nomenclature are diverging, or to put it more properly nomenclature based on fluctuating-improving research and nomenclature for stable usage (as per Killi-Data or any database of names and also for aquarists) are strongly diverging (that is not a serious problem if we keep for naming fish in databases some kind of stability) ; third, is to stress that we are not opposed to split genera (with less numerous species names) in Killi-Data and obviously we are not opposed in principles to split the genus Rivulus… when it will be ready, i.e. when solid research results with stable diagnosis on external characters (not a mere DNA or biogeographical phylogenetic tree with branches that are named by some authors or oppositely remain un-named by other authors), that may happen for Rivulus, but that is not for tomorrow at least for 2 important reasons : (1) that genus is virtually unknown in the huge Amazonian basin, and that is not a detail to understand the phylogeny of the subgroups in the Amazon basin, the heart of the genus (remember that not so long ago an argument to separate, among Rivulus, the pictus-punctatus superspecies (or Melanorivulus) from the geayi superspecies (or Laimosemion) was that they were distributed well apart, but with recent collections we know that they are bordering each other and could even be sympatric in the Amazon basin, remember that the number of minute species of Rivulus in the Amazon basin is numerous and that they are not related to each other as previously hypothesized, and probably some of them are not even related to already named groups like Laimosemion or Owiyeye, (2) the most confused situation comes with Laimosemion for which the initial diagnosis (by Huber, in 1999) is obsolete and the single primary (osteological) diagnostic character proposed by Costa (in 2006) is unseen in at least 2 species, including the recently Laimosemion paryagi (described as Laimosemion because this corresponds to the latest generic publication) in which the authors state that the diagnosis of Laimosemion is not founded at this stage and that the tree branches of the Guyanas Laimosemion are extremely long, implying a long separation along a unique morphological differentiation, and leading the way to (maybe) a further split of Laimosemion itself in the future… the situation is not that easy and simple (to use an ironic understatement) and, as usual more research is needed, which Killi-Data will follow carefully to provide with a good and hopefully more stable -but not permanent- picture (Jean Huber, finalized October 2012)



Q(38): What do you think of Costa's reshuffle of status of 3 species of Kryptolebias (previously Rivulus) ocellatus or hermaphroditus or hermaphoditus  (38)? (Anonymous, Switzerland, March 2012)

A: Well, the situation has become extremely complex since Costa's publication [Costa, W.J.E.M. 2011a. Identity of Rivulus ocellatus and a new Name for a hermaphroditic species of Kryptolebias from south-eastern Brazil (Cyprinodontiformes: Rivulidae). Ichthyol. Explor. Freshwaters, 22 (2): 185-192, figs.] and vanishes all past considerations since Seegers (1984) including a past ICZN decision (and regrettably Costa's move has not been opened to Seegers or others for argumentation) ; Costa (2011a), after having examined the unique type of ocellatus, considers that the to-date identification of ocellatus by Seegers is erroneous and the description by Seegers of caudomarginatus corresponds to the true ocellatus, then caudomarginatus described by Seegers is a junior synonym of ocellatus, and ocellatus in the previous sense by Seegers is described as hermaphroditus : he supports his argumentation on 3 points vs. caudomarginatus (number of scales of the longitudinal series equal to 43 and lower in ocellatus, depth of body in male lower in ocellatus, Ventral fin in male of ocellatus shorter, not reaching the base of the second or third Anal-fin ray), but the last 2 points are minor and variable upon individuals, and the first point, major, is debatable (most authors since Regan have disclosed more than 48 scales in LL series for ocellatus, because the rows and the scales are very irregular), and the only value of Costa's analysis is based on his own observation of the type specimen (Seegers is the only person to have seen it too, in modern times) ; in details, according to Costa, just counting scales in a straight line along the lateral line scales and excluding scales posterior to the hypural plate, there are 42-46 scales in caudomarginatus, while in ocellatus sensu Seegers, there are 47 or 48 scales while the holotype of ocellatus (ZMB 7448) has 43 scales ; pending further observations (notably from Seegers and also from third parties), Costa's decision is followed herein as the latest published evidence (with reservation)… however the biological situation is far more complex than nomenclature : Tatarenkov et al., also in 2011, have published a molecular study that (as a side note) mentions that a population from Puerto Rico (and 3 others, not too far) of Kryptolebias conform with that of hermaphroditus and not with marmoratus as it would be expected ; whether there are true Kryptolebias marmoratus on Puerto Rico is unknown at this time ! (Jean Huber,  finalized April 2012, updated October 2012, July 2013)



Q(37): Where to position Rivulus mahdiaensis, in the subgenus Laimosemion or Owiyeye  (37)? (Anonymous, Germany, April 2011)

A: Well there is no present answer because 2 positions have been published, one by the describers Suijker & Collier (2006), following a molecular study that clearly places their new species close to Rivulus lyricauda a member of the subgenus Laimosemion and the other by Costa [ref. Costa, W.J.E.M. & F.L. de Souza. 2009. Rivulus (Owiyeye) mahdiaensis Suijker & Collier, 2006. D.K.G. (Deutsche Killifisch Gem.) J., 41 (5): 64-65, 2 figs.], following an osteological study which shows that mahdiaensis possesses the vertebrate characters typical of Owiyeye, absent in Laimosemion ; therefore, it is impossible to clear the situation, until another study sorts the problem… but ironically you have to remember that at the end of 2011, Costa published a (preliminary) revision of Rivulus in which he synonymizes Owiyeye into Laimosemion !) (Jean Huber,  finalized April 2011, updated February 2012)



Q(36): Why not follow Collier and co-workers American revolutional findings who synonymize Pseudepiplatys into Epiplatys (36)? (Anonymous, USA, September 2010)

A: Yes, that molecular study is very important [ref.: Collier, G.E., W.J. Murphy & M. Espinoza. 2009. Phylogeography of the Genus Epiplatys (Aplocheiloidea: Cyprinodontiformes). Mol. Phylogenetics and Evol., 50 (1): 190-196] and the authors are among the pionneers in the gene techniques named molecular research ; their findings indeed show that the species annulatus is nested within other Epiplatys species groups (and the case of Aphyoplatys would be similar) and the various named subgenera of Epiplatys are not congruent with that phylogenetic tree that is produced by DNA ; however Killi-Data is based on definable units and Collier and co-workers do not go beyond their results except synonymizing Pseudepiplatys and do not give a new systematic analysis of their subunits of Epiplatys with new diagnoses for their new groupings as subgenera, and presently nobody is able to do so, so that it is better to keep the previous situation until a new study with solid diagnoses (and then Killi-Data will follow) ; in other words, if the previous situation is kept, then we have a clear organisation (which may be erroneous), if we synonymize Pseudepiplatys into Epiplatys, then the diagnosis of Epiplatys with its various subgenera does not fit anymore and is unstable, and if we downgrade Pseudepiplatys as a subgenus of Epiplatys, then the diagnoses of the various subgenera of Epiplatys are unstable ; by the way who told you that molecular results were the end of the line : it is a very important technique, but not the ultimate and not a technique more accurate at the generic level than, say, osteology (or morphology) ; and Aarn & Sheperd would not be in agreement who published in 2001 evidence from osteology that annulatus was sufficiently different to deserve a genus status (but they were unable to provide with solid diagnoses for Epiplatys subgenera) and, on the other hand, Costa (but without yet evidence) maintains Pseudepiplatys as a valid genus ; the situation is exactly the same with Aphyosemion subgenera and to define correctly Mesoaphyosemion is not coming soon (and in that case Killi-Data does not yet follow the molecular results by Sonnenberg, a German researcher, therefore nothing to do with that Collier is an American !) (Jean Huber,  finalized September 2010, updated October 2012)



Q(35): The diapauses in Nothobranchius, could you sum up and update the situation (35)? (Anonymous, Italy, January 2009)

A: Well, this is an uneasy question because the studies are rather old (it is very difficult to undertake a study in embryology because measured criteria must be independant and isolated for the experimentation, for example, a recent author has suggested that embyonic development may be influenced by the presence or the absence of adult fishes swimming in the close environment !) ; what is known is that there are pauses in the development of many Killifishes (not only the annual species) but it is not known if those pauses (diapauses) are similar for annual or non-annual, for east-african, west-african or south-american species ; to make things simple (but still accurate), it is possible to distinguish 3 states of diapauses in Nothobranchius and 8 paths to meet them : (1) after 2 to 3 days after spawning, the eggs may enter in a first diapause (Diap. I) or (2) continue to develop up to hatching or (3) to another diapause state (Diap. II), then hatch or (4) (Diap. III) then hatch; (1) after Diap. I, the eggs develop and (5) may directly hatch or (6) enter in Diap. II, then or (7) enter in Diap. III or not, then hatch or (8) enter in Diap. III ; in total, there are 8 situations when the eggs may hatch after no diapause, or 1, 2, 3 diapauses ; the factors influencing these diapauses are complex and our understanding is its infancy ; the probably major factor is the presence or absence of oxygen (diapause I is bound to anaerobic conditions - no oxygen -) ; for example, if the parents spawn in a different substratum than mud (e.g. silt or sand) that is aerobic development may not be stopped by a diapause (that may explain why in some locations fry and juveniles are seen when unexpected) ; temperature and moist during the embryonic development (season) may be important too (to induce or levy diapauses) ; all these (uncertain) factors make that an expert breeder will start with the standard formula (time, temperature of incubation) given by previous breeders, then will vary variables in order to keep all hasard on his side ; fortunately, Watters (2009) has recently published information regarding Nothobranchius diapauses that are exactly what you require ; basically, based on these old studies and his field experience, he thinks that most (say 90%) of Nothobranchius eggs undergo the full 3 diapauses in nature because the weather is seasonal and because the soil is made of vertisols (dark colored, clay rich, but not with kaolin, rather with a sort of clay called montmorillonite, of the smectite group of clays, which swells, falls into cracks, induces alkalinity of water, and develops anaerobic conditions when saturated ; according to him, the major factors to play a role are changes in oxygen, carbon dioxyde and moisture level ; typically, 3 days after spawning, the eggs enter into diapause I (they can stay dormant for many months, even years), from which they exit when the soil loses its anaerobic condition, i.e., weeks after the water has disappeared (cracking soil), to enter a new phase of development of about 5 days, then enter into diapause II and stay there until another change in environment (the onset of the new rains) when they resume development up to the pre-hatch stage, from which stage they will then enter into diapause III (because the first rains wet the soil but do not produce a body of water) and they will stay into that diapause III stage for 2 to 4 months (with a progressive dimunition of yolk sac volume between these 2 lag times) and exit from diapause III (and hatch) when the conditions return to anaerobic with enhanced levels of carbon dioxyde and decreased levels of oxygen (when the standing water has been permanent for 2 to 4 weeks) ; obviouly these observations are not the same for other weather conditions and soils (like in western Africa and South America) and for captivity conditions in aquarium (with peat moss and aerobic conditions which make the whole process much quicker) (Jean Huber,  finalized January 2009, updated August 2009)



Q(34): The new subgenera of Simpsonichthys, should we use them as aquarists (34)? (Stuart Lord, New Zealand, February 2008)

A: Yes, Wilson Costa has recently described (2006) 3 new subgenera of Simpsonichhys ; then, in the 2006 understanding, Simpsonichthys is divided into 5 subgenera, including Spectrolebias, previously considered as a separate genus ; each species of the genus is allocated to either the typical subgenus Simpsonichthys, or Spectrolebias, or one of the three new subgenera, Xenurolebias, Ophthalmolebias, and Hypsolebias ; this is just the result of his own research, and since it is based on scientific evidence, everybody, including aquarists who consider their hobby seriously, should consider this… however, no scientist has ever required that his findings are strictly followed by aquarists when it relates to the subgenus level (it is already very good if the dedicated aquarists use the species name and the population name correctly with no misspelling, and ideal when they use the most accepted genus name  !). Therefore, please use what you feel is appropriate for the subgenus level, and if you use those new subgenera, write them between brackets such as Simpsonichthys (Hypsolebias) ocellatus… and be prepared to change should a new research improvement arise ; there is though one case where you must change because it concerns the genus level : Spectrolebias semiocellatus must be renamed Simpsonichthys semiocellatus or Simpsonichthys (Spectrolebias) semiocellatus… until there is another publication on further research that confirms or change the picture… that's the way research is moving ; as a very practical illustration of the above statements, Costa (2010) has reviewed again those groups and re-established Spectrolebias as a full genus and uplevelled all previous subgenera to full genera inducing a major list of name changes… however, since 2009, Killi-Data is not immediately following research results at the genus level for the sake of stability notably when diagnoses are weak, when no new findings are evidenced, when the move is more based on a splitting attitude (with little value added) ; aquarists will still have to make efforts to memorize new names but only when the change is solid (for example if you follow Costa (2006) then the name will be with subgenus will be Simpsonichthys (Hypsolebias) ocellatus or without subgenus it will be Simpsonichthys ocellatus, if you follow Costa (2010) then the generic name of the previous situation may be added after between brackets, e.g. Hypsolebias ocellatus [Simpsonichthys] (not in-between like for a subgenus) ; the only regrets that could be formulated are that no other researcher with other techniques is publishing on these fishes for generic systematics ! (but there is hope since a Brazilian team around Costa is starting to publish molecular results on South American killifish and that hope came true with a publication by a Brazilian molecular team, without Costa, whose results are not at all in line with Costa's publications and hence give reason to Killi-Data to slow down on generic name changes… until next publication !) (Jean Huber,  finalized February 2008, updated December 2010, October 2012, July 2016)



Q(33): What is the correct year of publication for the description of Nothobranchius cardinalis (33)? (Anonymous, Canada, February 2008)

A: Your question is easy to answer in this specific case (but not in all cases !) ; the reference is Journal of the American Killifish Association 40 (5 & 6, September-October-November-December 2007): 128-145, 9 figs., 3 tabs. Description of Nothobranchius cardinalis spec. nov. (Cyprinodontiformes: Aplocheilidae), an annual fish from the Mbwemkuru River basin, Tanzania. Brian R. Watters, Barry J. Cooper and Rudolf H. Wildekamp. With the new Code of Nomenclature, valid from January 1. 2000, rules are simple with no exceptions (this is not the same story for names described before): the official date of publication is the day, month, year, when the printed magazine is actually distributed to its public target (i.e. people or/and organisations who paid the fee for it). Editors know this and they try to make sure that the date of publication printed on the printed magazine is the same as the actual date of publication according to nomenclature… but sometimes there are events that make the actual publication not in line with the printed date on cover (usually delayed because of the printer, or because of a strike in distribution, or because of a last minute failure). These problems may have 2 adverse consequences : (1) when the year end is concerned, for the official year of description, (2) when the same taxon is described with 2 different names by 2 sets of describers, in order to decide for the elder and only valid name (e.g., the case of Moema quiii vs. Moema ortegai, the latter having been actually distributed after the former… that had actually been distributed before its printed date of publication !). Your case of N. cardinalis is concerned with the first issue : the JAKA magazine states 2007 on cover, but it has been delayed and the distribution could only occur in January 2008 (and Brian Watters, one of the authors, has immediately sent a note to other researchers to inform them about this) : therefore the species must be labelled Nothobranchius cardinalis  Watters, Cooper & Wildekamp, 2008 (no blame on JAKA about this, it is frequent and does not only concern aquarium magazines) ; moreover, the Internet may today bring additional confusion : the date of publication on Internet is not the good one by nomenclature (only the actual distribution date of the printed material is the good one) ; even magazines that take preventive actions by issuing their last number well ahead of the year end may face pitfalls : at about the same time as cardinalis, that problem occurred with Ichthyological Exploration of Freshwaters with the last issue usually published on December 1. of each year and the abstracts and electronic text available on the Internet at that date… very bad luck this year, because their printer faced a major collapse and the actual distribution was only achieved on January 9. 2008 (instead of December 1. 2007 !) and 6 new killifish names (by Costa and by Sonnenberg) are then moved to 2008 (Sonnenberg, pers. comm.). This is not a big deal and the correct information is given in Killi-Data.  (Jean Huber,  finalized February 2008)



Q(32): What is the relationship between Oryzias melastigma (melanostigma?) and Aplocheilus panchax? (Gary Hoover, USA, April 2007)

A: Aplocheilus melastigmus has been first described by McClelland, in 1839, in the genus Aplocheilus. This observation may be seen as anecdotal but it is not : in more recent times (i.e., the beginning of the 20th century), species from Asia were described in Haplochilus or Panchax or Oryzias, because those genera were mixed and their definition ambiguous (believe me, many articles have then been written on the issue, often with polemics), until the matter was progressively settled after 1945 and 2 valid genera with their assigned species were cleared out either in Aplocheilus or in Oryzias. For example, the common lampeye aquarium fish from Java island was moved from Aplocheilus javanicus Bleeker, 1854 to Oryzias javanicus. Similarly (without much thought), Aplocheilus melastigmus was moved to Oryzias (and erroneously the species ending was changed in melastigma, while it must have stayed constant because the 2 genera are of masculine gender!). The name means : with black (from melas, melaina) pricked mark (from Greek: stigma), in reference to the male spot of the Dorsal fin. The type locality is Calcutta, northeastern India (without more details). And in this region, both true Oryzias and Aplocheilus species (Apl. panchax) live (and are often collected together), which you would agree makes the matter even more complicated (un-needed!). No types of melastigmus have been disclosed (unfortunately). The matter was frozen : conservatively scientists and aquarists consistently used the name Oryzias melastigma for a lampeye fish of costal eastern India, up until Roberts (1998). The renown American ichthyologist, within a global review of the genus Oryzias, has demonstrated that McClelland's melastigmus species cannot be an Oryzias member because the character linked with the name (the black pricked marking) is unknown in Oryzias throughout its range ; hence the species was hypothetically synonymized with Aplocheilus panchax, on the basis of the single dorsal spot, only, and of its origin. The common lampeye fish of aquarists that was incorrectly named Oryzias melastigma was renamed as Oryzias dancena (described in Cyprinus by Hamilton in 1822), an even older name with its type locality as estuary below Calcutta, India (and also no types known!). According to Eschmeyer (online), its distribution is India, Bangladesh and Myanmar, and habitat is freshwater, brackish. In conclusion (since 1998, only… it might be challenged by another scientific study!), the trade aquarium lampeye fish from India should be named Oryzias dancena (Hamilton, 1822) and the killifish from northeastern India is still named Aplocheilus panchax (Hamilton, 1822), with melastigmus being one of its (many) junior synonyms. (Jean Huber,  finalized April 2007)



Q(31): Why Plataplochilus valid today and as a synonym of Procatopus in the past ? (anonymous, Spain, December 2005)

A: This is an important question and you are right to raise it because no major new data has emerged since 40 years or so on the subject. Lambert (1963) and Lambert & Clausen (1967) revalidated and redefined Plataplochilus on the basis of micromorphological characters, separate from Procatopus ; but this view has been challenged and not accepted by all researchers and for a long period the 2 options of validity or synonymization were maintained. Only the systematic philosophy and the tools to analyse the characters explain the 2 options in 40 years. The systematic philosophy is today more like splitting than lumping, i.e. considering many valid genera than few, and this is just a fact that cannot be avoided (and in the future it may change again) : in the past, the tubular system on the head was thought as a variation from the open system and this variation was considered as a minor factor in the evolution (phylogeny), all the more that intermediates were found and thus Plataplochilus was synonymized with Procatopus (e.g., by Huber, 1981) ; today those characters are put in a digital matrix with a 0 or 1 state and the results are that both genera belong to the same phylogenetic line (= have the same ancestor) but are sufficiently separate and thus Plataplochilus are separated from Procatopus (e.g., by Huber, 1999, Ghedotti, 2000). Still the major difference is that Lambert & Clausen thought Plataplochilus was not belonging to the same lineage than Procatopus and thus you see things have dramatically changed actually : today Plataplochilus, Procatopus, Hypsopanchax, Hylopanchax, Lamprichthys, Rhexipanchax and Pantanodon are hypothesized to be related while in the past Lamprichthys and Pantanodon were even set in 2 different subfamilies, themselves isolated from Plataplochilus-Procatopus… the next issue is a molecular analysis of the deep-bodied lampeyes to confirm, or not, the present status. On aquaristic grounds, there is no problem to keep using the genus name Procatopus for all, but to communicate better with your fellows, just state Procatopus miltotaenia (Plataplochilus) or Plataplochilus miltotaenia (Procatopus) so that everybody understands you !  (Jean Huber, finalized January 2006, November 2009)



Q(30): Fluviphylax obscurum vs. Fluviphylax obscurus. On page 123 of the original description, Costa indicated the specific name obscurum is to be treated as an adjective, so he should have used obscurus. According to the ICZN Code the specific name must be emended. In the same work he used zonatus also as an adjective with the correct ending. What is your opinion? (Richard van der Laan, May 2005)

A: Indeed, the original spelling of the name, obscurum, has been a problem: obscurum, following the etymology (darkness), or, obscurus, according to the status of adjective also given in the description (Fluviphylax having a masculine gender)  ; it should be an adjective (from Latin: obscurus, a, um, meaning dark), in reference to the dark pattern of sides in larger specimens, with strong melanism. Hence the correct spelling is Fluv. obscurus (Wilson Costa, June 2005)



Q(29): At what point in time do researchers consider variability as a stable character : for example, why Aphyosemion primigenium GBN 88/10 has not been described as a new species yet ? (Anonymous, Portugal, March 2005)

A: Your question is important but uneasy ! Let's first go back to basics : a species is represented by a group of populations that share similar characters, notably genetic homogeneity, and that can be distinguished from another group of populations. For Killifish, genetic sterility cannot be a distinguishing character, as it should be following Mayr's definition of the biological species, because then there will be, for example, 1000 species in Aphyosemion, most of them being not distinguishable. Therefore, for Killifish, the researcher must find a stable character that separates a group of population from another. Obviously it is not easy to determine if a population constitutes a stable variant that may be candidate for a different name, or a simple variation within a continuum (grading differences). Experience plays a major role in assessing that issue, and, of course, the availability of many collections in the concerned region. Your example for primigenium is characteristic : the species is diagnosed by 3-5 series of longitudinal red lines on male sides, like striatum, and a "closed" pattern of Caudal fin (a blue circumcaudal border and a red parallel sub-border). The population coded GBN 88/10 is different because it shows only 1-2 series on sides and it is collected at the extreme south of the distribution of primigenium in Gabon, near the Congo border. Is it a simple variation at the population level, or is it the first population of a group, mainly living in Congo, with a single line ? That's the question ! Only collections in that region can allow to answer (Jean Huber,  finalized March 2005)



Q(28): From a Peruvian perspective, a) What altitude limits are Rivulus found at?, b) At higher altitudes, what water temperatures would likely limit Rivulus presence?, c) What kilometer distances or areas do a given species normally cover?, d) Are species limited to certain watersheds or do they often spread over different drainages ? (Lance Peck, October 2004)

A: Rivulus distribution patterns are irregularly known : most of our knowledge is from the Iquitos area in the low lands of upper Amazon river and from few localities also in the low lands and in high Andean valleys too. Rivulus species can be collected from sea level to an altitude of 1200-1500 m. Water temperature is not a limit (in Argentina, some are collected near to freezing). They are not available in the Andes (4000 meters) because when the mountain uplifted (about 30 Million years ago) they were not there, but their then marine relatives, today in the genus Orestias (40 species in Lake Titicaca) are there. All Rivulus species trapped at the foot of high Andean valleys have been found new (1 per valley): Puyo (Riv. jucundus), Tayuza (Riv. monticola), Tingo Maria (Riv. derhami). Kilometer distance … you probably mean distribution range : it depends on past ecological conditions, not on present river systems. To make it simple, during the last glacial peaks, species were isolated in refugia, then began to expand again with forest (and rain), up to the range of a similar species (from the same group) originating from another refugium (see Huber, 1998 "Comparison"). This can be 400+ kms in Paraguay and probably much less in Peru with a very high speciation. For example the fauna in Pucallpa is different from Puerto Maldonado. Besides, the fact that 2 distinct new Aphyolebias are found in the Reserve, about 150 km away from Puerto Maldonado, implies that Rivulus, there, should be different too. In 2011 (late), Valdesalici & Schindler described Rivulus parlettei from southern Peru and they mentioned another refugium recently discovered in the lowlands !(Jean Huber,  finalized November 2004, January 2012)



Q(27): Which Killifish is, a) a fish suited for an experienced aquarist but with no prior experience with killies, and b) reasonably available, and c) preferably not seasonal for an Orinoco-biotope tank ?(Ed Prust, October 2004)

A: The following Killifish are known from Venezuela (plus the other countries of distribution): Austrofundulus limnaeus [Colombia; Guiana; Venezuela], Austrof. transilis [Venezuela], Cyprinodon dearborni [Colombia; Ned. Antillas; Venezuela], Fluviphylax pygmaeus [Brasil (Amazonas); Colombia; Peru; Venezuela], Gnatholebias hoignei [Venezuela], Gnat. zonatus [Colombia; Venezuela], Kryptolebias marmoratus [Belize; Brasil (Atlantic); Cuba; Guadeloupe; Guyane; Honduras; Jamaïca; Martinique; Mexico; Ned. Antillas; Puerto Rico; Santo Domingo; USA (Florida); Venezuela], Micromoema xiphophora [Venezuela], Rachovia brevis [Colombia; Venezuela], Rac. hummelincki [Colombia; Venezuela], Rac. maculipinnis [Colombia; Venezuela], Rac. pyropunctata [Venezuela], Rac. stellifer [Venezuela], Renova oscari [Venezuela], Rivulus (Laim.?) gransabanae [Venezuela], Riv. (Laim.?) lyricauda [Venezuela], Riv. (Laim.?) nicoi [Venezuela], Riv. (Laim.?) tecminae [Venezuela], Riv. (Od.) hartii [Ned. Antillas; Venezuela], Riv. (Od.) immaculatus [Venezuela], Riv. (Od.?) deltaphilus [Venezuela], Riv. (Od.?) caurae [Venezuela], Terranatos dolichopterus [Venezuela], Tomeurus gracilis [Brasil (Amapa); Guiana; Guyane; Suriname; Venezuela] . Because the Orinoco river is so important in the country, all species are reported to live herein or should live not too far. If you do not want annuals or brackish water species, the list is limited to Fluv. pygmaeus, Tom. gracilis (both in large rivers), Rivulus gransabanae, Riv. lyricauda, Riv. nicoi (unkown live, maybe annual), Riv. tecminae, Riv. hartii, Riv. immaculatus, Riv. deltaphilus, Riv. caurae. In terms of aquarium trade availability, none is readily available except hartii (but it is probably the less sure for Orinoco). Within Killifish Associations, you should find (with some difficulty) the newly described caurae and also deltaphilus, and hypothetically the very rare gransabanae, tecminae, immaculatus and lyricauda, plus the gorgeous and newly described Rivulus staecki. (Jean Huber,  finalized October 2004, updated May 2011)



Q(26): Regarding Epiplatys lokoensis, it is considered as a valid distinctive species in Germany. Nobody who has seen the fish can believe what Romand wrote on its identity with bifasciatus or barmoiensis, therefore why do you follow him in Killi-Data? (Anonymous, October 2004)

A: The case of Epiplatys lokoensis is not an easy one. The fish has been described as new on the basis of a few juvenile specimens [Berkenkamp,H.O. & V. Etzel. 1978a. Epiplatys lokoensis spec. nov. eine neue Hechtlingsart des Überartenkreis Ep. bifasciatus aus Sierra Leone. D.K.G. (Deutsche Killifisch Gem.) J., 10 (9): 152-162, 6 figs.]. Nobody has ever published an account on adult specimens based on topotypes or material from other locations. Raymond Romand [1980b. Comments on Epiplatys lokoensis Berkenkamp & Etzel (1978) from Sierra Leone (Pisces, Cyprinodontidae). Rev. Zool. Bot. Afr., 94 (3): 591-599, 9 figs., tab.] has criticized the description and after a detailed review in an international publication has concluded that lokoensis is a synonym of barmoiensis or an hybrid between bifasciatus and barmoiensis. Wildekamp [1996. A World of Killies. Atlas of the Oviparous Cyprinodontiform Fishes of the World. Vol. 3. Amer. Killifish Assoc. Publ.: 330pp, figs.] support the point of view of a valid species, but it is only an opinion based on the analysis of the 2 papers. This is respectable (and nobody will quarrel about Wildekamp's expertise), but this is not the way Killi-Data operates. Killi-Data does not take a decision or do not derive an opinion from both papers (and never does that, in any case). Killi-Data takes into account the last published evidence on the case (i.e. scientifically argumented and evidenced) and in this case it is simply Romand's (not Wildekamp's). Berkenkamp and Etzel never answered to Romand's arguments and thus they are not acknowledged as the last evidence in Killi-Data. Is it possible to move the case in Killi-Data ? Of course, yes : by a new published evidence. For example, by a new study of the types showing new stable distinctive characters for the species or by the study of new material from the type locality or from new collections. (Jean Huber,  finalized October 2004)



Q(25): Regarding Chromaphyosemion, it is considered as a valid genus, distinct from Aphyosemion, in Germany ; this move is based on a proposal by a German Scientist, Rainer Sonnenberg, in a scientific publication. Why don't you follow that move ? (Anonymous, June 2004, Pohlmann, September, 2005)

A: An easy answer on a difficult subject. Indeed, Rainer Sonnenberg (a German Ph.D. student) has proposed to erect Chromaphyosemion as a valid genus [Sonnenberg, R. 2000. The Distribution of Chromaphyosemion Radda, 1971 (Teleostei: Cyprinodontiformes) on the coastal Plains of West and Central Africa. Bonn zool. Monogr., (Proc. 4th Symp.): 79-94, 1 fig., 1 pl.] and this is a solid publication. Killi-Data is not partial by not following (yet) that move, it is just expecting the final results of Sonnenberg's research in his thesis and notably how he places and diagnoses Chromaphyosemion from all other present Aphyosemion subgenera. For example, it is unsatisfactory to erect Chromaphyosemion alone and leave Diapteron, Kathetys, Mesoaphyosemion or Raddaella untouched because the molecular evidence (DNA) places it well nested in Aphyosemion, within the phylogenetic tree: are they still subgenera or should they be distinct genera, too ? all of them or some, only ? You see, your question is very important, but a little bit premature… be patient : Killi-Data will, no doubt, follow his conclusions when all current Aphyosemion subgenera are rediagnosed. New development : in 2005-2006, it is going to be a major problem because Collier has announced that in his molecular study to be published soon, all are Aphyosemion ! Names are just a way of communication… if everybody creates and uses its own language, it is the Babel tower ! Personally I have no pre-defined idea with Killi-Data, I follow the latest scientific evidence… but there must be one (Legros et al. 2005, when describing melanogaster and punctulatum in the genus Chromaphyosemion did not bring anything new on this subject).(Jean Huber,  finalized September 2004, updated October 2005)



Q(24): Regarding your database entry for Terranatos dolichopterus : is it correct when it states that this fish is found in neutral to alkaline water ? Everything I have seen in the hobby thus far has indicated a preference for acid to neutral water.(Nick Anderson, June 2004)

A: A difficult question ! All savannah populations of dolichopterus are probably on clay soil, then neutral pH or somewhat alkaline. Like most annuals. But it can be hypothesized that the forest population of Isla Raton dwells acid waters ! Our knowledge on biotopes of annuals is still poor on a year long basis (apart from spotty measurements). Water is stagnant and its level is gradually declining from peak of rainy season : at beginning, it can be inferred that water characteristics are close to those of rain water (but pH should not stay acid for more than a few hours after rain is falling from the sky), then the influence of soil increases more and more with oligo-elements and minerals being captured ; even for forest non annuals in "black waters", it has been recently shown that to use markedly acid waters on a permanent basis (e.g., pH< 5, for Aphyosemion, Rivulus… ) was an error ; a final consideration for all forest dwellers is the influence from flora : all plants produce pollens (tons of them) which often end up in waters where Killies live ; it can easily be hypothesized that pollens influence a lot the micro-characteristics of the shallow slow-moving waters (see Huber, 1998 : "Comparison"), and those pollens have been distinctive between Old World and New World since the dawn of Killifish… (Jean Huber,  finalized June 2004)



Q(23): What is the current list of DNA publications on Killifish ? (Steffen Hellner, February 2004)

A: Molecular Biology, as it is globally understood, encompasses studies of portions of genome ; it may be part of nucleus (DNA) or of mitochondria (DNA, RNA), the latter being frequently preferred because genes are known to evolve more slowly; this is a promising technique : it gives phylogenetic answers to issues at the generic levels, superspecies (or clades) levels ; however, it is not the ultimate technique because it does not give answers to all questions, it may give different answers than other established techniques, it has artefacts, it has limitations (haplotypes) and it raised important concerns with the case of lacustrine speciation (Cichlids) ; the first paper on Killifish is only dated 1993 : not many species have been studied yet and for a very small portion of their genomes. There is also an asset which is shared with modern morphological and osteological techniques : data are poured in a matrix that a computer analyses and these data are available to other researchers (to add species or characters) ; attempts to gather data from molecular, morphological and osteological have given disappointing results, yet. Here is the list of papers, you are looking for :
* Bernardi, G. 1997. Molecular Phylogeny of the Fundulidae (Teleostei, Cyprinodontiformes) based on the Cytochrome b Gene. In: Kocher T.D. & C.A. Stiepen. Molecular Systematics of Fishes. Academic Press Ltd, Harcourt Brace & Co., 314pp.
* Bernardi, G. & D.A. Powers. 1995. Phylogenetic Relationships among Nine Species from the Genus Fundulus (Cyprinodontiformes, Fundulidae) inferred from Sequences of the Cytochrome B Gene. Copeia, (2): 469-473, 1 fig., appx.
* Bernardi, G. & D. Talley. 2000. Molecular Evidence for reduced Dispersal in the coastal California Killifish, Fundulus parvipinnis. J. Exp. Mar. Biol. Ecol., 255: 187-199, figs.
* Echelle, A.A., R.A. Van den Bussche, T.P. Malloy Jr, M.L. Hatnie & C.O. Minkley. 2000. Mitochondrial DNA Variation in Pupfishes assigned to the Species Cyprinodon macularius (Atherinomorpha, Cyprinodontidae): taxonomic Implications and Conservation Genetics. Copeia, (2): 353-364, 2 figs., 2 tabs.
* Garcia, G., G. Wlasiuk & E.P. Lassa. 2000. High Levels of mitochondrial Cytochrome b divergence in Annual Killifishes of the Genus Cynolebias (Cyprinodontiformes, Rivulidae). J. Linnean Soc. London Zool., 129, 93-110, 1 tab., 6 figs.
* Ghedotti, M.J. & M.J. Grose. 1997. Phylogenetic Relationships of the Fundulus nottii Species Group (Fundulidae, Cyprinodontiformes) as inferred from the Cytochrome b Gene. Copeia, (4) December: 858-862, fig.
* Grant, E.C. & B.R. Riddle. 1995. Are the endangered Springfish (Crenichthys Hubbs) and Poolfish (Empetrichthys Gilbert) Fundulins or Goodeids?: a Mitochondrial DNA Assessment. Copeia, (1), Feb.: 209-212, 2 figs.
* Hrbek, T. & A. Larson. 1999. The Evolution of Diapause in the Killifish Family Rivulidae (Atherinomorpha, Cyprinodontiformes): a molecular phylogenetic and biogeographic Perspective. Evolution, Lawrence, Kansas, 53: 1200-1216, 7 figs., 3 tabs.
* Hrbek, T. & A. Meyer. 2003. Closing of the Tethys Sea and the Phylogeny of Eurasian Killifishes (Cyprinodontiformes: Cyprinodontidae). J. Evol. Biol., 16: 1-20, 7 figs., 3 tabs.
* Kreiser, B.R. 2001. Mitochondrial Cytochrome b Sequences Support Recognition of Two Cryptic Species of Plains Killifish, Fundulus zebrinus and Fundulus kansae. Am. Midl. Nat., 146: 199-209, 3 figs., 3 tabs.
* Meyer, A. & C. Lydeard. 1993. The Evolution of Copulatory Organs, internal Fertilization, Placentae and Viviparity in Killifishes (Cyprinodontiformes) inferred from a DNA Phylogeny of the Tyrosine Kinase Gene X-src. Proc. R. Soc. Lond., B, 254: 153-162, 4 figs.
* Murphy, W.J. & G.E. Collier. 1996. Phylogenetic Relationships within the Aplocheiloid Fish Genus Rivulus (Cyprinodontiformes, Rivulidae): Implications for Caribbean and Central American Biogeography. Mol. Biol. Evol., 13 (5): 642-649, 3 figs.
* Murphy, W.J. & G.E. Collier. 1997. A Molecular Phylogeny for Aplocheiloid Fishes (Atherinomorpha, Cyprinodontiformes): The role of Vicariance and the Origins of Annualism. Mol. Biol. Evol., 14 (8): 790-799, 6 figs., 2 Tabs.
* Murphy, W.J. & G.E. Collier. 1999. Phylogenetic Relationships of African Killifishes in the Genera Aphyosemion and Fundulopanchax inferred from mitochondrial DNA Sequences. Mol. Phylogenetics and Evol., 11 (3) (April): 351-360, 5 figs.
* Murphy, W.J., T.N.P. Nguyen, E.B. Taylor & G.E. Collier. 1999. Mitochondrial DNA Phylogeny of West African Aplocheiloid Killifishes (Cyprinodontiformes, Aplocheilidae). Mol. Phylogenetics and Evol., 11 (3) (April): 343-350, 3 figs.
* Murphy, W.J., J.E. Thomerson & G.E. Collier. 1999. Phylogeny of the Neotropical Killifish Family Rivulidae (Cyprinodontiformes, Aplocheiloidei) Inferred from Mitochondrial DNA Sequences. Mol. Phylogenetics and Evol., 13 (2) (November): 289-301, 5 figs.
* Parker, A. 1997. Combining Molecular and Morphological Data in Fish Systematics: Examples from the Cyprinodontiformes. In: Kocher, T.D. & C.A. Stiepen. Molecular Systematics of Fishes. Academic Press Ltd, Harcourt Brace & Co., 314pp. 
* Parker, A. & I.L. Kornfield. 1995. Molecular Perspective on Evolution and Zoogeography of Cyprinodontid Killifishes (Teleostei; Atherinomorpha). Copeia, (1): 8-21, figs.
* Perdices, A., J.A. Carmona, C. Fernandez-Delgado & I. Doadrio. 2001. Nuclear and mitochondrial Data reveal high genetic Divergence among Atlantic and Mediterranean Populations of the Iberian Killifish Aphanius iberus (Teleostei: Cyprinodontidae). Heredity, 87: 314-324.
* Strecker, U., C.G. Meyer, C. Sturmbauer & H. Wilkens. 1996. Genetic Divergence in an extremely young Species Flock in Mexico, formed by the Genus Cyprinodon (Cyprinodontiformes, Teleostei). Mol. Phylogenetics and Evol., 6 (1): 143-149. (Jean Huber,  finalized February 2004)



Q(22): What is your view about Fp. gresensi and other mirabilis, moensis, traudeae, intermittens : species or subspecies ? (Anonymous, December 2003)

A: What a difficult question ! Indeed, Berkenkamp has just described (2003) as Fundulopanchax gresensi the population of Takwai previously identified as aff. mirabilis and up-levelled as distinct species all subspecies of that species, while Radda had described his 4 names as subspecies and did not change their status over time. The first author gives the following reasons for his move : all populations breed true to their characteristics and they are geographically isolated. On the contrary the second author used the subspecies level because all populations readily interbreed with complete fertility and are vicariant (geographical neighbors). While there are some definitions of the species status that reach (with difficulty) international consensus, there is none for the subspecies status. Some schools believe that a subspecies is a group of populations that are mutually inter-fertile and that replace each other geographically. However, the key issue is whether those populations can meet each other or not. If no, some schools of thought believe that they are true distinct species then (many examples are known with components of the genus Cyprinodon in the US). If you are an experienced "killifish nut", then you know that things are never simple with these fishes : for example, Fp. oeseri was readily accepted as a distinct species because it was collected in an island separated from mainland Africa where its counterpart, marmoratus, is collected, even if both do interbreed easily ; however, bad luck (!) a new population near Mundemba has been found sympatrically with marmoratus that is strikingly similar to oeseri. What can we do with gresensi, mirabilis and alike ? If we had detailed geographical micro-distribution (borders) and DNA data, maybe we could take a rational decision that would not be too fragile, but this is not the case. To decide on the basis of the present knowledge for Berkenkamp or for Radda would be respectable (you can do that), but opinion-based (not evidence-based) and this is not Killi-Data philosophy (and I do not want to de-stabilize the board of scientific advisors by asking them now to rule out this conflicting case). Therefore, the best proposal I can think of, presently, is to provisionally accept the options of each author independently (gresensi has a distinct species : Berkenkamp speaks of semi-species "status nascendi", the others by Radda, conserved as subspecies of mirabilis). Some people may find awkward this proposal or even the worse they could think of… May I propose another solution for your fish exchanges (which I cannot use in Killi-Data online, because a choice has to be done with computerized stuff) ? This is just to put the subspecies level as an option, i.e. between parenthesis ; for example Fp. gresensi (mirabilis), Fp. moensis (mirabilis). Salomon would surely approve ! Recently a new study has brought more light on the issue [Collier, G.E. 2010. The Genus Fundulopanchax: Taxonomic History and Molecular Phylogeny. J. Amer. Killifish Assoc., 43 (1): 3-24, 15 figs.] and it is mostly followed by Killi-Data : with molecular data, it appears that mirabilis, moensis, gresensi deserve the full species status and that traudeae, intermittens are not sufficiently distinct from mirabilis (they are temporarily kept as subspecies of mirabilis pending further field surveys) (Jean Huber,  finalized December 2003, updated November 2010).



Q(21): What is valid for Killi-Data, in the case of Callopanchax huwaldi ? Valid for DNA, invalid for morphology ? Berkenkamp or not Berkenkamp ? (Reinhardt Kämpfe, November 2003)

A: Killi-Data is evidence based, not opinion based. This means that as soon as a publication bringing a scientific demonstration on a systematic issue is available, Killi-Data accepts it. If this publication reflects its author's opinion without rational argumentation, then it is not taken into account, even if the author is followed by the aquarists association of his country. To answer to your specific question on Cal. huwaldi, let's enlarge it to another -difficult issue- the status of Aphyosemion kekemense, to show that the previous and present position of Killi-Data is not against Berkenkamp for huwaldi and in favor of Scheel for kekemense (and of course, not "all the time" against Berkenkamp). Cal. huwaldi, soon after its description, has been synonymised by many authors on the basis of the variability of color pattern and of the sympatric occurrence of color phases (blue and red) in occidentalis and toddi, and Killi-Data followed that stance. Within a DNA work, Murphy et al. (1999) showed that huwaldi is a genetically distinct species from occidentalis and toddi and that the fish named "sp. Guinea" was also genetically distinct, but their paper had no systematic purpose for these species and then Killi-Data did not change the status of these species. Now, Berkenkamp & Etzel have just published a paper [2003, D.K.G. (Deutsche Killifisch Gem.) J., 35 (6): 205-215, 7 figs., 4 tabs.] bringing a comparative diagnosis of the 4 Callopanchax species and proposing to revalidate huwaldi on the basis of the female color pattern (new diagnosis); this has been taken into account and immediately accepted in Killi-Data. Note that things are not as simple as that, because they do not consider the "sp. Guinea" has valid, but include it in occidentalis, to the contrary of DNA results ! The case of kekemense is similar. The description by Radda, A.C. & J.J. Scheel, in 1975, is scientifically solid with a rational diagnosis and a distinct genotype ; some authors, but others wrote the contrary, mentioned that they were of the opinion that it is not valid because there were other populations with many bars scattered in the distribution of bualanum (or elberti if you prefer), however, no publication has brought scientific evidence to support this (or the contrary) ; therefore, Killi-Data maintains the valid status of kekemense, until the availability of a systematic revision of these taxa with a rational argumentation (notably a comparative diagnosis). These 2 cases have been brought to exemplify the process of decision of a valid status or not in Killi-Data, i.e. distinction between evidence-based and opinion-based. A final word to mention that Killi-Data does not judge the value of the published evidence because that would introduce an opinion in the process : time will tell … and also present evidence can be broken by future evidence ! A new species described by a professional or amateur researcher in a scientific International magazine with anonymous reviewers and with a solid diagnosis is readily accepted as valid, but this may not be forever, whoever is the author. All critics that were brought to Berkenkamp, including by Germans themselves (see Zeitschrift für Fischkunde, 6 (1): 114-115, by Schindler, I. 2003, on Rivulus villwocki), emphasized that many of his descriptions lack a formal and solid diagnosis, therefore are not fully evidence-based, despite his hard work and sincerity. With at least 2 notable exceptions, those descriptions published in Tervuren Museum scientific magazine "Rev. Zool. Bot. Afr." for heinemanni and hildegardae (Jean Huber,  finalized December 2003).



Q(20): Regarding Rivulus ornatus, I have seen some renown aquarists, Wim Suyker and Henri de Bruyn, who name this species as "ex ornatus". Is this information really correct. Has the R. ornatus taxon been revised or is to be revised actually? (Sebastian Calero, August 2003)

A: This is a complicated case and a confusion due to the simultaneous publications of 2 authors (end of 1992 and beginning of 1993, without knowing the project of each other): my Rivulus book (Huber, J.H. 1992. Review of Rivulus. Ecobiogeography - Relationships. Cybium Suppl., Société Française d'Ichtyologie Publ.: 586 pp., 40 pls., 85 figs., 8 tabs, 13 maps.) where I designate lectotypes of ornatus and obscurus and Costa's paper in DKG Journal (Costa, W.J.E.M. 1993. Zur Identität und Verbreitung von Rivulus ornatus und Rivulus punctatus. D.K.G. (Deutsche Killifisch Gem.) J., 25 (3): 44-46, 2 figs.) where he mentions that the true ornatus is obscurus according to material from near Manaus (3.130S, 60.020W) and the aquarium ornatus is then an unknown species, without knowing my lectotype designation for ornatus from Silves, i.e. not in the area of Manaus. This is unfortunate and lies only in the lack of communication between Costa and Huber. But the problem comes from the fact that in the descriptions by Garman (1895), he used several lots for ornatus with 3 different and distant localities and one lot for obscurus with a type locality near Manaus, identical with one of the 3 for ornatus: in details for ornatus, the 3 localities are Silves, Lake Saraca, N. Brasil 2.880S, 58.350W; Parana do Janauari, Brasil 3.200S, 60.080W (near Manaus); Lago Cudajas, now L. Badajos, N. Brasil 3.250S, 62.780W and for obscurus, Lago Januaria, vicinity of Manaus, N. Brasil 3.200S, 60.080W. By selecting Silves as the official type locality for ornatus, without knowing Costa's research, Huber made Costa's publication inappropriate because Costa allocated ornatus to a fish we do not know if it is identical with the fish from Silves. Therefore the best is to stick conservative (i.e. my view, keeping valid both names, not because it is better, but because it is anterior to Costa's by a few months and because it is legally correct according to ICZN rules) until live material is collected from the 2 fixed type localities, Silves for ornatus, and Januaria, near Manaus for obscurus and the issue can be fixed. Costa may be right or may be wrong, this would not be contradictory with Huber's designation: we just have to wait for material from Silves. If he is right (the 2 fish are identical), then (Bill Eschmeyer, pers. comm. November 2004), he will be considered by ICZN rules as the first reviser, and among the 2 taxa described by Garman in the same publication, ornatus will have the priority and obscurus will be a synonym. If he is wrong (the 2 taxa correspond to 2 different fish and species), then Huber's proposal to consider both names as distinct will be correct. Finally, another issue is the identification of the aquarium strains of ornatus: one as an aquarium import from Obidos (?), lower Amazon (NSC-2), Brasil, 1.920S, 55.520W and the other from Padre Isla, Iquitos, Peru, 3.620S, 73.700W. Both are far from the type locality and may or may not represent the true ornatus: it is only by conservatism that it is advised to keep the name ornatus for them. For obscurus, the situation is a little bit clearer: Garman described his taxon on the basis of a female, but collections in the Manaus area have shown that a very colorful male fish with a black mid-side band, related to R. atratus, had a female with a distinct pattern similar with the one described by Garman and therefore has been identified, probably correctly, as obscurus! (Jean Huber, finalized, September 2003; updated November 2004).



Q(19): After the cautious reading of the description of Nothobranchius nubaensis, there are concerns about its validity: what is your opinion on a point concerning the ICZN rules on nomenclature. Seemingly types have to be designated (and deposited) but in this case they have not. Bellemans only indicates that this will happen in future. This poses the following questions: 1. Does the lack of type material invalidate the description? 2. If so, what is now the status of the proposed name and is it still available if Bellemans re-described the fish in future?  3. If type material is subsequently deposited, does the original paper then become a valid description ? Apart from deposition, there is also the question of designation: Isn't it compulsory under ICZN rules. Bellemans does not do this and in fact quite clearly states that "types will be designated" He therefore appears to be against the Rules, doesn't he? (Anonymous, July 2003)

A: No problem, the answer lies in the ICZN book. The deposition of types for a new species is recommended upon description but not mandatory. Therefore the name nubaensis is said "available" regarding that issue, i.e. the description is valid in terms of nomenclature (provided -which is the case- that the compulsory data are given for the description, such as a full diagnosis). In addition, Bellemans states that he will deposit types in the future and gives details on where and why he delays the deposition. When Bellemans publishes a full description with the types designation, it will be a new scientific paper (some people speak of a re-description to acknowledge its importance), but the original description is single, i.e. that just published in the BKA separatum. If you scrutinize the data base of Killi-Data online, you'll see that quite a number of species-taxon, notably old, does not hold types. It is not a big issue when the type locality is well known (the risk -very seldom- is only with cryptic species that are sympatric, such as escherichi sensu Seegers and striatum, probably). If the type locality is not given, then neotypes should be designated from a locality with specimens that fit best with the description (that's what I did with Aphyosemion ahli). The designation of neotypes may be risky though: everybody knows that Seegers designated neotypes for Rivulus santensis from Rio de Janeiro, because they looked identical to those from the type locality in Santos. But, Costa later described R. janeiroensis from Rio, pushing Seegers's neotypes to invalidity. No, the designation, like the deposition, of types is not mandatory: it is strongly recommended but that does not make a rule. In the ICZN code there is a clear distinction between rules and recommendations. The new ICZN code (2000) which prevails for the nubaensis description keeps the duality of rules and recommendations. Besides no Institution will accept to give a number for types (i.e. designation), without having them at hands or in the hands of another institution. Therefore, yes, nubaensis is an available name according to nomenclature and Bellemans appears to be perfectly in line with the rules. Finally it is also advised that the publications (especially in magazines with small distribution, such as BKA Killi News), be sent by their editors to at least 5 international Institutions (and that had been done, to our best knowledge) ; however, still in 2008, i.e., 5 years after the provisional description Bellemans had not proceeded further with the full description and had not deposited the types as mentioned in the description ; therefore in 2009, Valdesalici, Bellemans, Kardashev & Golubtsov published a full description of nubaensis, mentioning it was provisionally described by Bellemans (2003), but this name is not available, because (1) the mention of a provisional description may questionably be considered to violate ICZN Article 16.1 (error 15.1), (2) no type has been designated in the paper (and deposited in MRAC between 2003 and 2009), the author rather states that types will be designated (ICZN Article 16.4.1 of new code of 1999 clearly requires an explicit fixation of a holotype, or syntypes, even if not immediately deposited) and several specimens are mentioned without giving numbers ; these reasons would not be sustainable (the first reason is even not acknowledgeable) and Wildekamp (also Huber) would be correct if Bellemans would not have been part of the quoted new describers, because he could have deposited a single type (in MRAC) even after, as announced in the provisional description (Bellemans's loose attitude is then a pity) ; if Bellemans would not have been part of the new describers, then this would raise a major conflict, but since he is, there is no way but acknowledging the new describers and new date for the species… but that might be challenged in the future (Jean Huber,  finalized July 2003, updated December 2009, October 2012).



Q(18): I maintain and breed a strain of Procatopus nototaenia coming from the Fifinda river that I caught with Patrice Lambert in February 2001. I need more information about the geographic distribution of  nototaenia. The information that I have now are :
- Proc. nototaenia lives in the south of Cameroun from the south of the Lokundjé to the Kienke, the Lobé river and the Ntem. (Killi Data on line).
My questions are : - Is nototaenia present in the Lokundjé ? If I have found it in the Fifinda, i.e. north of the Lokundje, it could mean that nototaenia lives in the Lokundjé, then that the geographic distribution of similis begins in the north of the Lokundjé. I hope you could give me answers or any information about the Procatopus nototaenia distribution (Olivier Buisson, March 2003)

A: You are pointing at a problem in Procatopus taxonomy. In fact you are saying, where ends the distribution of P. nototaenia and where starts P. similis. This question I cannot answer you. First, because I think (but think only) that the two are one and the same species and that differences exist only at the population level. Second, because I never made an in-depth study of the problem and never was in that area of Africa to collect. If we want to distinguish Proc. nototaenia from Proc. similis, one of the most striking features is the extended fin rays of the male of nototaenia. But also populations in the eastern Niger delta, identified as similis, do have those extensions. Another mark of identification could be the straight dorsal profile that nototaenia usually shows, versus the more curved profile of similis. The reddish stripe on the back of nototaenia, sometimes mentioned, is also found in some of the populations of similis. Therefore, I advise you to look at all possible photographs of males from that area and distinguish them on the basis of extended Anal fin rays. This may give the best (up to now) clue to establish a border between the two species (Ruud Wildekamp,  finalized March 2003).



Q(17): What is your opinion on the molecular clock and dispersalism vs. vicariance? In my book on Cichlids of Madagascar that has just been published, we suggest that because there is no primary freshwater fish available in the large island and because Killifish and Cichlids are secondary freshwater fishes, then they should have reached Madagascar through the sea, at a much later period than the continental drift of Madagascar from Gondwana, therefore through dispersal. In details, firstly, if the presence of Cichlids and Killies in Africa on the one hand, and Madagascar and India on the other hand, was the consequence of tectonic movements, then Cypriniformes (Barbs) should be found in Madagascar, for example. However they seem to have never been there. I thought, on the contrary, that the presence of Etropline Cichlids (Etroplus et Paretroplus) in India and Madagascar (and, probably too, the presence of Aplocheilus and Pachypanchax), and in these 2 regions only, could be, and even ought to be the result of a tectonic drift between India and Madagascar that according to some geologists occurred about 100 million year ago, i.e. after the split of Gondwana and after the break of the block "India-Madagascar" from present Africa. In a first step, I have then viewed the issue without considering the molecular clock. However, Miguel Vences and colleagues have shown that the divergence between Etroplus and Paretroplus could be old of about 20 million years, whereas the separation between India and Madagascar could be old of between 90 and 60 (minimum) million of years … what are your comments ? (Patrick de Rham, January 2003)

A: The question of rates of molecular evolution and the molecular clock is pretty complex and controversial. First of all, one needs to use genes or DNA sequences that evolve sufficiently slowly so as not to be too "noisy" (the noise obscures potential signal). The assumption of the molecular clock is that there is a linear relationship between the amount of molecular divergence and the amount of time. One has to test that, if that is not the case, then one needs to apply correction in form of a function to the molecular data. Another assumption is that the molecules evolve at about the same rate in all lineages studies. This often is not the case, it is much harder to deal with. One often has to assume a "local" molecular clock rather than a "global" molecular clock. If one is able to overcome these issues, one can then "calibrate" the molecular clock. This is done using fossils or some kind of a vicariant event (one fixes a point of certain amount of molecular divergence to correspond to a certain time divergence and extrapolates the rest) or one uses "standard" molecular clock (usually something like 1x10 -8 for mtDNA) without any specific knowledge for the group of organisms studied. This is the basic approach, but there are other non-parametric or semi-parametric approaches that are able to deal with molecular rate heterogeneity in the data. (I have used these approaches on the Aphanius data and got the same results as reported in the paper).
There are lots of papers that deal with very old divergences, but only a relatively few with fish. I am including a pretty good paper that deals with fish outside Killifish (Molecular Phylogeny of Osteoglossoids: A New Model for Gondwanian; Origin and Plate Tectonic Transportation of the Asian Arowana, by Yoshinori Kumazawa and Mutsumi Nishida. Mol. Biol. Evol. 17(12): 1869-1878. 2000; E-mail: I know of the cichlid paper. I have read it and thought about it, and in general it does make sense to me. There are a couple of issues that I think will need to be addressed in cichlids, but otherwise I think it is pretty solid work. Still no matter what happened with cichlids, it may have no bearing on killifish. It is entirely possible that cichlids have dispersed, and that killifish have drifted on continents. What one group does has no influence on what the other group does. (Tomas Hrbek, finalized January 2003) The question of secondary or primary freshwater fish seems unimportant to the molecular clock issue. Among Madagascan Killifish, there are 2 groups -Pachypanchax and "Pantanodon"- which seem to have very different destiny. For the former, MtDNA has shown its link with Aplocheilus and with the plate tectonic events (see Murphy, W.J. & G.E. Collier. 1997. A Molecular Phylogeny for Aplocheiloid Fishes (Atherinomorpha, Cyprinodontiformes): The role of Vicariance and the Origins of Annualism. Mol. Biol. Evol., 14 (8): 790-799, 6 figs., 2 Tabs.), but the authors do not use a molecular clock. For the latter, if it is finally attached to Killifish, the pattern of vicariance is ambiguous and reversed because the other (true) component of Pantanodon (P. stuhlmanni from coastal Tanzania) seems to be derived (i.e., not primitive) (see Huber, J.H. 1998d. A Comparison of Old World and New World Tropical Cyprinodonts. A parallel Outlook of similar and distinctive Characteristics regarding Distribution, Evolution, Ecology, Behavior, Morphomeristics, Genetics and Color Pattern. Soc. fr. Ichtyologie: 109 pp., 17 figs.) (Jean Huber, finalized February 2003)



Q(16): Is the identity as Plataplochilus mimus correct or legitimate for the Gabunese somewhat deep-bodied population of Tchibanga? I believe that the population from Kinguélé, sympatric with A. striatum and Ep. sexfasciatus, cannot be mimus and is closer, by morphology, to ngaensis. What do you think of this and also of Roman's identifications in Rio Muni of Plataplochilus sp. with a similar morphology to the Tchibanga population? (Eduard Pürzl, January 2003)

A:  The fish from Tchibanga (SW Gabon, about 2.760S 11.080E) cannot be mimus that had been described from north of the Ogooue, at PK 28 Libreville to Kango, N.W. Gabon (0.367N 9.700E), completely in the coastal plain. You also quote another Tchibanga in the coastal plain, more northerly (1.13S 9.55E), but this is still south of the Ogooué river and in that case, according to Lambert (1967) it should be identifiable to chalcopyrus, another putative junior synonym of ngaensis. You are right to mention that the Tchibanga population is somewhat deeper than mimus. If it was collected in the southern Tchibanga, it should be identified as Plataplochilus aff. cabindae (type locality: Lucola river, near Cabinda, Angola, 5.533S 12.250E), as proposed by Lambert in 1967. 
Plataplochilus mimus is very similar (hence the name !) to ngaensis (and it is a supposed junior synonym) but German aquarists have collected a few somewhat different populations near Kougouleu and Song at the foothills of Cristal Mountains with a red chevron near the caudal peduncle in dominant male of wild fish (not in F1 generation), and they tend to identify it Platap. aff. mimus. Your photo of a fish from Kinguélé also shows that red chevron. Kinguélé is situated in N.W. Gabon, near the foothills of the Cristal Mountains, like the type locality of ngaensis (Atogondema, Nduya river, N.W. Gabon, 0.850N 10.250E). It would be legitimate to identify the foothills populations as ngaensis(and not mimus) and to use mimus for populations of the coastal plain near Libreville, but that is not the case and I did not want to introduce confusion in Killi-Data, since nobody knows what really is ngaensis: therefore I kept conservatively the used names and allocated your Kinguélé photos to mimus and the Libreville photos to ngaensis. On the other hand, Roman's identifications are probably erroneous because he identified his fishes from criteria of morphometrics and patterns used for riverine standard fishes (not killifish) and it is known that the morphology of Plataplochilus (and related genera like Procatopus) is extremely variable at the population level. The male color pattern and the adult body depth and size seem more appropriate as diagnostic for the time being. But I must admit that our knowledge of Plataplochilus is extremely poor. As you know in my 1981 paper I recognized only 4 valid species, ngaensis, cabindae, miltotaenia and terveri, but Wildekamp has demonstrated rightly later that loemensis is valid too and distinct from cabindae. All the rest, we don't know and notably we are ignorant if the color variation that is observed in Plataplochilus and that led to the description of pulcher, chalcopyrus, mimus corresponds to specific diagnosis (like in Hypsopanchax) or polymorphism (like in Procatopus). And our knowledge is scarce concerning the live pattern of Rio Muni's populations, not to mention that the color patterns of topotypical cabindae and ngaensis are unknown ! Only DNA techniques may bring a beginning of an answer to that difficult problem. (Jean Huber, finalized February 2003)



Q(15): What is your opinion on Fp. gularis and Fp.deltaensis? Should they be separate sp. and if so how would you separate them? (Tim Addis, January 2003)

A: Well, this is an uneasy question, not because this is difficult to answer, but because data are insufficient for a perfect answer. Fp. gularis and deltaensis are today well known, well described with precise type localities, distant by about 80 km (Agberi, southern Nigeria, 5.233N 6.383E and ca. 200 yards south of road Sapele-Benin to Warri in western Nigeria delta, Midwestern State, Nigeria, 5.820N 5.770E, respectively), of course. They can be separated by the male color pattern: deltaensis is a yellow phase with a red median line on sides and gularis is a blue phase with very few red spots scattered on sides (Boulenger's description), to many spots and even broken red blotches more or less along mid sides. Whereas deltaensis is only known from a few localities, gularis is known from many locations in western Nigeria and Bénin. No study has been performed since Scheel and no collecting trip has recorded samples since many years, except one. That important collection, reported by Wildekamp (1995), records deltaensis from nearby the type locality of gularis. In addition, crossings between the two are fertile beyond the third generation in aquarium. 
These observations may legitimately induce the conclusion that the 2 names correspond to the same species and that deltaensis is a color variation and a junior synonym of gularis, as Wildekamp has proposed. A respectable move. 
However, in Killi-Data, conservatism prevails in order to minimize the risk of inappropriate changes and deltaensis is provisionally and cautiously kept valid for 4 reasons: first, gularis from its type locality and according to Boulenger's description (very few spots in male) is unknown live; second, it is not known if the few locations of deltaensis corresponds to a vicariant distribution or to a sympatric pocket distribution within the wider range of gularis; it is not known, either, if the probable sympatric occurrence of the 2 around Agberi is the rule or an accident within the Niger delta which is known as a very complex region for Killifish, maybe because of the past occurrence of an epicontinental sea there; third, it is not known if the genetics of the type population of Agberi are identical to those of the type locality of deltaensis and detailed molecular studies have not been performed over the entire range (preliminary reports show that both species are DNA-distant); fourth, the case deltaensis/gularis should be viewed with a wider point of view, i.e. including fallax Ahl and fallax sensu Seegers (with a publication in preparation from BMNH samples). All in all, very little is known on these fishes and on that region of Nigeria, probably one of the most important for Killifish in terms of speciation. Political instability, only, explains that poor situation. Let's be patient. (Jean Huber, finalized January 2003).



Q(14):  Is Megalebias robustus a valid species, as per reported in Seegers's  Aqualog and why? (anonymous, December 2002)

A:  This is a puzzling problem, since in Aqualog no evidence is given, but the mention that the name is valid with a photo of a male. I have sent a mail to Lothar Seegers to ask for an explanation of his revalidation of Megalebias robustus. But no reply yet, sorry. The fish that is shown in Aqualog (III) as robustus is identical to what Ruud Wildekamp calls the deep bodied elongatus in his book (World of Killies, volume II). It is an aquarium strain imported in Germany, without a precise origin. The first to report on it was the renown German aquarist Walter Foersch as elongatus.
Reference (with a color photo): Foersch, W. 1978. Wie laicht der gestreckte Fächerfisch? Beobachtungen bei der Pflege und Zucht von Cynolebias elongatus. Aquar. Mag., 12 (2): 86-93, figs. The question is: what is the real robustus according to the description ?
Cynolebias robustus has been described by Günther (1883) from San Antonio, a locality probably about 150 km to the East of that of elongatus.
Up to now, all researchers report that it is a synonym of elongatus from reading the description (morphology, type locality).
But only a study of the types that are stored in the British Museum in London can solve the question together with a recollection of Cynolebias robustus at type locality to compare it with types of elongatus
Late 2002, a collecting trip to the type locality of robustus has been organized by Argentinean aquarists and they found 3 species, the standard Megal. elongatus, Austrol. bellottii and a fish that is similar to Austrol. nonoiuliensis, identical to robustus, and that has been redescribed. In conclusion, Foersch's fish is related to elongatus but its origin is unknown. Then, robustus is a valid Austrolebias species, closely related (maybe identical) to nonoiuliensis. (Jean Huber, finalized, December 2002, updated January 2003).



Q(13): Why is there a difference in the name and date of describer for Austrolebias carvalhoi, between Killi-Data online and Costa's publications? (Pablo Calvino, November 2002)

A: The difference between the original describer of Austrolebias carvalhoi in Killi-Data online (Costa, 1995) and publications by Costa (Myers, 1947) lies in the evaluation of the minimum description that is required to declare a name as available (i.e. as "officially described", according to the ICZN rules, published by the Commission of Nomenclature).
According to Costa and other scientists, a few words of description are sufficient to declare a name as available.
According to the author and other scientists, a diagnosis, i.e. a description that gives separating characters from related (or not) species is necessary to declare a name as available. The ICZN requires that a diagnosis is given (from 1930) to declare a name as available. The meaning of the word "diagnosis" is then not understood identically by one researcher or another.
This is not, however, a big issue for the carvalhoi case, since the validity of the name has not been discussed.
Only the author's name is discussed:
Either, Myers (1947) as per Costa, or Costa (1995) as per Huber (2000) in the book Killi-Data.
The original publication by Myers contains a few words and the question is whether that constitutes a diagnosis.
The decision of Killi-Data is more to show that, for a paper published after 1930, the single full diagnosis that is available is Costa's description in 1995.
Update (2006): Lazara (1979) reported on a pers. comm. by Myers to him on carvalhoi : «… one of my species, but never described, so no one can identify it!» and concluded that «Myers did name the fish, but without a proper published description, a name has no status» ; Myers's remark was probably the result of the difficulty of separating carvalhoi from the variable, then known adloffi in terms of colour pattern, by Myers. However, to be complete, it must be mentioned 2 other publications. First, Vaz-Ferreira & Sierra (1971) attempted a redescription of the taxon from material collected in southern Uruguay (1200 km away from type locality), but it was a misidentification and that material was later described as cinereus ; second, Lazara (1981) studied the case again and came to the opposite conclusion that Myers's name was available, despite his 1979 analysis : his proposal was based on the descriptive sentence («from highlands of upper Rio Iguassu near Porto Uniao, a small species of the deep-bodied type, dull in color, the male with numerous vertical bands wider than those of adloffi»), on the availability of types in Rio de Janeiro Museum and Stanford University and on the fact that the then available alternative, an article using the name carvalhoi by Vaz-Ferreira & Sierra (1971) was not ICZN available. Lazara's proposal should not be overlooked, because Myers's descriptive sentence includes a diagnostic element vs. adloffi. Against Lazara's proposal are the following arguments : (1) the diagnostic character vs. adloffi is of little value, (2) the putative types are not mentioned at all in Myers's article, (3) Myers himself considers his article as not a valid description, (4) Costa (1998) wrote the first article who presents a formal redescription of the taxon most probably based on the original material from Porto Uniao in Rio de Janeiro Museum and in that article he writes that the «first diagnosis dates of his 1995 publication», (5) Myers's material was collected near Porto Uniao (without precision) and it cannot be discarded that in its vicinity another cryptic species is distributed since it is frequently the case for the genus (the region is entirely virgin of collections except Costa's single locality). In total, a very difficult case : nomenclaturally, carvalhoi Myers may be seen as either unavailable, including according to Myers himself and the first publication that meets the availability requirements would be by Costa (1995), using «types» in MNRJ ; but to the contrary, it may be seen strictly as available when the single character is accepted as diagnostic. Indeed, Myers's disclaimer is indirect and 32 years subsequent to the original publication. Systematically, the situation has changed significantly when Costa (1998) was able to collect a fish in the same region as Myers and redescribed the taxon in details : it appeared that the probability that Costa's fish and Myers's fish are identical is significant and that the taxon is a valid species is high. The final word on this issue will surely be derived from a new study based on an in-depth collection of material in the region : if a single Austrolebias species is living in the region, then there will be no doubt on Myers's so called «diagnostic» character and the balance will move to availability (diagnosis vs. unwillingness of authorship) ; but, if there are two similarly patterned species or if the congeners of the region are DNA variable or polychromatic, then Myers's so called diagnostic character will be valueless and only the subsequent description by Costa, 1995 may be considered as available. Until then, Huber (2006) proposed to follow the consensus of authors (Lazara, 1981; Wildekamp, 1995; Costa, 1995) i.e. accept carvalhoi Myers, 1947 as presumably available and a valid species. To keep Costa as the actual describer and respect Myers's position of unavailability is equally reasonable, but it does not take into account the poor and debatable, but formal, diagnosis of that taxon, even if that diagnosis is no better than the absence of diagnosis in other taxa erected by Myers in the same period, like for antenori, regani, aureoguttatus (Jean Huber, finalized, December 2002, updated July 2006).



Q(12): With a friend of mine I plan to go to Costa Rica for holiday to visit the country and, at the same time, to collect Killifish. We need authorizations to take the fish back to France. As an experienced collector in that country, what would you advise us to obtain the various legal authorizations? Everywhere in the past (Gabon, Vietnam) I have just collected the fish and took them home in my luggage saying nothing to the country customs. Can we do that and what do we risk in South America? 
Notably, we fly back to France after a transit in the US and it may be an additional problem, what do you suggest? Finally where would it be best to go collecting? (Hervé Gonin, August, 2002)

A: It is best to be completely legal. If you collect the fish illegally and export them illegally, you risk having them confiscated at the airport when you leave Costa Rica. I have been asked for copies of my paperwork, and with security as it is now you won't be able to hide your fishes. You can try to do things illegally, but it is much better to be entirely correct.
If you collect illegally and are caught, do not claim to be a scientist or affiliated with a scientist or collecting for scientific purposes. They're much harder on scientists who don't follow the rules.
I don't know for sure about transit proceedings, but I expect you'll change planes in the U.S. I know it is ridiculous, but transit passengers have to clear US customs. Declare the fish on your US customs declaration. It is better to declare them than not declare them and have the customs agent find them. If you declare them, customs and the Fish & Wildlife Service will probably take no interest in them at all (except for protected species). But, do NOT try to smuggle plants, birds, mammals, or dendrobatid frogs through the US! 
Best advice: make your collections legally. Directions to the gate-keeper for permits in Costa Rica can be found at:  (also for other South American countries). There, you can learn how to initiate the process of getting permits to collect fishes in and export them from Costa Rica. I urge you to obtain permits. Visit this web site, get the forms, and write write write. You won't need a phyto-sanitary certificate, but you will need an animal health certificate. I think that can be arranged with the export permit. It would help if you had a scientist in France vouch for you, the ideal would be to have someone at M.N.H.N. design a project that needs fresh, better yet, live, material from Costa Rica. Also you'll need a letter of support from someone in Costa Rica. I think you'll find CR much more developed than Vietnam and even, in places, a little francophone.
About where to go? I have little idea. Your collection of data from Killi-Data is a good starting point, but since I have not been to CR since 1999 I really can't make suggestions. Everything there is changing rapidly. Best that you just go and look. I think late October is not the best period. That's approximately the end of the rainy season, and field conditions might not be good. Very wet, fish hard to find, and so on. In general, it seems best to visit CR early in the dry season. I also think it is a little crazy to rent a 4x4 in CR. Everything of interest to aquarists, and nearly everything of interest to scientists, can be found within a short distance of roads that are passable with a small "Sedan" car. In 1999, I rented a small Hyundai Sedan because of lack of money (naturally one wants a spacious vehicle with good ground clearance "just because"). I drove on unpaved roads for much of the trip, only once had to drive very thoughtfully, and I hit all but two of my Rivulus targets. The fishes I couldn't find were not in the places where I'd got them before or in similar situations close by. I didn't fail to get them because I couldn't travel on the roads. Where there were roads at all, my car could go. The Hyundai was not a handicap at all. About where not to bother going for Rivulus. Sierpe drainage. All brackish, no Rivulus. Pacific slope of Guanacaste province. All dry, no Rivulus, except in the headwaters of the Rio Corobici (?) at Tilaran. Pacific slope north of the mouth of the Rio Terraba (this region includes Guanacaste). No Rivulus. About Oxyzygonectes, I haven't seen the fish in CR since 1979. I have no idea where to look for it in CR. I did look for it in mangroves along the Nicoya Peninsula (just to see, no hope of catching adults single-handed) in 1999. I didn't see any at all. (Dan Fromm, finalized September 2002).



Q(11): Would you be able to advise me as to the status of Epiplatys sp. Lake Fwa ? The Americans consider that the name sp. Lake Fwa should stand until a decision is reached on whether it should be named a new sp. Ed Pürzl considers the only difference between Lake Fwa & Ep. chevalieri nigricans is that Lake Fwa is more slender. Wildekamp (in World of Killies) considered Ep. chev. nigricans endemic in and near Lake Fwa. Seegers shows 3 photos in Aqualog: 2 are slender, 1 is not. (Tim Addis, July, 2002)

A: This question is a good mirror of our poor knowledge of the Killifish fauna of the Congo basin, and not only for the genus Epiplatys. After having collected there back in the late seventies and studied the material, I have been convinced that nigricans is a junior synonym of chevalieri. Up to now no new published evidence has been put forward to change my view. Ruud Wildekamp has proposed that western populations (i.e. west of Congo river) are chevalieri while eastern ones are nigricans, on the basis of color pattern differentiation of some populations. But that was tentative to maintain a conservative position and he did not used the same model for the multifasciatus group (boulengeri / multifasciatus). There are differences between populations of chevalieri/nigricans from the Congo cuvette, but we simply don't know if these differences are stable, if they correspond to strong genetic divergence, if they are linked to geographical criteria (there is absolutely no barrier, but the concept of refugium may apply, like for lake Tumba). All we know are a few populations from the Kisangani area, from the Brazza area, from the Likouala area and from isolated spots like the one from lake Fwa, thousands of kilometers apart. And neither molecular studies, nor detailed morphological measurements have been performed on these populations. It is extremely limited. And some people may add it is totally insufficient. Indeed in conclusion, I believe the "American" conclusion to name the fish Epiplatys sp. -Lake Fwa is reasonable and I fully agree on its pragmatism. (Jean Huber, finalized, July 2002)



Q(10): There are some photographs of a Rivulus species we found around the village "Mabura" some years ago which have some resemblance with Riv. dibaphus and geayi. The pictures show a barred pattern in the male Caudal fin. In Suriname, we found a similar species in the drainage system of the Marrowijne border-river to French Guiana (Palemeu). Where do you think you can place this species? (Frans Vermeulen, February, 2002)

A: The Guyana and Suriname populations of Rivulus showing similar color pattern in male with geayi mark the limits of the taxonomy in Killifishes. Indeed it is known since Scheel that Killifishes are extremely variable in genotypes. A new name could be proposed for these aberrant populations, but the added value would be limited and even zero, because then there will not be any limit in describing new names (e.g., with allopatric, genetically distinct, populations of Aphyosemion calliurum that cannot be separated by character of coloration). And cherry on the cake, there are also similar populations in western French Guyane, also isolated within the range of the very similar agilae. Maybe in the future, notably after DNA experiments, the understanding of this situation will be improved. A reasonable explanation has been forwarded with the past climates hiccups of the dry maximums and the refugium concept (Huber, 1998), but it needs further study. Until then, it is more realistic to name these aberrant populations Riv. aff. geayi. (Jean Huber, finalized, May 2002)



Q(9): Wim Suijker went to Guyana 2 years ago and had the goal to reach the terra typica (type locality) of Rivulus holmiae, described in 1909. We did before some attempts to reach that goal but were not able to reach the spot that times due to several problems. He did succeed and now it comes. The name of "Holmia", the place where Eigenmann found his Riv. holmiae, is renamed in an other name "Sinnapouw", but it is sure it is the same location. Wim was not able to find other fishes than the species we know as Riv. waimacui. There were no other discoveries. It cannot be true that the species has disappeared in the 90 years in between, certainly if you remark that there is no human involvement in that region. I asked Wim if he had looked very thoroughly and in every possible location. He did. Wim and I did put a lot of our time and money in the search of this species and we were not successful all that time. What do you think about this. Is it possible a scientist like Eigenmann could have made a mistake like that ? More to the east in Surinam are species which are very close to the description of Eigenmann's Rivulus holmiae. Rivulus igneus is more like that Riv. holmiae. (Frans Vermeulen, February, 2002)

A: With Riv. holmiae, we have a very good description for the beginning of the twentieth century, plus numerous type specimens of both sexes in good conditions. By morphology, holmiae is close to igneus and to immaculatus. It differs from igneus by the frontal scalation, the male color pattern and minor morphological characters. It does not differ from immaculatus except by having a supracaudal ocellus in female (but this is a variable character). In addition, holmiae and immaculatus are living in the plateau. I do not mean that both are synonyms, but our thinking may be biased by the historical collections in the lowlands of Guyana and Suriname that were identified as holmiae and that may be closer to hartii (or to igneus). Riv. waimacui is very different from the types of holmiae (also as a living fish, as you have demonstrated). No precise argumentation can be forwarded whether Eigenmann made a mistake about the type locality or not. He was a young researcher then and stayed in the area for a long time. Eigenmann is acknowledged as a serious researcher and an error in labelling is unlikely, and no researcher would follow a person who would believe of an error on the basis of being not able to recollect a fish. In addition, we have examples (e.g. in Diapteron) where a fish could not be found at the same locality a few years after description, but an in-depth survey by natives proved the contrary soon after. Therefore, the reasonable and operational conclusion is to keep considering holmiae as valid and distinctive from igneus and immaculatus, and also from the very different waimacui, until fishes are collected not far from its type locality and different from waimacui. Then to collect intensively in the type area, in the region in-between with immaculatus and to compare the material with the lowlands in Guyana. I know that after your considerable efforts this answer may be frustrating but just imagine the situation with Riv. micropus, with a type locality as uncertain as Rio Negro (2000 km !)… and in that specific case (all books have been destroyed in the Vienna Museum during the 1945 war) Costa had no choice to designate a new type locality (identical to its junior synonym, compressus). (Jean Huber, finalized, May 2002, November 2011).



Q(8):  You know my dedication to Roloffia: I have always been sorry by the quarrels surrounding this name. Now it seems to be also the case with Aphyosemion bualanum / elberti and Lazara does not seem to follow your preference of bualanum in the special JAKA issue of 2001. Could you clarify? (Royal Ingersoll, April, 2002)

A: I have always been sorry with the quarrels surrounding any name, too. First comment : times change and quarrels are rare today compared to during the 1970'ies because many cases have been clarified (not all) and because researchers, amateurs or professionals, are judged on published evidence and not on emotional opinions without solid arguments. My paper on bualanum/elberti and on escherichi/microphtalmum in Aphyosemion is absolutely in line with this philosophy : it is based on the study of the types. But since Seegers has also studied the same types, I am suggesting that new evidence is searched with new techniques (e.g., DNA) to solve these thorny problems, and even if I believe my observations are correct, I ultimately propose that readers use as valid whichever taxon they prefer. In Killi-Data, my point of view is followed because, with computers, you have to make a choice, because it is the last published evidence and because I follow conservatism in line with ICZN recommendations. But Seegers's point of view is duly mentioned. Maybe it is also useful to add, for emotional people, that I consider Lothar Seegers as a very good researcher and that I followed his results most of the time. Regarding Lazara's position, there is a confusion. Although the Kathetys special issue of JAKA has been published in 2001 (reprinted in BKA Killi News of July 2002), i.e. long after my bualanum publication, it is a simple copy of what appeared in KMI for which the publication dead line has been December 1999 and this may explain why my publication (also in JAKA!) is not quoted. This does not mean that Lazara agrees with my observations. Time will tell! (Jean Huber, finalized, April 2002; updated July 2002). Note : this answer is dedicated to Royal's memory who died a few weeks after he asked this consensus question, very well in line with his character).



Q(7): What do you think of the codes and should we use them in place of population names? (Michel Leclerc, November, 2001)

A: My approach to this sensitive subject is pragmatic : if you can mention both, the population name and the code of collection, this is best. But I would suggest not to use the code alone. Codes, although interesting, have major drawbacks. First, printing errors are extremely easy (e.g., change of TAN95/7 for TAN95/1 or 4) and consequences may be disastrous: if the same species has been collected at the two locations then crossing are unavoidable and sterile strains may be the outcome, and, if the species are different a lot of confusion will show up. Population names may also be altered frequently, but errors are easy to reconcile with the truth: for an experienced breeder, the "Estrias" population of australe is to be corrected into the right "Cap Esterias" at first glance. Second, a code does not say anything, to the contrary of the population name that gives a clue of the place of collection, e.g. Makokou in Gabon, even if in exceptional cases the village may bear the same name in several countries! Proponents of codes alone argue that it is not always possible to give a population name because no village is in vicinity. This can be easily solved by giving a name like West Makokou or PK 20 Makokou or 20km West Makokou by the collector himself. Proponents of codes alone also argue that codes allow to avoid the mixing of populations of the same locality, collected on different years. Well, let's ask ourselves what are the objectives of aquarium population names: unambiguous identification obviously and also development of a long term aquarium strain: if the two fish are from exactly the same place, then mixing them will have new blood benefits. I believe strongly that aquarists must only remember, as a required minimum, the species name and the population name (and not the generic name that changes too often). And today the population name is becoming extremely important, not only to avoid mixing intersterile populations (as it was pinpointed in the past), but also because species names do change not rarely: e.g. Cynolebias albipunctatus from Itacarimbi has been renamed Cyn. perforatus, or Simpsonichthys flavicaudatus from around Manga, Itacarambi has been renamed Simp. flagellatus… with codes only, it would be a nightmare ! Proponents of codes finally argue that with commercial importations, there is no other way than using codes. Yes, absolutely! But let's move and keep asking importers to obtain the information, or else, give a fantasy name, e.g. Schleser's strain for a well known strain of a South American annual from the trade import. Besides, this idea can also be applied for populations collected in the middle of nowhere (e.g. 200 km away from any named place), with the decision left to the collector. That's what Christian Cauvet actually did for Scr. cauveti - Barrage Loire Atlantique: anecdotally, the name is not that of a village, but that of the notice board near the creek of collection. The French name indeed means "the Dam that was financed by the administrative district of Loire Atlantic, as a cooperationproject with Guinea"! (Jean Huber, finalized, December, 2001; updated July 2003).



Q(6): How can be measured the geographical coordinates of a collecting spot without GPS, for example my spots of southeastern Brazil of last year?
What do we we use to measure the eggs: millimeter-scaled paper (fast, but not very accurate) or better with a micrometer? About the incubation time of the eggs of annuals, do you expect data with standard temperature and wetness of peat (e.g., 25°C, tobacco dryness as usual) or is it enough to give a "medium time"? (Stefano Valdesalici, April, 2001)

A: If you do not have a GPS device or if you are not any more in the field, you only can compute geographical coordinates of a collecting spot approximately, from your map and a decimeter (you have to measure the distance of one degree in mm and of each of your spots from the basal line of the south and the west and compute them in hundreds of degree of latitude and longitude by dividing the measured distance of your spot with that of 1 degree). For example, your map of the Sao Paulo area in Brazil shows squares of 111 mm for one degree of latitude and longitude (sometimes, these are not squares, but rectangles corresponding to more than 1 degree and then the computation must be adjusted), and your spot is at 61 mm of 22° south line and at 14 mm of 46° west line: geographical coordinates are easily obtained by dividing 61 by 111 (0.55) and 14 by 111 (0.13), i.e. 22.55 S and 46.13 W.
Re diameter of eggs, a cheap plastic micrometer is ideal (about 5 Euros or 4 US Dollar), but you can also use millimeter-scaled paper (0.1 mm accuracy is fine because the egg diameter also varies according to external factors, the first being the population itself!).
For the rest (incubation time, time of sexation, maturity, adult stage, normal and maximum life expectancy), a medium estimate of the optimum is enough in normal conditions because it varies according to many factors (an approximation is better than nothing). Obviously, you are not going to give data corresponding to abnormal conditions, e.g. 30°C of water temperatures for Austrolebias species or Orestias species, 18°C for Simpsonichthys species (etc.) (Jean Huber, finalized, April 2001).



Q(5): What is the best way to abbreviate generic names? (Roger Brousseau, February 2001)

A: Generic name abbreviations are standardized for scientific publications: first is stated the name in full (e.g., Rivulus), then for any further mentions is stated the first letter (e.g., R.), followed by the concerned species name. However, this may become awkward when several genera are mentioned that start with the same letter (e.g., Renova). The main problem arises when lists of species in several genera are built, then common sense should prevail: old, frequently used genera may be abbreviated strongly with a single letter (e.g., A. for Aphyosemion or F. for Fundulus), but other genera with the same starting letter will be abbreviated with two or three or four letters (but not more, because it is no more an abbreviation and it looses its interest) and it can be used to differentiate two similar names (e.g., Ep. for Epiplatys, as commonly used, and, Epis. for Episemion, subsequently described). A spiny problem occurs when 4 or more starting letters are identical (e.g., Fundulus, Fundulosoma, Fundulopanchax), then usage and pragmatism should prevail (respectively, F., Fund., and inevitably Fp., with the first and the last genera being the most speciose). Describers should avoid to create taxonomic names with such strong similarities, but this is the trend today when several generic names are stemmed from the same root, indicating their relationship, as Austrolebias, Megalebias, Stenolebias… (although this relationship may prove later to be erroneous, like Pachypanchax and Micropanchax). This difficulty has been exacerbated recently with the description of Austrolebias whereas Austrofundulus was already known and its standard abbreviation has had to be changed in Killi-Data: Austrof. for Austrofundulus and Austrol. for Austrolebias and (update) even more with the description of Notholebias by Costa in 2008 which is easily (and regrettably) confused with Nothobranchius (Jean Huber, finalized, March 2001, February 2009).



Q(4): What is the status of the Diapteron population PEG98/5 which shows bars on sides, an orange Anal fin and a faint orange band on lower Caudal fin of male, i.e. intermediate between abacinum and seegersi? (Jaroslav Kadlec, July 2000)

A: There are currently 5 species in the subgenus Diapteron (molecular studies have shown that Diapteron is close to Chromaphyosemion and Kathetys, two subgenera of Aphyosemion, and ecobiogeography together with a PAUP analysis of external characters have shown the same, adding Episemion to that phylogenetic group). In this latter publication (Huber, 1998), it is suggested that Diapteron species are very young, possibly less than a few thousand years. The species are, though, intersterile in crossings, except if male unpaired fins are cut to confuse the selecting female. Genetically, however, the picture is much more complex with reported intersterility within one taxon (e.g., cyanostictum) or interfertility of first generation for some distinctly colored populations of distinct taxa. The best is then to consider these 5 "species" on the basis of the single (rather stable) diagnostic characteristic, i.e. the male color pattern (and similarly for many superspecies of the genus Aphyosemion): (1, 2) the dominant and widespread blue (cyanostictum) and yellow (georgiae) phenotypes with blue spots on sides and fins on red background (note that some populations of georgiae show orange instead of yellow, completely on Anal fin and lower Caudal fin); (3,4) the locally confined to the dead-end eastern distribution, blue (abacinum) and yellow (seegersi) phenotypes with blue bars on sides and fins on red background; (5) the locally scattered double banded at Caudal fin yellow (fulgens) with red spots on red background (derived from georgiae). The recently collected population "PEG98/5" fits with seegersi in a large sense (in the same way than the orange georgiae fits in georgiae): it shows yellow orange markings on lower unpaired fins like seegersi, though fainter. Therefore, to that complex picture, the only current practical answer is to name population "PEG98/5" from northeastern Gabon: Aphyosemion (Diapteron) aff. seegersi, where aff. is an abbreviation for affinis meaning similar to. The population is distinctive from abacinum by color pattern of the distal part of fins in male. Further studies in the field and by crossings with abacinum and the typical seegersi (when it will be ultimately brought back alive from Congo) may give a better answer, but nothing is less certain (Jean Huber, finalized, September 2000).



Q(3) How do you explain the differences between the books Killi-Data 2000 and the preliminary list of the Killifish Master Index, published in the Journal of A.K.A. (June 2000) by Baker-Carr and Lazara? (Anonymous, August 2000)

A: Globally, Killi-Data 2000 and KMI list 2000 are not that different, since 90% to 95% of the detailed listings of names are identical, which is indeed a mark of similar views. Differences may be explained by (1) interpretations of publications (when authors do not state precisely the consequence of their findings in terms of taxonomy), whether it must be splitting or lumping, (2) information regarding soon to be published works, available to Killi-Data and not to KMI, or vice versa, (3) decisions how to handle debated (discussed or controversial) cases, one way or the other, (4) last update limit (November 1999 for Killi-Data 2000 vs. February 2000 for KMI list) with a few new names, described in-between. But these 5-10% differences will not be explained, case by case, until the actual publication of Lazara's new KMI that is scheduled soon (published in December 2001). With Killi-Data online, which is constantly updated and which details all taxonomic views per taxon, all differences will disappeared or at least will be clarified (Jean Huber, finalized, August 2000, updated June 2002).



Q(2): How do you find the time to do all that stuff and to process Killi-Data with so few typos? (Jorge San Juan, March 2000)

A: My job includes a lot of travelling, since many years and I use a laptop computer (Toshiba, Dell) and a pen scan (weekly train return trip to Bruxelles or flight trips to the UK, Germany, Swiss, Italy, Spain, etc) : to cumulate from a few minutes to an hour or so, each time, makes a tremendous amount of work in total. Processing Killi-Data properly requires method, only: a critical path has been designed for any new or modified data and none is added without having terminated the full process of each. Also the analysis of any new information is unique for Killi-Data, different for any other list of valid species for Killifish. The analysis is based only on published evidence (Is the information scientifically demonstrated ? Is there a comparative diagnosis ? Does the publishing magazine edit a board of scientific advisor or anonymous reviewers of articles ?) The analysis is not opinion based and obviously not based on my opinion. If the analysis ends up with the fulfilment of all basic requirements, then the publication is fully acknowledged in Killi-Data online, even if its results are contrary to other authors' previous publications (including those by myself, of course). All lists of Killi-Data are based on these strict principles and must not be considered as my own lists. In fact, it is just the contrary, my preferred list of valid names, for example, would be quite different from the list available in Killi-Data. By doing this, I aim to serve best the scientific community and to promote best the progress of knowledge (Jean Huber, finalized, July, 2000 ; updated January 2004).



Q(1): What do you consider should be the basic ideal paper library for a Killi-hobbyist? How to get these undisputable books? (Jean-Paul Cicéron, February, 2000)

A: First of all, a Killi-hobbyist should be a member of his country Killifish Association, at least, and preferably of other Killifish Associations (same language or not) and read the regular newsletter and the beginner's guide. Secondly, he probably should buy two series of complementary publications: (1) Wildekamp's books ("A World of Killies" with 4 out of 5 volumes being published, and then all genera covered except for Rivulus to Valencia, scheduled for volume 5), encompassing digest information of each valid species of Killifish; (2) Seegers's books ("Aqualog" with all 3 volumes being published), encompassing the colour photos of nearly all known species of Killies with little accompanying information (pictorial codes). Alternatively, he may prefer to purchase the print-out of the present database (in part, obviously… but already 1028 pages!), as Killi-Data Catalogue 2006. If he is an advanced aquarist, he should add to his library the latest version of Killi-Data (2000, 2007) or of KMI (2001) or of Lexikon (2005), which index all available information on Killifishes, plus many other items. Third, he may be willing to read and keep some basic books on some Killifish groups, such as Rivulus (Huber, 1992), Cynolebiatins (Costa, 1995), Rivulins of the Old World (Scheel, 1968, 1990), Nothobranchius (Jubb, 1969; Seegers, 1985; Watters, 1997), Epiplatys (Neumann, 1980, 2000), the Aphyosemion cameronense superspecies (Eberl et al., 1997) or on the Killifish fauna of some countries, such as Cameroun (Amiet, 1987), Spain for Aphanius iberus and Valencia hispanica (Villwock et al., 1999), the South American annuals (Brousseau, 1990, revised 2001) or a Comparison between Old World and New World Tropical Cyprinodonts (Huber, 1998). Finally, there are a number of specialised books of real interest, such as the comparative experience of breeding Killifishes by international expert breeders (Langton, 1997) and the list of collecting expeditions by Killi-Hobbyists with code names ("Wild Collections of Killifish 1950-2000", Third Edition, Langton, 2001). To purchase these books, please note:
* Wildekamp's "A World of Killies" are available through most Killifish Associations and notably their editor, the AKA. 
* Seegers's "Aqualogs" and "Nothobranchius" are available through most Killifish Associations and notably the DKG. 
* Huber's Killi-Data Catalogue 2006 is available through the present author (155 €, fully in color, laser print only) with a message in the CONTACT forms
* Huber's "Review of Rivulus", "Killi-Data 2000" and "Comparison" are available through most Killifish Associations and if not there by the initial author, see ORDER and CONTACT
* Scheel's "Rivulins of the Old World" are available through most Killifish Associations and also their editor, TFH publications (second hand, for 1968) and sometimes through 
* Costa's "Cynolebiatins" is available through its editor, TFH publications and sometimes through 
* Nielsen's "Simpsonichthys e Nematolebias" (in Portuguese) is available through its editor, CABRAL publications and sometimes through 
* Lazara's "KMI" is available through its editor, the AKA. 
* Neumann's "Lexikon" is available through its editor, the DKG. 
* Neumann's "Prachtgrundkärpflinge. " (in German) is available through its editor, the DKG. 
* Watters's "Nothobranchius" is available through its editor, the AKA. 
* Neumann's "Epiplatys" is available through its editor, the DKG. 
* Villwock's "Fartet" is available through the Spanish Killifish Association, SEK.
* Eberl's "cameronense" is available through most Killifish Associations and notably the DKG and sometimes through
* Amiet's "Cameroun Aphyosemion" is available through most Killifish Associations and notably the KCF and sometimes through
* Brousseau's "South Americans Annuals" is available through its author:
* Langton's "Expert breeders" and "Wild Collections" are available through its author:
(Jean Huber, finalised, July, 2000; updated November 2002, October 2003, September 2004, April 2006, January 2007, December 2009).


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