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VVictorianictorian CCichlidichlid SSocietyociety 444:014:01 IIncorporatedncorporated MMarcharch 20152015 2015 committee@.org.au CONTENTS ::

COMMITTEECOCOMMIITTTETEE (E((Exeutive):Exexeuuttive)): Calendar ...... 2 [email protected]@ccicchlidds.s org.g au GregGrG egg NicolacopoulosNicicolo acacopopououlloos Cichlids ...... 3 [email protected]@ccicchlidds.s org.g au GrahamGrGrahaham RRoweowowe e [email protected]@ccicchlidds.s org.g au DavidDaDavivid GreenGrG eee n Newsletter Columnists Wanted – admin ...... 3 [email protected]@ccicchlidds.s org.g au TonyToTonyny FFergusoneerrgugusson COMMITTEE-AT-LARGE:COCOMMMMITTTEEE-ATAT-L-LARARGEG : Molecular and Fossil Evidence Place the Origin [email protected]@ccicchlidds.s org.g au DarylDaDaryyl HutchinsHuH tctchhiinss of Fishes Long After Gondwanan Rifting – [email protected]@ccicchlidds.s org.g au AndrewAAnndrd ewew CChallingsworthhahalllliningsg wowortrth Matt Friedman et al ...... 4-14 SUB-COMMITTEES:SUSUB-B-COOMMMMITITTETEESES: [email protected]@ccicchlidds.s org.g au PeterPeetteer RoRobinsonobbiinnssonon The EDASoV Show 2015 ...... 14-15 [email protected]@ccicchlh idds.s org.g au DarylDDaarryyl HutchinsHHuutcchih nnss QCG International Cichlid Conference 2015 .. 14-15 [email protected]@ccicchlidds.s org.g au DarylDDaaryyl HutchinsHuHutctchihinsns [email protected]@ccicchlidds.s org.g au VacantVaacacantn The Aquarist’s Notebook – Dr Albert J Klee .. .. 16-19 webslavewewebsb [email protected]@ccicchlh idds.s org.g au DarylDaDarryyl HutchinsHuH tcchihinss LIFELLIIFEFE MMEMBERS:EEMMBEBERRSS: Adventures In Failure: Benign Neglect GrahamGrGrahahamam RRowe,oowwe,e HHeinzeieinznz SStaude,taauuddee,, KevinKevevinin Archibald,Arcrchhibab ldd, KeithKKeeitith Patford,PaPattfforord,d, vs Over-Reacting – Michael Larsen ...... 20-23 DDaDannyannnny Genovese,GeGenonovveesee, DaDDarylaryyl HuHHutchinsutcchih nsns aandndd JohnJohnn MMcCormickcCcCoro mim ckk. FELLOWFFEELLLLOOWW OOFF THTTHEHE SOCIETY:SOSOCCIIEETTYY: GrahamGrG aha amm RRoweowwe ACA Convention 2015 ...... 24

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Associate (non-voting, no Joining Fee) $15 per Auction INSIDE FRONT: CORRESPONDENCE: Paratilapia sp East Coast Small Spot, female – John McCormick. THE SECRETARY INSIDE BACK: :: Victorian Cichlid Society Inc Steatocranus glaber – Alf Stalsberg. 2 Alern Court, Nunawading 3131 Victoria Australia [email protected] © All images copyright by their respective owners. [email protected] for information. IMAGE: John McCormick cichlidpower.org.au 1 AUTHORS Newsletter WANTED Columnists URGENTLY! Wanted OR NOT! Are you ready to dip your toes into TThehe eCCichlidsichlids the Brave New World of electronic IT’S YOUR CHOICE publishing? In a modest, beginner’s CCompendiumompendium way, of course. You can help without AFTER ALL THE EDITOR getting out of your chair. DOES HAVE A MAGIC WAND It just does not get any easier than 4480+80+ this. Any member who so desires mmagazines.agazines. can be authorised to log-in to the the MMATE’SATE’S RRATESATES MailChimp system and compose their own section of the eNews newsletter. CCOMPUTEROMPUSSERVICESTEREVICRES EEveryvery eeditiondition (So can aquarial superstars, but that’s another story.) ooff ‘‘TheThe CCichlidichlid eNews is much less formal and CCOMPUTEROMPUTER RREPAIRS,EPAIRS, easier to write for than the magazine, MMAINTENANCEAINTENANCE MMonthly’,onthly’, as it is not a venue for full-blown ‘iCCichlid’ichlid’ aandnd articles; anything even vaguely fi sh- © Copyright, Victorian Cichlid Society Incorporated 2015 & UUPGRADESPGRADES related is ok. If people were breeding fi shes and Any non-profit group may re-use material from this magazine for non-commercial purposes, unless it is ‘eNNews’ews’ eeverver AALSOLSO registering their spawnings, no doubt copyright by the author, provided due credit is given to the author and iCichlid. A copy of the relevant the Breeder’s Achievement Award DDesktopesktop PPublishingublishing SServices:ervices: pproduced.roduced. publication (a pdf is fine) is to be forwarded to the author/s, and/or [email protected]. Enquiries re Chairperson would be taking great delight in entering reports this way. the use of material in any other way may be directed to [email protected]. SScan/Convertcan/Convert DDocumentsocuments ttoo PPDF;DF; OOCRCR DDocumentsocuments ttoo EEditableditable FFiles;iles; SStilltill oonlynly $$2525 The same goes for the person/s in charge of the Show Table, social stuff, NB: the Society’s Website, including this magazine, is archived by the State Library of Victoria’s Pandora DDTPTP TTuition;uition; DDataata IInput;nput; system, an online archive in which selected websites/online publications are preserved and made (($30$30 fforor nnon-members)on-members) etc. But you could probably think of RRepair/Installepair/Install NNetworks/etworks/ a multitude of other uses for it that permanently available to the public for research and reference. CComputers.omputers. oonn aann 8 GGBB haven’t even occurred to me. There are standard boxes in the UUSBSB drive.drive. template just waiting for your input. 00430430 032032 330404 But if they are all used-up, it is a ((OtherOther mmediaedia oonn rrequestequest simple matter of dragging a suitable box from the right-hand panel and .... oorr BBYO.)YO.) placing it in the left-hand column. VCS 2015 CALENDAR Click the box and start typing! What n/a eNews iCichlid eNews iCichlid eNews iCichlid eNews iCichlid eNews iCichlid eNews could be easier? JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC E-mail [email protected] ✔ ✔ Auction Auction AAGMGM and you could be in business in n/a << Meetings/Auction >> No Meetings << Meetings/Auction >> minutes. p

2 facebook.cichlids.org.au cichlidpower.org.au 3 From rspb.royalsocietypublishing.org © 2013 the Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License creativecommons.org/licenses/by/3.0/. MMolecularolecular andand FossilFossil EvidenceEvidence PlacePlace thethe OriginOrigin ofof Figure 1. Congruent molecular and palaeontological time scales CCichlidichlid FishesFishes LongLong AfterAfter GondwananGondwanan RiftingRifting place the origin of cichlid fishes in the Late Cretaceous-Eocene Matt Friedman1, Benjamin P Keck2, Alex Dornburg3, Ron I Eytan3, interval, substantially post- Christopher H Martin4,†, C Darrin Hulsey2, Peter C Wainwright4 and Thomas J Near3 dating Gondwanan rifting. (a) Molecular phylogeny for Cichlidae 1Department of Earth Sciences, University of Oxford, South Parks Road, Oxford OX1 3AN, UK calibrated using fossils belonging 2Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN 37996, USA to non-cichlid groups (full 3Department of Ecology and Evolutionary Biology and Peabody Museum of Natural History, Yale University,New Haven, CT 06520, USA phylogeny provided in electronic 4 Department of Evolution and Ecology, University of California, Davis, CA 95616, USA supplementary material, figures S1 and S2). (b) Bayesian point estimates and 95% CIs for the timing of cichlid origin based on the distribution of cichlid fossils and the availability of freshwater sedimentary deposits of Triassic- Recent age on Gondwanan landmasses that bear articulated fish remains. The top estimate is derived from the record of landmasses inhabited by extant cichlids, and the bottom estimate is derived from the record of all Gondwanan landmasses. The density of all Gondwanan horizons bearing articulated freshwater fish fossils is indicated by the histogram at the bottom of the figure (densities including disarticulated material are ichlid fishes are a key model system in the study of adaptive radiation, speciation and evolutionary given in electronic supplementary developmental biology. More than 1600 cichlid species inhabit freshwater and marginal marine material). Grey bars indicate total horizon density. Pink bars indicate environments across several southern landmasses. This distributional pattern, combined with parallels the density of the subset of fossil between cichlid phylogeny and sequences of Mesozoic continental rifting, has led to the widely accepted fish horizons that bear cichlids. C (c) Bayesian point estimates and hypothesis that cichlids are an ancient group whose major biogeographic patterns arose from Gondwanan vicariance. 95% CIs for the timing of cichlid origin based on successive fossil Although the Early Cretaceous (ca 135 Ma) divergence of living cichlids demanded by the vicariance model now outgroups to the clade. The represents a key calibration for teleost molecular clocks, this putative split pre-dates the oldest cichlid fossils by two estimates reflect competing hypotheses for the earliest fossil nearly 90 Myr. Here, we provide independent palaeontological and relaxed molecular-clock estimates for the time examples of some outgroups. of cichlid origin that collectively reject the antiquity of the group required by the Gondwanan vicariance scenario. The top estimate is based on the oldest proposed outgroup ages and The distribution of cichlid fossil horizons, the age of stratigraphically consistent outgroup lineages to cichlids and the bottom estimate is based on relaxed-clock analysis of a DNA sequence dataset consisting of 10 nuclear genes all deliver overlapping estimates for the youngest proposed outgroup ages. Cichlid illustrations, from top crown cichlid origin centred on the Palaeocene (ca 65-57 Ma), substantially post-dating the tectonic fragmentation to bottom: Etroplus, , , Hemichromis, of Gondwana. Our results provide a revised macroevolutionary time scale for cichlids, imply a role for dispersal in Steatocranus, Altolamprologus and generating the observed geographical distribution of this important model clade and add to a growing debate that Tropheus. Continental arrangements based on palaeogeographic questions the dominance of the vicariance paradigm of historical biogeography. reconstructions by R. Blakey, originally presented in [7].

4 facebook.cichlids.org.auforums.cichlids.org.au cichlidpower.org.au 5 1. Introduction places this continental fragmentation event or inferring that the advanced morphol- share only one common feature in their Cichlid fishes, along with Darwin’s finch- in the Early Cretaceous (ca 135 Ma) [11]. ogy of the earliest fossil cichlids implies calculations: both are necessarily con- es and Caribbean Anolis lizards, repre- However, the stratigraphically oldest fossil a long, and as yet unsampled, palaeon- strained by the minimum age for cichlids as sent a key vertebrate model system for cichlids are Eocene in age (approx. 46 Ma) tological history of the group [9,10,13]. imposed by the oldest fossil example(s) of understanding the evolutionary assembly [12,13], implying a gap of approximately 90 The seemingly ambiguous signal of the group. As an independent assessment of biodiversity [1,2]. Despite the group’s Myr in the early history of the group. This, palaeontological data with respect to of the divergence times estimated from prominence in biological research, a con- along with the absence of Early Cretaceous the question of cichlid origin is symp- palaeontological data, we conducted a sistent macroevolutionary time scale and fossils belonging to more inclusive and tomatic of a qualitative approach to relaxed-molecular-clock analysis for cich- biogeographic history for cichlids has taxonomically diverse clades that con- an inherently quantitative problem. lids and Ovalentaria [19], a percomorph remained elusive [3-6]. For nearly four tain cichlids, has led some to abandon Invoked stratigraphic gaps are neither lineage that includes cichlids. Ourdataset & Reptiles decades, the study of deep cichlid evo- the orthodoxy of Mesozoic vicariance in ad hoc contrivances nor trivial incon- includes 10 protein-coding nuclear genes lutionary history has been dominated by favour of Cenozoic dispersal to explain the veniences to be dismissed as for 89 cichlids and 69 non-cichlid species ATAT CRANBOURNE vicariance models of biogeography that present-day distribution of cichlids [12,14]. non-evidence; they are hypotheses of Percomorpha. link the present-day distribution of the The complete absence of fossil cichlids amenable to statistical interrogation. 2. Material and methods group to the tectonic fragmentation of from many former Gondwanan landmasses In order to provide a robust time scale for (a) Estimating time of evolutionary origin using the For the right advice and all your the supercontinent of Gondwana during would seem equally problematic for the cichlid diversification and select between distribution of cichlid-bearing fossil horizons needs the mid to late Mesozoic (ca 135-90 Ma; vicariance hypothesis, but has received competing biogeographic hypotheses, we One method of estimating credible inter- figure 1) [8,9]. Continued investigation of surprisingly little attention. For example, applied three semi-independent approach- vals (CIs) on stratigraphic durations draws cichlid intrarelationships, including phy- the Australian fossil record contains sev- es in estimating the age of crown-group on the number of fossil horizons within logenetic analysis of molecular sequence eral fish-bearing freshwater deposits of Cichlidae. Our first two methods are pal- the sampled range of the group of inter- data, has shown congruence between the Mesozoic and Cenozoic age, but no fossil aeontological, and draw on (i) the distribu- est. The simplest approach assumes that OPEN 365 DAYS order of divergences among geographi- cichlid is known from the continent. While tion of fossil horizons yielding cichlids and fossil horizons are distributed at random cally restricted cichlid clades and proposed it is clear that assembly of the composi- those that might plausibly yield cichlids (i.e. [20,21], but the potential for fossil recovery 8 am-6 pm sequences of continental break-up [10]. tionally distinctive Australian freshwater fish-bearing freshwater deposits on former undoubtedly varies over time as a conse- The vicariance hypothesis of cich- fish fauna has a complex history stemming Gondwanan landmasses) [17], and (ii) the quence of a heterogeneous rock record. lid historical biogeography has become from isolation, aridification and marine stratigraphic distribution of more inclusive Marshall [17] developed a more general Full range of aquarium, tropical and coldwater fi sh; so entrenched that the rifting history invasion coupled with the persistence of teleost lineages (meaning clades of higher method that permits non-uniform preser- Large range of reptiles and accessories; of Gondwana is routinely used to cali- ancient lineages [15], this complexity does taxonomic rank) that contain cichlids [18]. vation by using an empirically informed Tanks and cabinets; Full set-ups from 2ft to 8ft; brate teleost molecular clocks [3,6], with not undermine the prediction of the vicar- These techniques relate directly to two function that quantifies potential for fos- the consequence that this hypothetical iance model that cichlids should have contrasting arguments that emerge repeat- sil recovery. We have applied this logic Wide range of live and frozen foods; Plants; scenario now directly influences estimates been widely distributed across Gondwanan edly in palaeontological debates concern- in conjunction with a Bayesian approach Full range of conditioners; Free water testing. of the evolutionary time scale for more landmasses during the Mesozoic [9]. ing the chronology of cichlid evolution: that provides a statistically appropriate than half of all modern vertebrate diversity. These palaeontological arguments have either that the record of freshwater fishes framework for discussing the probability 14 Cooper Court, Cranbourne, VIC If the distribution of modern cichlids been dismissed as ‘non-evidence’ by generally, and cichlids specifically, is suf- of clade origin within certain stratigraphic is attributable purely to Gondwanan break- advocates of cichlid vicariance [16]. Some ficiently poor that the absence of Mesozoic intervals [20]. Our results are conditioned up, then it necessarily follows that the com- authors have even suggested that the cichlid fossils is unsurprising, or that the on the prior assumption that cichlids are 03 5995 5066 mon ancestor of all living cichlids originated fossil record supports the notion of cich- minimum age of origin for a series of more post-Palaeozoic in age (i.e. they originated aquariumsatcranbourne.com.au no later than the final separation between lids deep within the Mesozoic, citing the inclusive lineages of teleost fishes pre- in the Triassic or later), which is consistent Madagascar-India and South America- high probability of non-preservation for cludes the origin of cichlids deep within with the fossil record and does not exclude Africa-Arabia. Current geological evidence freshwater taxa of Cretaceous age [9] the Mesozoic. Significantly, these methods the possibility of Gondwanan vicariance.

6 facebook.cichlids.org.au cichlidpower.org.au 7 We assembled a database of known fos- based on (i) the fossil records of Gond- An account of the outgroups used in analy- Table 1. Posterior molecular age estimates for major lineages of Cichlidae. sil occurrences of cichlids on Gondwanan wanan landmasses currently inhabited by sis is provided in the electronic supple- Ages refer to crown groups. landmasses based on the literature cichlids (South and Central America includ- mentary material. In some cases, there is PPRODUCTRODUCT 95% highest posterior (see electronic supplementary material). ing the Caribbean, Africa, Madagascar, disagreement surrounding the identity of Clade mean age (Ma) density interval (Ma) Different geological formations (or localities India, Arabia; cichlid fossils are known from the earliest representatives of these line- Cichlidae 64.9 57.3-76.0 where there is no formalized lithostrati- all of these regions except Madagascar EEFFICACYFFICACY ages. To accommodate uncertainty, two Etroplinae (India and Madagascar) 36.0 30.3-42.2 graphic framework) were assumed to and India), and (ii) these records combined sets of calculations were completed: one Ptychochrominae (Madagascar) 38.2 31.7-46.4 represent distinct sampling horizons. The with those of Australia and Antarctica, AALERT!LERT! using the oldest proposed minimum age unnamed Afro-American clade 46.4 40.9-54.9 function for the potential recovery of fos- former Gondwanan landmasses that lack for a clade and the other applying the (neotropics) 29.2 25.5-34.8 sil cichlids was estimated by tabulating cichlids but would be predicted to have youngest. Collectively, these paired analy- Pseudocrenilabrinae (Africa) 43.7 38.2-51.6 the number of sedimentary horizons (for- once been inhabited by the group under IIMPORTANTMPORTANT ses provide upper and lower estimates unnamed east African clade 8.0 6.9-9.5 mations or localities) that meet three key the vicariance hypothesis. For both, we NNOTICEOTICE TTOO AALLLL of CIs for divergence times given present most recent common ancestor of L. Malawi and L. Victoria radiations 2.3 1.7-3.1 criteria. First, candidate deposits must calculated CIs based on the record of all understanding of both the fossil record and Crater Lake Barombi Mbo (Cameroon) 1.4 0.8-2.3 be present on former Gondwanan land- cichlid fossils and estimated range exten- OOWNERSWNERS OFOF teleost interrelationships. These age esti- masses. Second, candidate deposits must sions based on articulated cichlid remains TTHEHE eCCichlidichlid mates are conditioned on a prior assump- represent freshwater environments. Third, alone combined with appropriate recov- tion that divergence occurred after a user- ing. Alignments of the DNA sequences (d) Fossil calibration age priors candidate deposits must have the potential ery potential functions generated from the CCOLLECTIONOLLECTION specified hard upper bound. This bound from the individual genes were constructed For each fossil calibration prior, we iden- to yield fossils of cichlids, were this group subset of deposits that yield complete fish applies to the divergences of all groups from the inferred amino acid sequenc- tify the calibrated node in the percomorph present. Sites yielding fish fossils (includ- specimens. This modified procedure is DDueue toto improvementsimprovements considered, not only the focal clade. es. Thirty data partitions were designated phylogeny, list the taxa that represent the ing but not restricted to cichlids)meet this more conservative and reflects the very real ttoo tthehe wwayay wwee We have therefore selected the Carbon- that corresponded to the three separate first occurrence of the lineage in the fos- final criterion. The nature of fossils (articu- possibility that the earliest cichlids might ppublish,ublish, allall previouslypreviously iferous (Serpukhovian) Discoserra, a puta- codon positions for each of the 10 protein- sil record, describe the character states lated or fragmentary) from sites satisfy- be recognized only on the basis of articu- iissuedssued discs,discs, andand tive stem neopterygian [22], as defining an coding genes. A phylogeny of the aligned that justify the phylogenetic placement of ing these conditions was also recorded. lated remains, as their isolated fragments upper bound of 322.8 Ma (see the elec- DNA matrix was inferred using maximum- the fossil taxon, provide information on Because of uncertainty surround- might be too character-poor, too general- UUSBSB ddrivesrives areare nownow tronic supplementary material). likelihood and relaxed-clock analyses using the stratigraphy of the rock formation(s) ing age assessments, uniform recov- ized or both to permit reliable taxonomic oobsolescent.bsolescent. (c) Collection of sequence data, phylogenetic a random local molecular-clock model in bearing the fossil, give the absolute age ery potential was assumed within each attributions. WWhichhich mmeanseans tthathat ttheyhey wwillill analyses and relaxed molecular clocks the computer program BEAST v. 1.6 (figure estimate for the fossil, outline the prior epoch-level stratigraphic bin, with rela- (b) Estimating time of evolutionary origin using the ccontinueontinue toto wwork,ork, bbutut iitt iiss nnoo Standard phenol-chloroform extraction 1; electronic supplementary material, figure age setting in the BEAST relaxed-clock tive recovery potential given by the num- distribution of ages of outgroups to cichlids llongeronger possiblepossible toto updateupdate thethe protocol or Qiagen DNeasy Blood and S1) [23,24]. DNA sequences are depos- analysis and provide any additional notes ber of candidate horizons present in a Hedman [18] devised a Bayesian ccollectionollection bbyy ssimplyimply addingadding fi lleses Tissue kits were used to isolate DNA from ited on GenBank KF556709-KF557487. on the calibration [25]. Each calibration is given interval divided by its duration. approach for constraining the time of ori- ttoo tthehe eexistingxisting structure,structure, andand tissue biopsies sampled from 158 spe- Aligned gene sequences used in phyloge- numbered and the phylogenetic placement Ambiguity surrounds the age of many gin of a clade based on the distribution tthathat iiss nnotot hhowow iitt shouldshould be.be. cies of percomorph teleosts that included netic analyses, phylogenetic trees result- of the calibration is highlighted in the elec- freshwater deposits. In this study, of stratigraphic ages of successive out- SSo,o, iiff yyouou oownwn tthehe ccollectionollection onon 89 species of Cichlidae (electronic sup- ing from RAXML and BEAST analyses, tronic supplementary material, figure S3. im precisely dated deposits are given their groups. This method requires that out- ddiscisc oror USBUSB youyou cancan acquireacquire plementary material, table S6). Previously files formatted for BEAST analyses and Full justification of our calibrations is given oldest plausible age. This approach sys- groups appear in the fossil record in an a completelycompletely uup-to-datep-to-date 8GB8GB published PCR primers (see the elec- files used to estimate the age of cichlids in the electronic supplementary material. UUSBSB drivedrive forfor onlyonly $5$5 (which(which tematically biases analysis towards older order matching phylogeny and that they tronic supplementary material) were used using palaeontological data are available Because we look to provide a critical test of bbarelyarely recoversrecovers thethe costcost ofof thethe age estimates for the time of clade origin, pre-date or are contemporary with the first to amplify and sequence exons from 10 from the dryad digital repository (dx.doi. competing models of cichlid biogeography, uupgradedpgraded drive).drive). thereby providing a more generous test appearance of the focal clade. Such perfect nuclear genes (ENC1, Glyt, myh6, plagl2, org/10.5061/dryad.48f62). Fossil-based we have not assumed Gondwanan vicari- of the Gondwanan vicariance hypothesis. congruence is rare in empirical examples, IIff youyou can’tcan’t makemake itit toto a meeting,meeting, e-maile-mail Ptr, rag1, SH3PX3, sreb2, tbr1 and zic1). age constraints were applied to 10 nodes ance a priori and did not use the timing of [email protected]@cichlids.org.au aandnd wwe’lle’ll These data were used to generate point and we adopt a proposed solution that aarrangerrange ssomethingomething fforor yyou.ou. Amplified gene copies were cleaned and in the percomorph phylogeny (electronic the fragmentation history of this supercon- estimates and 95% CIs for cichlid origin conservatively excludes inconsistent ages. used as templates for DNA cycle sequenc- supplementary material, figure S2). tinent to inform calibrations in the relaxed-

8 facebook.cichlids.org.au cichlidpower.org.au 9 molecular-clock analyses. Furthermore, we fossil record, but the upper bounds of crown acanthomorph lineages are likely Based on the timing of Gondwanan frag- 4. Discussion horizons or our database of outgroup- have not included any internal calibra- the CIs do increase by more than 10 Ma. to be sufficiently ancient to have vicariant mentation events, crown cichlids should (a) Congruence between palaeontological and based age constraints. We interpret the tions within Cichlidae, so that our relaxed- Depending on the scope of geographical Gondwanan distributions, as we estimate occur in the Early Cretaceous or Late molecular time scales for cichlid evolution convergence of these three semi-inde- molecular-clock estimate of the evolution- analysis, we estimate the time of cichlid the age of the group as between 106.4Ma Jurassic [3,6,9,10]; however, the Bayesian The application of two contrasting pal- pendent approaches, which all deliver age ary time scale for the group is truly inde- origin based only on articulated remains as (95% CI: 98.5-132.2Ma) and 109.2Ma (95% random local molecular-clock analyses aeontological approaches in calculating estimates for cichlids that are within error pendent of its fossil record, which contrib- ranging from 59.8 Ma (95% CI: 56.1-75.1 CI: 98.5-136.0Ma). The most restrictive place the origin of the modern cichlid temporal range extensions yields strikingly of one another, as a consequence of genu- utes to our palaeontological estimates of Ma; landmasses inhabited by modern cich- group containing cichlids that we can date radiation near the Cretaceous- congruent time scales for cichlid evolution. ine evolutionary signal that strongly contra- divergence times (see §2). lids) to 60.2 Ma (95% CI: 56.1-77.8 Ma; all with this method and which is of sufficient Palaeogene boundary (figure 1 and Both methods provide point estimates for dicts the time scales for cichlids demanded 3. Results Gondwanan landmasses). The Gondwanan apparent antiquity to have been affected by table 1; electronic supplementary mate- the origin of the group that range between by hypotheses of Gondwanan vicariance. the initial rifting of Gondwana is Eurypterygii, 57 and 60 Ma (Palaeocene), and strongly Our three approaches to estimating a vicariance hypothesis requires a pre- rial, figure S1), with a mean age esti- (b) The timing of cichlid diversification: congruence time scale of cichlid origin and diversifi- Eocene record of cichlids that is roughly the radiation containing Acanthomorpha, mate of 64.9 Ma (95% CI: 57.3-76.0 Ma). reject the possibility that crown cichlids and incongruence cation yield overlapping CIs that diverge 10-30 times worse than their recorded Myctophiformes and Aulopiformes [25]. The estimated age of the most recent are sufficiently old to have been affected Our estimates for the time of cichlid significantly from the predictions made by fossil history, with rescaled recovery poten- Our estimates for the time of origin for common ancestor (MRCA) of cichlids by the initial rifting of Gondwana. Instead, origin are congruent not only with one the Gondwanan vicariance biogeographic tials conditioned on point estimates for this major teleost clade range between and their sister lineage, Pholidichthys, is upper limits for the origin of cichlids lie another, but also with previous molecular hypothesis, and are discussed in turn in the origin of the group at 135 Ma ranging 131.1Ma (95% CI: 104.9-163.2Ma) and also younger (mean: 103.7 Ma; 95% CI: consistently within the late Late Cretaceous. time scales for the evolution of this group §3a,b (figure 1). from 2.8-3.3% (all fossils) to 6.6-6.9% 142.1 Ma (95% CI: 126.2-166.2 Ma). 92.0-118.4 Ma) than the initial rifting of This congruence is particularly compelling that do not assume a Gondwanan vicari- because the methods that yielded these (a) Palaeontological time scales for cichlid evolution (articulated fossils only) of their original (b) A molecular time scale for cichlid evolution Gondwana at approximately 135 Ma [11]. ance scenario a priori [4,5,28-33]. The old- comparable results share only one similar- The distribution of cichlid-bearing fos- values. Classical confidence intervals deliver The phylogeny of Ovalentaria and the The mean estimated age of the MRCA of est such estimates from previous work sil horizons, combined with an empiri- similar results to the Bayesian estimates (see major cichlid lineages inferred from the 10 the African and Neotropical cichlids was ity in their calculations: both are constrained are early Late Cretaceous [3], pre-dating cally informed function describing fossil electronic supplementary material, table S2). nuclear genes is similar to previous molec- 46.4 Ma (95% CI: 40.9-54.9 Ma), post-dat- by the minimum age for cichlids as imposed our proposed time of origin by roughly recovery potential, indicates an age of Analysis of outgroup ages provides broadly ular and morphological analyses [8,10,19], ing the final separation of Africa and South by the oldest fossil example of the group. 35-45 Myr. However, these more ancient origin for cichlids in the Late Cretaceous similar estimates for the timing of cichlid with Etroplinae (India, Madagascar) America by more than 40 Myr. The cichlid Our molecular time tree provides a dates derived from analysis of mitochon- or Palaeocene. If only the records of land- origin to those derived from the distribu- resolved as the earliest-diverging clade time tree confirms ages estimated in previ- mean estimate for the timing of cich- drial sequences, which are characterized masses that are currently inhabited by tion of cichlid fossil horizons, in terms of and Ptychochrominae (Madagascar) as the ous studies for the east African [26] (mean: lid origin in the Palaeocene, but can- by high rates of nucleotide substitution that cichlids are considered, the time of origin both the magnitude of point estimates sister lineage to the unnamed clade that 8.0 Ma; 95%CI: 6.9-9.5 Ma) and Cameroon not reject the possibility that the group might bias clock analyses towards older of the clade is estimated as 59.2 Ma (95% and the degree of uncertainty surround- contains the African (Pseudocrenilabrinae) crater lake Barombi Mbo [27] radiations arose as early as the Late Cretaceous. estimated times of divergence [25,34-36]. CI: 56.1-67.6 Ma). By contrast, a slightly ing them. We find a mean age of 60.7 Ma and Neotropical (Cichlinae) cichlid line- (mean: 1.4 Ma; 95%CI: 0.8-2.3 Ma), verify- This result is consistent with other recent Generally, the only molecular-clock analy- younger age estimate of 57.8 Ma (95% CI: (95% CI: 46.8, 90.1 Ma) using the old- ages (figure 1; electronic supplementary ing relatively young ages for these remark- molecular-clock estimates for the origin of ses to deliver time scales consistent with 56.1-62.4 Ma) is obtained if the record of est possible fossil ages for outgroups. material, figure S1). The 10 nuclear gene able examples of adaptive radiation (figure cichlids that do not assume Gondwanan the predictions of the vicariance hypoth- all Gondwanan landmasses is considered. The time scale for cichlid origin is predict- phylogeny preserves the parallels between 1 and table 1; electronic supplementary vicariance for the group a priori, and which esis were themselves calibrated using a Restricting the scope of analysis to con- ably more recent using the youngest pos- patterns of cichlid interrelationships and material, figure S2). The age estimate in the range in age from Late Cretaceous to combination of age constraints from the sider articulated remains alone provides sible fossil ages for outgroups, but only the fragmentation history of Gondwana 10 nuclear gene inferred time tree closest Eocene [28-33]. In terms of point estimates fossil record and Gondwanan fragmenta- a more conservative means of estimating slightly so, with a mean age of 57.0 Ma that has led to the prominence of vicari- to the timing of Gondwanan fragmentation and surrounding uncertainty, our revised tion events [3,5,6,37]. There is no pub- the time of origin for cichlids, because (95% CI: 46.8-81.2 Ma). Using this same ance biogeographic scenarios for this lin- is that of the inclusive (mean: 123.5 Ma; molecular time scale is entirely consistent lished relaxed-molecular-clock analysis early members of this group might not be approach, it is also possible to determine eage [9]. However, the Bayesian random 95% CI: 111.4-136.2 Ma), but unnamed, with the ages derived from analyses of the that results in an Early Cretaceous or recognized on the basis of less diagnos- probable times of origin for a series of more local molecular-clock analyses yield age percomorph clade that contains more than fossil record alone (figure 1). It is important Jurassic origin of cichlids that is inde- tic skeletal debris. Point estimates for the inclusive clades that contain Cichlidae: estimates for the origin of cichlids consist- one-quarter of all living vertebrate species to note that our relaxed-molecular-clock pendent of the ages implied by the timing timing of cichlid origin under this approach Ovalentaria, Percomorpha, Acanthopterygii, ent with those derived from analysis of (approx. 16 570 species), including cichlids analysis shares no palaeontological data of the fragmentation of western Gondwana. do not change drastically from those Acanthomorpha, Eurypterygii, Euteleostei fossils alone (figure 1 and table 1; elec- (see electronic supplementary material, fig- in common with either our analysis of Our palaeontological time scales obtained using the entirety of the cichlid andTeleostei. This exercise implies that no tronic supplementary material, figure S1). ure S2). the distribution of cichlid-bearing fossil for Cichlidae constrain only the ori-

10 facebook.cichlids.org.au cichlidpower.org.au 11 gin of the group, but our time-calibrat- ing phylogenies (e.g. bootstrap resam- lution reject Gondwanan vicariance as a also possible that freshwater plumes, such versions of this work, and we thank A Murray and M and beyond. Syst. Biol. 61, 1001-1027. (doi:10.1093/sysbio/sys060) Vences for their insightful reviews and J Hutchinson 20. Strauss D, Sadler PM. 1989 Classical confidence intervals and Bayesian ed phylogeny permits investigation of the pling scores or Bayesian posterior prob- viable mechanism for the modern distri- as that produced by the modern Congo probability estimates for ends of local taxon ranges. Math. Geol. 21, 411- for editorial assistance. 427. (doi:10.1007/BF00897326) timing of deep divergences within the clade abilities). We also note that some phyloge- bution of the group, but they demand what River [56], provided corridors of brackish 21. Marshall CR. 1990 Confidence intervals on stratigraphic ranges. Funding statement: Paleobiology 16, 1-10. (table 1). We estimate the divergence of netic hypotheses derive from successive can only be considered a series of highly surface water that could have permitted This research was supported by the Peabody 22. Hurley IA et al. 2007 A new time-scale for ray-finned fish evolution. Proc. Museum of Natural History and grants awarded to R. Soc. B 274, 489-498. (doi:10.1098/rspb.2006.3749) South American and African cichlids as reweighting exercises [39], while others unlikely transoceanic dispersal events. Like migration by freshwater taxa across a nar- PCW and TJN from the National Science Foundation, 23. Drummond AJ, Suchard MA. 2010 Bayesian random local clocks, or one rate to rule them all. BMC Biol. 8, 114. (doi:10.1186/1741-7007-8-114) Eocene, with the origin of the African assume restricted placement of fossil the fossil record, the salinity tolerance of rower marine barrier during the Palaeogene. USA (DEB-0444842, DEB-0716155, DEB-0717009, 24. Drummond AJ, Rambaut A. 2007 BEAST: Bayesian evolutionary analysis by DEB-0732642, ANT-0839007, DEB-1060869, DEB- sampling trees. BMC Evol. Biol. 7, 214. (doi:10.1186/1471-2148-7-214) cichlid crown within the same interval. species prior to analysis [13]. There is cichlids has been subjected to contrasting Coeval examples of migration provide 1061806 and DEB-1061981). MF is supported by 25. Near TJ, Eytan RI, Dornburg A, Kuhn KL, Moore JA, Davis MP, Wainwright PC, Friedman M, Smith WL. 2012 Resolution of ray-finned fish phylogeny This is consistent with the placement no doubt that Lumbrera cichlids are sig- interpretations; it has been cited as both circumstantial evidence for the possibil- the Natural Environment Research Council (NERC and timing of diversification. Proc. Natl Acad. Sci. USA 109, 13 698-13 NE/I005536/1). CHM was supported by the National 703. (doi:10.1073/pnas. 1206625109) of the middle Eocene (approx. 46 Ma) nificant on account of their antiquity and consistent [12] and inconsistent [10] with ity of trans-Atlantic dispersal, and geo- Geographic Society (Young Explorers Grant) and the 26. Koblmu¨ller S, Egger B, Sturmbauer C, Sefc KM. 2010 Rapid radiation, † graphical factors during the Eocene would American Philosophical Society (Lewis and Clark ancient incomplete lineage sorting and ancient hybridization in the Mahengechromis as an early crown pseu- geographical provenance. However, in the marine dispersal. Experimental evidence endemic Lake Tanganyika cichlid tribe Tropheini. Mol. Phylogenet. Evol. Field Grant). appear to have facilitated the crossing, 55, 318-334. (doi:10.1016/j.ympev. 2009.09.032) docrenilabrine [12]. By contrast, our esti- absence of demonstrably robust phyloge- points to high salinity tolerance in some 27. Schliewen UK, Klee B. 2004 Reticulate speciation in Cameroonian crater References: lake cichlids. Front. Zool. 1, 1-12. (doi:10.1186/1742-9994-1-5) mated Eocene-Oligocene age for the South netic placements of these fossil lineages cichlids [50,51], but the fact that no cichlid but cichlid migration across the south 1. Wagner CE, Harmon LJ, Seehausen O. 2012 Ecological opportunity and 28. Alfaro ME, Santini F, Brock C, Alamillo H, Dornburg A, Rabosky DL, sexual selection together predict adaptive radiation. Nature 487, 366-369. Carnevale G, Harmon LJ. 2009 Nine exceptional radiations plus high American cichlid crown contradicts pub- within a group well known for convergent inhabits the open ocean indicates that long- Atlantic and other marine barriers never- (doi:10.1038/nature11144) turnover explain species diversity in jawed vertebrates. Proc. Natl Acad. 2. Kocher TD. 2004 Adaptive evolution and explosive speciation: the cichlid Sci. USA 106, 13 410-13 414. (doi:10.1073/pnas.0811087106) lished interpretations of the fossil cichlids morphological evolution [42], their exact distance marine migration is improbable. theless remains an extraordinary claim. fish model. Nat. Rev. Genet. 5, 288-298. (doi:10.1038/nrg1316) 29. Santini F, Harmon LJ, Carnevale G, Alfaro ME. 2009 Did genome 3. Azuma Y, Kumazawa Y, Miya M, Mabuchi K, Nishida M. 2008 Mitogenomic However, the evolutionary time scale duplication drive the origin of teleosts? A comparative study from the ‘Faja Verde’ level of the Lumbrera implications for the timing of major events Dispersal across the south Atlantic would evaluation of the historical biogeography of cichlids toward reliable dating of diversification in ray-finned fishes. BMC Evol. Biol. 9, 164. of teleostean divergences. BMC Evol. Biol. 8, 215. (doi:10.1186/1471- inferred for cichlids on the basis of both (doi:10.1186/1471-2148-9-194) Formation of Argentina. These fossils are in cichlid evolution are likely to remain appear to be especially unlikely, given 2148-8-215) 30. Near TJ et al. 2013 Phylogeny and tempo of diversification in the 4. Vences M, Freyhof J, Sonnenberg R, Kosuch J, Veith M. 2001 Reconciling fossils and molecules demands that this superradiation of spiny-rayed fishes. Proc. Natl Acad. Sci. USA 110, 12 often cited as early-middle Eocene in age ambiguous. that it measured roughly 1000 km [52] fossils and molecules: Cenozoic divergence of cichlid fishes and the 738-12 743. (doi:10.1073/pnas.1304661110) biogeography of Madagascar. J. Biogeogr. 28, 1091-1099. (doi:10.1046/ [13,38,39], leading to calibration minima of in width by the time of the inferred diver- hypothesis is given serious considera- 31. Crottini A, Madsen O, Poux C, Strauss A, Vieites DR, Vences M. 2012 (c) Comparison with other putative examples of j.1365-2699.2001.00624.x) Vertebrate time-tree elucidates the biogeographic pattern of a major 5. Genner MJ, Seehausen O, Lunt DH, Joyce DA, Shaw PW, Carvalho GR, Turner 49 Ma in recent molecular clock studies [6]. Gondwanan vicariance gence between South American and tion rather than being dismissed a priori. biotic change around the K-T boundary in Madagascar. Proc. Natl Acad. GF. 2007 Age of cichlids: new dates for ancient lake fish radiations. Mol. Sci. USA 109, 5358-5363. (doi:10.1073/pnas. 1112487109) However, the hard minimum for the age Among vertebrates assumed to have lim- African cichlids in the Eocene (figure 1). Our estimation of consistent palaeonto- Biol. Evol. 24, 1269-1282. (doi:10.1093/molbev/msm050) 6. Lo´pez-Ferna´ndez H, Arbour JH, Winemiller KO, Honeycutt RL. 2013 Testing 32. Betancur RR et al. 2013 The tree of life and a new classification of bony of these fossils is 33.9 Ma, which derives ited dispersal ability across marine barriers, Despite the presence of a substantial logical and molecular ages for the origin of for ancient adaptive radiations in neotropical cichlid fishes. Evolution 67, fishes. PLoS Curr. (doi:10.1371/currents.tol.53ba26640df0ccaee75bb165 1321-1337. c8c26288) from radiometric dating of overlying tuff cichlids are not unique in showing a broad marine barrier, it is clear that at least cichlids adds to a growing number of stud- 7. Blakey RC. 2008 Gondwana paleogeography from assembly to 33. McMahan CD, Chakrabarty P, Sparks JS, Smith WL, Davis MP. 2013 breakup — a 500m.y. odyssey. Geol. Soc. Am. Spec. Pap. 441, 1-28. Temporal patterns of diversification across global cichlid biodiversity layers [40]. This more appropriate minimum distribution across southern landmasses two groups of terrestrial mammals — pri- ies reporting close congruence between (doi:10.1130/2008. 2441(01)) (Acanthomorpha: Cichlidae). PLoS ONE 8, e71162. (doi:10.1371/journal. 8. Stiassny MLJ. 1991 Phylogenetic intrarelationships of the family Cichlidae: pone.0071162) age estimate only partially reconciles our combined with a fossil record that com- mates and hystricognath rodents — dis- divergence time estimates from ‘rocks’ and an overview. In Cichlid fishes: behaviour, ecology, and evolution (ed. MHA 34. Brandley MC, Wang Y, Guo X, Nieto Montes de Oca A, Feria-Ortiz M, Keenleyside), pp. 1-35. London, UK: Chapman & Hall. Hikida T, Ota H. 2011 Accommodating heterogenous rates of evolution in time scale with previous phylogenetic inter- mences long after the tectonic break-up of persed from Africa to South America at ‘clocks’, in cases where these approaches 9. Chakrabarty P. 2004 Cichlid biogeography: comment and review. Fish Fish. molecular divergence dating methods: an example using intercontinental 5, 97-119. (doi:10.1111/j. 1467-2979.2004.00148.x) dispersal of Plestiodon (Eumeces) lizards. Syst. Biol. 60, 3-15. pretations of the Lumbrera cichlids, each Gondwana. Several groups of freshwater approximately this time [53]. More gener- had previously delivered wildly different 10. Sparks JS, Smith WL. 2004 Phylogeny and biogeography of cichlid fishes (doi:10.1093/sysbio/syq045) (Teleostei: Perciformes: Cichlidae). Cladistics 20, 501-517. (doi:10.1111/ 35. Phillips MJ. 2009 Branch-length estimation bias misleads molecular of which has been placed within the South fishes, reptiles, mammals and plants show ally, there is strong evidence from other evolutionary time scales [57]. This conver- j.1096-0031.2004.00038.x) dating for a vertebrate mitochondrial phylogeny. Gene 441, 132-140. gence would seem to signal the end of an 11. Jokat W, Boebel T, Konig M, Meyer U. 2003 Timing and geometry (doi:10.1016/j.gene.2008.08.017) American crown in association with specific disjunct distributions, with members pre- groups and plants for surprisingly of early Gondwana breakup. J. Geophys. Res. 108, 2428. 36. Zheng Y, Peng R, Kuro-O M, Zeng X. 2011 Exploring patterns and extent of bias in estimating divergence time from mitochondrial DNA sequence † era dominated by debates on the relative (doi:10.1029/2002JB001802) cichline tribes ( Protocara as either a geo- sent in South America and Africa, but only high levels of biotic interchange between 12. Murray AM. 2001 The fossil record and biogeography of the Cichlidae data in a particular lineage: a case study of salamanders (Order Caudata). merits of molecular and fossil data, per- (: Labroidei). Biol. J. Linn. Soc. 74, 517-532. Mol. Biol. Evol. 28, 2521-2535. (doi:10.1093/molbev/msr072) phagine or a stem member of an unnamed a few instances seem definitively attribut- South America and Africa throughout the 13. Perez PA, Malabarba MC, del Papa C. 2010 A new genus and species 37. Miya M et al. 2010 Evolutionary history of anglerfishes (Teleostei: mitting molecular biologists and palaeon- of (Perciformes: Cichlidae) from the early Eocene of southern Lophiiformes): a mitogenomic perspective. BMC Evol. Biol. 10, 58. clade comprising , Geo- able to driftbased vicariance [14,43,44]. Late Cretaceous and Palaeogene [54,55]. South America. Neotrop. Ichthyol. 8, 631-642. (doi:10.1590/S1679- (doi:10.1186/1471-2148-10-58) tologists to move forward on addressing 62252010000300008) 38. Malabarba MC, Malabarba LR, Del Papa C. 2010 phagini, and Heroini [39,41]; Instead, molecular clock analyses for a Geological evidence indicates the pres- 14. Lundberg JG. 1993 African-South American freshwater fish clades and eocenicus, n. sp. (Perciformes: Cichlidae), an Eocene cichlid from †Gymnogeophagus eocenicus as phylo- range of groups with apparent vicariant ence of a chain of now-submerged islands questions related to the timing of major continental drift: problems with a paradigm. In Biological relationships the Lumbrera Formation in Argentina. J. Vert. Paleontol. 30, 341-350. between Africa and South America (ed. P Goldblatt), pp. 156-199. New (doi:10.1080/02724631003618348) genetically nested within a living genus distributions across southern continents across the south Atlantic during the events underpinning the origin of modern Haven, CT: Yale University Press. 39. Malabarba MC, Zuleta O, del Papa C. 2006 Proterocara argentina, a new 15. Unmack PJ. 2001 Biogeography of Australian freshwater fishes. J. fossil cichlid from the Lumbrera Formation, Eocene of Argentina. J. Vert. [38]; and †Plesioheros as a crown heroine [45-48] paint a picture of widespread Palaeogene [52]. These islands coincided biodiversity. Biogeogr. 28, 1053-1089. (doi:10.1046/j.1365-2699.2001.00615.x) Paleontol. 26, 267-275. (doi:10.1671/0272-4634(2006) 26 [267:PAANFC] 16. Sparks JS, Smith WL. 2005 Freshwater fishes, dispersal ability, and 2.0.CO;2) [13]). It is difficult to evaluate confidence ‘pseudo-congruence’, where similar bio- with strong east-to-west palaeocurrents Acknowledgements: nonevidence: ‘Gondwana Life Rafts’ to the rescue. Syst. Biol. 54, 158-165. 40. del Papa C, Kirschbaum A, Powell J, Brod A, Hongn F, Pimentel M. 2010 We thank J Friel of the Cornell University Museum of (doi:10.1080/10635150590906019) Sedimentological, geochemical and paleontological insights applied to in the evolutionary relationships proposed geographic patterns originate at different across the south Atlantic and both have 17. Marshall CR. 1997 Confidence intervals on stratigraphic ranges with continental omission surfaces: a new approach for reconstructing an Vertebrates, USA, and AC Bentley and E O Wiley of nonrandom distributions of fossil horizons. Paleobiology 23, 165-173. Eocene foreland basin in NW Argentina. J. S. Am. Earth Sci. 29, 327-345. for these fossils because published anal- times that may be disjunct with the age been invoked as key elements of a selec- the Biodiversity Institute of the University of Kansas, 18. Hedman MM. 2010 Constraints on clade ages from fossil outgroups. (doi:10.1016/J.Jsames.2009.06.004) USA for generous gifts of tissue specimens. G Paleobiology 36, 16-31. (doi:10. 1666/0094-8373-36.1.16) 41. Smith WL, Chakrabarty P, Sparks JS. 2008 Phylogeny, , and yses using morphological data do not of specific palaeogeographic events [49]. tive dispersal route from Africa to South 19. Wainwright PC, Smith WL, Price SA, Tang KL, Sparks JS, Ferry LA, Kuhn KL, evolution of Neotropical cichlids (Teleostei: Cichlidae: Cichlinae). Watkins-Colwell assisted with museum collections. Eytan RI, Near TJ. 2012 The evolution of pharyngognathy: a phylogenetic Cladistics 24, 625-641. (doi:10.1111/j.1096-0031.2008. 00210.x) provide support for nodes in accompany- Our consistent time scales for cichlid evo- America during the Eocene [12,52]. It is G Lloyd and D Bellwood commented on previous and functional appraisal of the pharyngeal jaw key innovation in Labroidei 42. Ru¨ber L, Adams DC. 2001 Evolutionary convergence of body shape and

12 facebook.cichlids.org.au cichlidpower.org.au 13 trophic morphology in cichlids from Lake Tanganyika. J. Evol. Biol. 14, 325-332. (doi:10.1046/J.1420-9101.2001.00269.X) 43. de Queiroz A. 2005 The resurrection of oceanic dispersal in historical biogeography. Trends Ecol. Evol. 20, 68-73. (doi:10.1016/j. tree.2004.11.006) 44. Beaulieu JM, Tank DC, Donoghue MJ. 2013 A Southern Hemisphere origin th for campanulid angiosperms, with traces of the break-up of Gondwana. One day only ... Saturday March 7 2015 Glenn Briggs needs no introducƟon to Australian fishkeepers, BMC Evol. Biol. 13, 80. (doi:10.1186/1471-2148-13-80) 45. Gamble T, Bauer AM, Colli GR, Greenbaum E, Jackman TR, Vitt LJ, Simons starƟng fishkeeping in 1975 with a typical tropical community set up. AM. 2011 Coming to America: multiple origins of New World geckos. J. presented by:- Cichlid Press Australia and His fishkeeping hobby grew in size and to assist in paying for fish Cichlid Press Australia Evol. Biol. 24, 231-244. (doi:10.1111/j.1420-9101.2010.02184.x) Glenn took a job at the local aquarium store. This soon led to full Ɵme 46. Rowe DL, Dunn KA, Adkins RM, Honeycutt RL. 2010 Molecular clocks keep The Eastern Districts Aquarium Society The Eastern Districts Aquarium Society dispersal hypotheses afloat: evidence for trans-Atlantic rafting by rodents. work with several aquarium retailers before he changed direcƟon and J. Biogeogr. 37, 305-324. (doi:10.1111/j.1365-2699.2009.02190.x) started work in the finance industry. 47. Cook LG, Crisp MD. 2005 Not so ancient: the extant crown group of (as part of their 60th. Birthday celebrations) SPECIAL EVENT Nothofagus represents a post-Gondwanan radiation. Proc. R. Soc. B 272, AŌer being married in 1986, Glenn greatly expanded his fish keeping, 2535-2544. (doi:10.1098/rspb.2005.3219) building a fish room of 50 aquaria, housing Cichlids, naƟves and killifish. Garden ponds 48. Knapp M, Stockler K, Havell D, Delsuc F, Sebastiani F, Lockhart PJ. 2005 housed local naƟve fishes. His wife , Jenny soon realized there was no way to avoid fish Relaxed molecular clock provides evidence for long-distance dispersal of Nothofagus (Southern Beech). PLoS Biol. 3, 38-43. (doi:10.1371/Journal. food in the freezer or blackworms in the fridge so she bought him his own fridge ! In International Pbio.0030014) 1998 Glenn returned to the aquarium trade working for many years at Aquarium Indus- 49. Donoghue MJ, Moore BR. 2003 Toward an integrative historical biogeography. Integr. Comp. Biol. 43, 261-270. (doi:10.1093/icb/43.2.261) tries with Rick Datodi. At present he works part Ɵme at A.I. He sƟll maintains a large 50. Suresh AV, Lin CK. 1992 Tilapia culture in saline waters: a review. number of aquariums and ponds keeping especially Purple SpoƩed Gudgeons (Mogurnda Aquaculture 106, 201-226. (doi:10.1016/0044-8486(92)90253-H) spp), Rainbows and Pygmy Perch. A foundaƟon member of ANGFA NaƟonal, Glenn has Cichlid Conference 51. Uchida K, Kaneko T, Miyazaki H, Hasegawa S, Hirano T. 2000 Excellent salinity tolerance of Mozambique tilapia (Oreochromis mossambicus): enjoyed parƟcipaƟon in the organisaƟon for many years, he was made a life member in elevated chloride cell activity in the branchial and opercular epithelia 2007. 14th March 2015, in Brisbane of the fish adapted to concentrated seawater. Zool. Sci. 17, 149-160. (doi:10.2108/Zsj.17.149) Featuring: 52. de Oliveira FB, Molina EC, Marroig G. 2009 Paleogeography of the South Atlantic: a route for primates and rodents into the New World? In South Costs: Non-members $55.00 per person Ad Konings - Juan Miguel Artigas Azas - Spencer Jack American primates (eds PA Garber, A Estrada, JC Bicca-Marques, EW Heymann, KB Strier), pp. 55-68. New York, NY: Springer. 53. Loss-Oliveira L, Aguiar BO, Schrago CG. 2012 Testing synchrony Members EDAS/A S of V $40.00 per person in historical biogeography: the case of New World primates and Hystricognathi rodents. Evol. Bioinform. 8, 127-137. (doi:10.4137/Ebo. S9008) 54. Ezcurra MD, Agnolin FL. 2012 A new global palaeobiogeographical BOOKINGS ARE ESSENTIAL model for the Late Mesozoic and Early Tertiary. Syst. Biol. 61, 553-566. (doi:10.1093/sysbio/syr115) 55. Morley RJ. 2000 Origin and evolution of tropical rain forests, p. 362. Chichester, NY: Wiley. Enquiries and Bookings: 56. Measey GJ, Vences M, Drewes RC, Chiari Y, Melo M, Bourles B. 2007 Freshwater paths across the ocean: molecular phylogeny of the frog Email: [email protected] Ptychadena newtoni gives insights into amphibian colonization of oceanic Renowned internationally known aquarists islands. J. Biogeogr. 34, 7-20. (doi:10. 1111/J.1365-2699.2006.01589.X) at the Tavernetta Function Centre 57. Erwin DH, Laflamme M, Tweedt SM, Sperling EA, Pisani D, Peterson KJ. Phone Daryl : 9874 1850 0438 799933 2011 The Cambrian conundrum: early divergence and later ecological 144 Dorvile Road, Carseldine QLD success in the early history of . Science 334, 1091-1097. Ad Konings and Or Phone John: 5429 1516 0407 513 549 (doi:10.1126/science. 1206375)  Bank details : BSB 633-000 A/C No. 124299744 Full Day Conference with Morning Tea, Afternoon Juan Miguel Artigas Azas Tea, Lunch and Evening Meal. Thanks The generosity of the following organisaƟons is gratefully acknowledged Also includes Trade Room, Tea and for their support in making this event possible: Coffee throughout the day, Conference also our own fish guru ...Glenn Briggs The Eastern Districts Aquarium Society. The Aquarium Society of Victoria Shirt and Lucky Door Prizes. Cichlid Press Australia Cristys Spit Roasts Cranbourne Aquariums Club Members $130 and $150 for non-members. Aqua Pics Subscape Aquariums Coburg Aquariums Aquarium Industries A.S. of V. Hall … 29 Grant Street Clifton Hill A.S. of V. Hall … 29 Grant Street Clifton Hill Co-Ordinator: Sue Broadbent 0409639603 or [email protected] Saturday Mar. 7th 2015 ~ 9.30 am to 6.30 pm A.S. of V. Hall : Melway Ref: 2 C K 2 Postcode: 3068 Saturday Mar. 7th 2015 ~ 9.30 am to 6.30 pm

14 facebook.cichlids.org.au cichlidpower.org.au 15 It was distributed to over 300 stores and ceased publication in December 1975, a run of Volume II Number 2 eight years and seven months THE AQUARIST’S NOTEBOOK is free! It is for a total of 103 issues. available, just for the asking, at your local pet shop or fish store. It is your dealer’s way of saying “Thank you!” for your patronage. Written over 30 years ago, Because THE AQUARIST’S NOTEBOOK is published at regular intervals, and because It is some of the scientific referenc- Another little bit of a permanent reference work chock full of valuable ideas and fascinating articles, you es may be outdated. However, history for you: will want to ensure that you do not miss a single issue. Keep your reference library com- the material was aimed at plete… visit your dealer regularly for your free copies! beginners and the informa- In the late spring of 1967 tion and recommendations are Albert J Klee conceived the as valid today as they were idea of a brand new month- then. ly publication, unique to the Two of the cichlid-related aquarium world. Physically, it editions are recreated here consisted of a single sheet of with the permission of Dr Klee, heavy stock, printed on both a very helpful man who is still sides, with spot-colour on the imparting his knowledge with first page and punched so that a refreshingly correct turn of it could be saved in a loose- phrase that is so rare these leaf binder. No photographs days. were used, but there was an abundance of black and white More examples of this inter- illustrations. esting publication will be The publication was named reproduced in future editions The Aquarist’s Notebook and of iCichlid, presuming there is was sold in bulk to pet shops other copy to fill-out the rest of and fish stores and distrib- the pages. uted free to their customers. ANSWERS TO QUICKIE QUIZ The idea being that it would Barbus everetti - clown barb QUICKIE QUIZ Brachydanio rerio - zebra danio encourage return business for On the left are the partial scientific names of maronii - Barbus tetrazona - tiger barb the shops and also provide certain fishes; on the right are their partial MORE popular names. Match them up. Gymnotus carapo - knife fish ➔ useful and interesting material 1. everetti a. zebra 2. rerio b. knife THE AQUARIST’S NOTEBOOK to their customers. 3. maronii c. tiger Vol. 2 No. 2 4. tetrazona d. clown © 1968 by 5. carapo e. keyhole The Aquarist’s Notebook Space was provided at P.O. Box 139 Fairfield, Ohio 45014 the bottom of the first page (Answers on next page) for the dealer to stamp the THIS COPY OF THE AQUARIST'S NOTEBOOK name, address, and telephone COMPLIMENTS OF: number of the establishment distributing the publication.

16 facebook.cichlids.org.au cichlidpower.org.au 17 THE AQUARIST’S NOTEBOOK is free! It is available, just for the asking, at your local pet shop or fish store. It is your dealer’s way of saying “Thank you!” for your patronage. Volume I Number 4 Because THE AQUARIST’S NOTEBOOK is published at regular intervals, and because It is a permanent reference work chock full of valuable ideas and fascinating articles, you will want to ensure that you do not miss a single issue. Keep your reference library com- plete… visit your dealer regularly for your free copies!

QUICKIE QUIZ 1. Cichlids have spines in their dorsal fins. (a) True, (b) False 2. The primary purpose of aeration is to force oxygen into the water via the air bubbles. (a) True, (b) False 3. Catfishes are more closely related to killifishes than they are to rasboras. (a) True, (b) False 4. The swordtail comes from Mexico. (a) True, (b) False ANSWERS TO QUICKIE QUIZ THE AQUARIST’S NOTEBOOK 1. True Vol. 1 No. 4? (Answers on next page) 2. False (to provide circulation which © 1967 by The Aquarist’s Notebook assists in oxygenation at the surface) P.O. Box 139 3. False 4. True THIS COPY OF THE AQUARIST'S NOTEBOOK Fairfield, Ohio 45014 COMPLIMENTS OF:

18 facebook.cichlids.org.au cichlidpower.org.au 19 From Cichlidae communique #206 – Reprinted from Cichlidae communiqué, publication of the Pacifi c Coast Cichlid Association. Interested parties are invited to write: PCCA, Membership Chairman, PO Box 28145, San Jose, CA 95159-8145 for more information. my favourite fish store, I Kite!” Kite is the name for the large Green cal filtration materials from the filters. Adventures In Failure: brought the breeding col- Severum; he’s really placid and just drifts Within three days, I saw that almost all of ony to an end by bringing around the tank like a big kite, hence the the fish were erupting with the parasites those females and juveniles name. I asked what “wrong” meant. She and the tell-tale signs on Kite were becom- to the shop. Yes, we’re said that his whole body looks like some- ing even worse. He looked like he was Benign Neglect talking dozens of fish. In one poured salt all over him. coated in a layer of plaster that was crack- the process, I noticed that Uh Oh!!! I know what that means. We ing away, and his fins were decaying at an vs Over-ReactingOver-Reacting they had some new arriv- have an “ich” infestation. alarming rate. als, in particular several Now, I’ve seen these before, and I’ve With that, I knew it was only a matter of Text and Photos by Michael Larsen Green Terrors (Aequidens treated them in the past, so I figured, well, time. Some of the smaller convicts were rivulatus). Contrary to this shouldn’t be too big a deal. Since I’d the first to go on Wednesday, then some This is a blog post I hoped I would never without any jokes or colour commentary. popular folklore, the needed to get medication anyway, I figured of the larger Convicts followed suit. Kite, name is a bit of a mis- have to write. To my tester friends, this Yesterday, my 260 litre fish tank became a I’d pick it up the next day and start treat- my oldest and longest-lived veteran of my nomer. They are cichlids, has only peripheral relation to my testing ghost town. ment when I got home. For good measure, tank, at 12 years, succumbed to the dis- so yes, when they are in career, but it’s something I’ve been deal- This tank has been up and running, in I’d do another large water change so that ease on Friday morning. Over the next “breeding mode”, they can ing with for the past several days, and it’s some way, shape and form, continuously, I could limit the spread of the problem. three days, roughly every 12 hours, another The female Convict guarding over the fry. be as aggressive as any taught me a few things, and reminded me with one major exception (for a swap-out Unfortunately, my hospital tank was not set fish would die until finally I was left with other cichlid species, but once again of several things I thought I back in 2007 due to a ruptured seam) for up. Even if it was, I’d need a larger tank to only three fish. in the way of tweaking and meddling on my no more so. As I was looking at them already knew. Normally, I take a somewhat 19 years. In that 19-year period, many fish put the Green Severum in; a 22 litre quar- Looking at the two surviving Green Terrors part, for years. That all changed Monday I thought “wow, what a great time to self-deprecating tone with posts like this, have come and gone, for a variety of rea- antine tank isn’t going to cut it for a 25 cm (two had likewise died during the week), the 11th of November, 2013. balance out the tank with a new species”. but I’m really and seriously bummed about sons (personal taste and interest, changes With Acara (Green Terrors being part of fish! Therefore, it meant I’d have to treat the and my longest-lived male Convict, I took this, so I’m just going to tell it straight in water chemistry moving from soft, acidic Wait, let me step back another few days, this family of cichlids) and Convicts, as whole tank. a look at the two Green Terrors and realised San Francisco to to Saturday, November 2, 2013. That day, well as my main tank denizen, a Green Turns out that hard, alkaline San I did something momentous, and poten- Severum (Heros efasciatus), coming from would have to be tially provided the catalyst that started this Bruno, etc). similar waterways in Central and South the main step any- whole thing. On that day, I made a deci- Generations of America, I figured they’d be a great addi- way, since when I sion to end a decade-long experiment. I’d fish have been born, tion to my tank. got home, I noticed kept a breeding colony of Convict Cichlids grown, given away, I bought four of them, introduced them that the signs of the ( nigrofasciatus) running in and repopulated to into my tank, and then spent the next cou- disease had spread that tank for that time, and it was success- other tanks. It was ple of days getting ready to go to Sweden. to several of my ful. In fact, it was too successful. I had run a vibrant communi- Since I was going to be gone for a while, Convicts as well. ty of predominantly out of tanks to house them, and shops I figured giving the tank a good thorough With that, I started cichlids, though it willing to take them (even for free; I was cleaning and a larger-than-average water measuring out the has housed other flooding the local market with Convicts). change would be good for all involved, medicine and dos- fish over the years With no more room to put them, I decided as well as a perfect time to introduce the ing the entire tank. as well. To put it it was time to make a change. I kept new fish. Now we fast-forward to Monday I also followed simply, it’s run pret- the largest and longest-lived males, sepa- evening, November 11th. As I came home the directions and ty much flawlessly, rated out all of the females and juve- from work that day, my younger daughter removed all of the The tank before the crash. and without much niles, and with a final “special delivery” to said “Dad, there’s something wrong with carbon and chemi- Heros efasciatus before the crash.

20 facebook.cichlids.org.au cichlidpower.org.au 21 when I checked the water late last week, one of the ways I observe and learn about the pH was 7.8 (and yes, that difference the semi-natural world. AAUSTRALIANU is significant). That large a pH swing could STRALIAN Often in tanks, there are two things that & NNEWEW GUINEA have triggered the change. Could the medi- GUINEA are killers; overt neglect and over meddling. FFISHESISHES AASSOCATIONSS cation have exacerbated the problem? OCATION The middle ground is sometimes referred to Entirely possible. One of the dangers of MMeetseets bi-monthly at the bi-monthly a FFieldiel Naturalist Club,t th e as “benign neglect”, where we do the bare d Naturalist C medicating a large tank is that the dosing 1 GGardeniaard Street, Blackburn.lub, enia Street, is very hard to determine. What would be minimum necessary, and let them sort it out Blackburn. CContact:ontact: angfavic.org angfavic.org normal for one fish might be too little for for themselves. I’ve never been guilty (not another, or way too much for a third. This that I know of at least) of “overt neglect”, is why it’s best to treat fish individually in a but yes, I’ve frequently been on the “benign hospital tank if you can. Treating a whole neglect” scale. Sometimes, that’s just the tank can have wildly varying results. It’s NNATIONALATIONAL best way to deal with things, but it makes also possible that the dosing of the tank AUSTRALIAN KILLIFISH us feel heartless when we do it. It seems IATION was too late for Kite, who already showed AASSOCIATIONSSOC this would have been a time where more advanced symptoms. nthly Male Convict. Harlock, the Green Terror, after the crash. “benign neglect” would have been far more MMeetseets BBi-monthlyi-mo homes. Could I have done anything different? iinn mmembers’embers’ homes. beneficial. As it stands, I now have a recov- CCall:all: Sure, and the last option is the one that 3408. Jenkin 9442 that, if anyone were likely to survive, they what may well be a colony of parasites that added them to the tank. What’s the worst really makes me cringe, but I know the ery project underway. I will rebuild, and I EEmmamma Jenkin 9442 3408. had the best chance. With that, I set up I will now wait out the next three weeks, to that could happen? I really hadn’t imag- truth of it, and didn’t heed it. I could have will renew this tank. New life will take root the 25 litre hospital tank, and pulled them make sure that they are all dead before I try ined “the worst” would mean “wipe out my left them alone. I could have resisted the here again, but sadly, it will be with a whole out to be monitored and treated. My oldest to start the tank up again. entire environment”, but yes, that is exactly urge to add some new fish to the tank after new family. All my “best friends” are gone, EEASTERNASTER DISTRICTS Convict, I had to leave him in the big tank, Many things have been going through what happened. Was it just the introduc- a major “depopulation”. I could have not and they are gone because I over-reacted. N DISTRICTS AAQUARIUMQUA and hope for the best. Alas, the end came my head since this all happened. What did tion of the fish? If that’s the case, why were bothered with the water change before RIUM Update: I’m happy to report that both SSOCIETYOC for him too, last night. they not sick? Why did none of them show leaving for Sweden. I could have let the IETY I do wrong? The answers are telling. Most of the Green Terrors seem to be doing disease just run its course. Yes it would MMeetseets ono the 4th Friday of the month at As of now, there are no fish of any kind critically, I broke one of my most important any signs of the disease? Could there have n the 4th Frid OK, even the one that’s missing half the NNunawadingunaw Civic Centre,ay of tWhitehorsehe mo have likely killed Kite, but he was 12 years ading Civic nth at been another potential cause? Yes. Pretty Centre, Whit in my main tank. They are all dead. As of rules, and I did it out of impatience. My RRoad,oad, Nunawading.N ehorse old, already beyond his life expectancy, scales on the left side of his head. He’s unawading. much all fish carry parasites. It comes with PPOO BoxBo 3005, Nunawading 3131 this morning, my two Green Terrors are hospital/quarantine tank wasn’t set up. It x 3005, Nuna and having had a really great run. All sorts being feisty, sparring with the other Green wading 3131 still holding on, one looking like it might hadn’t needed to be for some time. I was the territory. “Ich” is expressed when fish of coulda’, shoulda’ woulda’s, but no, the Terror, and thankfully, he’s even eating, be a hard recovery (he’s lost a fair amount not prepared, and I didn’t have the neces- have a stress episode, and those parasites thought of doing nothing terrified me. I did which means he really does have a fighting of scale over the left side of his head and sary materials in place. Cautious, intelligent are excited and activated. They then push what any irrational pet owner would do me would have set up that tank first, and out of the fish to reproduce and look for chance. I tend to name the fish that stand flank), and the other having what looks like when their animals are in distress. I tried to AAQUARIUMQUARIUM would have let it cycle for three weeks, new hosts. Could the two large-scale water out in my tanks, and thus, if this little guy a real fighting chance. My 25 litre tank is fix it with all the tools at my disposal. The SSOCIETYOCIETY then bought new fish, and had them wait in changes in less than two weeks have con- pulls through, since it’s likely he’ll have a IA not much, it’s definitely not the environs I net result is a ghost town. Over a dozen OOFF VICTORIAVICTOR just pulled them out of, but it will have to the quarantine tank for three weeks, before tributed to that? Well, yes, potentially. bit of scarring that will look like a Pirate’s valuable fish, but more to the point, fish nth ay of the mo be home for the next three weeks, and on introducing them to my main tank. The fact is, water pH is probably the one eye patch, I’m going to call this little fighter e last Thursd that were good and dear friends I’d raised MMeetseets onon theth last Thursdayli fofto nthe H ilmonthl. rant Street, C the plus side, they are currently still alive. Instead, because of a moment where I thing, outside of ammonia or nitrite spikes, “Harlock” after Leiji Matsumoto’s legend- aatt 2929 GrantG Street, Clifton Hill. for many years. Why am I mentioning all CCall:all: 0. ary space pirate. Here’s hoping I can make ck: 9874 185 For the first time in 19 years, though, my was making a major change in the ecology that will most stress/affect a fish. I try to this? Simple, this is my blog. This is one of DDarylaryl Maddock:Maddo 9874 1850. main tank is now devoid of life, except for of the tank, I figured “why not?”, and I just keep the pH as close to 7 as possible, but my biggest hobbies outside of testing, and good on that. j

22 facebook.cichlids.org.au cichlidpower.org.au 23 ::

The Cichlids are Coming!” the The Cichlids are Coming!” last Coming this summer to wordBy DDarylaryl HutchinsHutchins he fi rst Last Word for 2015! Hopefully Where Cichlids are Revered Tnot another Annus Horribilis as far as new content for this, your journal is concerned! I am hoping for an Annus Miribilis instead. That will require that you do something The Sheraton Hotel - Springfield, Massachusetts to help keep the covers of this journal July 30 — August 2, 2015 from coming into contact with one another. As it is electronic, that could be Hosted by catastrophic.

or the fi rst time in many, many years, I Fdid not produce an entry in the Elaine Vendor Room Show Competition Turner Memorial Art & Photographic Competition in December 2014 ... neither Hospitality Suite Kids’ Tank Decorating did anyone else. Boston Day Trip Old Sturbridge Village Banquet Giant Sunday Cichlid Auction While it is fairly typical of what was a year best forgotten, I fi nd it extremely Speakers: disappointing. Let’s see if we can turn Wayne Leibel Laif DeMason Oliver Lucanus Ad Konings that around in 2015 ... you have 11 months Dr. Paul Loiselle Rusty Wessel Al Sabetta Jaap-Jan de Greef to produce something, it’s not like there’s Charley Grimes Dr. Hubert Kuerzinger a rush or anything.

To register, reserve a room, become a sponsor, or volunteer to help, go to: any, many paragraphs have given M their lives in vain here, trying ACAconvention2015.com to encourage/goad/embarass/coax/ implore/inspire/spur/energise/push/stir Proudly sponsored by or sway you, yes you, into submitting content of some sort to this your, yes your, magazine.

Those words can’t just continue being :: ignored ad infi nitum. Eventually, the bucket will not return from the well. y IMAGE: Alf Stalsberg 24 facebook.cichlids.org.au : D 4 9 7 2 1 0 0 A A0012794D

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