Theraps intermedius © Lee Nuttall

An extensive review of the herichthyin clade of Middle American cichlids has resulted in the naming of eight new genera and a number of taxonomic modifications.

The herichthyins are a group of 45 cichlid species contained within the larger Heroni tribe of the Cichlinae subfamily. They are mostly distributed in Mexico and Guatemala and contain a number of aquarium favourites. Generic placement for a number of them has long been considered uncertain, however, with conflicting taxonomic designations having been proposed in a number of cases. In others, species have remained in the catch-all genus Cichlasoma or the placeholder ‘Cichlasoma’ incertae sedis.

In the new study, published last week in the journal ‘Zootaxa’, ichthyologist Caleb McMahan and collaborators compiled DNA sequence data and evaluated morphological characters from various earlier studies. The results were analysed alongside physical examination of specimens in order to support hypotheses and potentially obtain new diagnostic characters.

As a results, existing herichthyin groupings were rediagnosed, and new genera proposed based on genetic and morphological differentiation.

The herichthyins thus now comprise the following genera. Please note that only the principal defining characters for each grouping are included here, and that in all cases the first species listed is the type species:

Vieja maculicauda, type species of the genus. © Štefan Zelinka

Vieja Fernández-Yépez 1969 includes the type species V. maculicauda plus V. bifasciata, V. breidohri, V. fenestrata, V. guttulata, V. hartwegi, V. melanura, and V. zonata. These species are distinguished from most other herichthyins by possession of a large blotch covering the majority of the caudal peduncle, which extends anteriorly as a dark stripe in all members except V. maculicauda. They have all recently been included in the genus Paraneetroplus.

Maskaheros argenteus ©Unimati.dk

Maskaheros McMahan and Piller 2015 is a new genus containing M. argenteus and M. regani, which have previously been included in both Vieja and Paraneetroplus. The primary diagnostic characters are the presence of small, dark, irregular spots on the entire body and two broad dark interorbital bars.

Paraneetroplus gibbiceps © Lee Nuttall

Paraneetroplus Regan 1905 is now restricted to contain only P. bulleri, P. gibbiceps, and P. nebuliferus. These taxa differ from other herichthyins by the elongate body and caudal peduncle, and narrow, sub-terminal mouth.

Herichthys cyanoguttatus © Štefan Zelinka

A reduced Herichthys Baird and Girard 1954 includes H. cyanoguttatus, H. carpintis, H. deppii, H. minckleyi, H. tamasopoensis, H. tepehua, H. teporatus. They are diagnosed by possession of 6-7 vertical body bars which are often grouped in the posterior portion of the body, and a series of dark blotches below the upper lateral line.

Nosferatu molango © Mauricio De la Maza Benignos

Nosferatu De la Maza-Benignos et al. 2015 was raised in a study published earlier this year. It contains N. pame, N. bartoni, N. labridens, N. molango, N. pantostictus, N. pratinus, and N. steindachneri, all former members of Herichthys. They are diagnosed by possessing a relatively elongate body with variable, irregular stripes or blotches laterally, and conical unicuspid teeth in the anteriormost portion of the upper jaw.

Theraps godmanni © Lee Nuttall

Theraps Günther 1862 has also been reorganised and now comprises T. irregularis, T. godmanni, T. intermedius, T. microphthalmus, and T. nourissati. These are distinguished from other herichthyins by having bodies that are longer than deep, a small subterminal mouth, an upper jaw that extends beyond the lower, and spots on the unpaired fins.

Kihnichthys ufermanni © Mo Devlin

Kihnichthys McMahan and Matamoros 2015 is the second new genus raised in the paper and is monotypic, containing only K. ufermanni, a former member of both Vieja and Theraps. It is chiefly distinguished by presence of a dark blotch on the centre of the caudal peduncle base, and spatulate or chisel-like teeth in the anterior portions of both jaws.

Cincelichthys bocourti © Unimati.dk

The third new genus Cincelichthys McMahan and Piller 2015 includes C. bocourti and C. pearsei, both of which have most recently been considered members of Theraps. They are diagnosed by possessing spatulate, chisel-shaped teeth plus very deep, rounded to oval-shaped bodies with wide, irregular bars formed by black scales, and a small black spot in the centre of the caudal peduncle base.

A young individual of Rheoheros lentiginosus © Florent De Gasperis

Rheoheros McMahan and Matamoros 2015 is another new taxon, and contains R. lentiginosus and R. coeruleus, both formerly placed in Theraps. It is is principally diagnosed by a unique colour pattern of dark blotches and/or bars on an elongate body.

Oscura heterospila © Unimati.dk

Oscura McMahan and Chakrabarty 2015 is a new monotypic genus raised for the species formerly referred to as Theraps heterospilus, which now becomes O. heterospila. It is diagnosed by the combination of a round or oblong-shaped dark blotch occupying most or all of the caudal peduncle, plus small black spots the sides of the body.

Chiapaheros grammodes © Štefan Zelinka

Chiapaheros McMahan and Piller 2015 is another new, monotypic taxon containing C. grammodes. It is distinguished by presence of several, typically 7, thin dark lines on the sides of the head, plus an elongated predorsal profile and head.

Thorichthys ellioti, type species of the genus, is currently regarded as a synonym of T. maculipinnis. © Robert Smith

Thorichthys Meek 1904 consists of T. ellioti, T. affinis, T. aureus, T. callolepis, T. helleri, T. meeki, T. pasionis, T. socolofi. The authors note that T. ellioti, the type species, is currently regarded as a synonym of T. maculipinnis, and that further work is required to assess its taxonomic validity. Members of this genus are readily distinguished from all other herichthyins by possession five (versus four) mandibular pores along the dentary. They are also relatively small in overall size, have an elongate angular head profile, and a small terminal mouth.

Mesoheros festae © Enrico Richter

New genus Mesoheros McMahan and Chakrabarty 2015 contains the South American species M. festae, M. atromaculatus, and M. ornatus, all previously assigned to Cichlasoma. They are diagnosed by possessing an elongate body with 7 (rarely 6) dark spots or bars laterally, and a moderately small mouth that does not reach the anterior margin of the orbit.

Tomocichla tuba © Lee Nuttall

Tomocichla Regan 1908 now contains only T. tuba and T. asfraci. They are distinguished by the combination of compressed conical teeth in the anterior upper jaw, and dorsal and anal fins with markedly rounded posterior margins.

Herotilapia multispinosa © Unimati.dk

The monotypic Herotilapia Pellegrin 1904 is revalidated and contains H. multispinosa, unique among herichthyins in possessing tricuspid teeth.

Trichromis salvini © Ryan O’Donnell

Trichromis McMahan and Chakrabarty 2015 is another new, monotypic taxon and contains T. salvini, formerly Cichlasoma salvini. It is easily distinguished from other herichthyins by possession of two dark lateral stripes in the upper portion of the body.

‘Cichlasoma’ sieboldi © Unimati.dk

‘C.’ tuyrense and ‘C.’ sieboldii are retained in ‘Cichlasoma’ incertae sedis since they are not closely related to the herichthyins despite former generic placements in Vieja and Tomocichla, respectively.

Matt Ford

For further information see the full paper (link to abstract only): McMahan, C. D., W. A. Matamoros, K. R. Piller and P. Chakrabarty, 2015. Taxonomy and systematics of the herichthyins (Cichlidae: Tribe Heroini), with the description of eight new Middle American Genera. Zootaxa 3999(2): 211-234.

Additional references:

López-Fernández, H., K. O. Winemiller and R. L. Honeycutt, 2010. Multilocus phylogeny and rapid radiations in Neotropical cichlid fishes
(Perciformes: Cichlidae: Cichilinae). Molecular Phylogenetics and Evolution 55(3): 1070-1086.

McMahan, C. D., A. D. Geheber and K. R. Piller, 2010. Molecular systematics of the enigmatic Middle American genus Vieja (Teleostei: Cichlidae). Molecular Phylogenetics and Evolution 57(3): 1293-1300.

De la Maza-Benignos, M., C. P. Ornelas-García, M. de Lourdes Lozano-Vilano, M. E. García-Ramírez and I. Doadrio, 2015. Phylogeographic analysis of genus Herichthys (Perciformes: Cichlidae), with descriptions of Nosferatu new genus and H. tepehua n. sp. Hydrobiologia 748(1): 201-231.

Thanks to Lee Nuttall, Mo Devlin, Mauricio De la Maza Benignos and regular contributors Enrico, Štefan, Ryan, Robert, and Unimati.dk for use of images.

Betta hendra is a member of the B. coccina species group within the genus. © Gustav Eek

Two fish species previously referred to as Betta sp. ‘Sengalang/Palangkayara’ and Apistogramma sp. ‘masken’ in the aquarium hobby have been described on an official basis in the last couple of weeks. Continue reading »

Adult male; red operculum morph (?) © Tino Strauss

Scientists from the University of Alberta, Canada, have ascertained that the pH of water influences both sex ratio and male phenotype in the kribensis, Pelvicahromis pulcher. Continue reading »

Apistogramma spp. will be the focus of the BCA’s Spring convention 2013. © Kris Weinhold

The British Cichlid Association (BCA) has been a respected mainstay of the aquarium hobby for around 40 years, and for cichlid hobbyists resident in the UK the group’s 2013 conventions promise to deliver a real treat.

The annual Spring event takes place on Sunday 5th May and should prove unmissable for anyone interested in dwarf cichlids of the genus Apistogramma with guest speakers Mike Wise (USA) and Tom Christoffersen (NOR) among the world’s most respected names in the study and husbandry of this diverse group of dwarf cichlids.

Mike and Tom have made a number of expeditions to the Amazon region, and will share their sometimes very amusing rainforest experiences as well as revealing how they transport their fish back home from some of the most remote regions in South America and breed them in their tanks.

Check out Tom’s brilliant website for a flavour of what’s in store.

The Autumn convention is scheduled for Sunday 8th September and true to form the BCA have selected a contrasting yet equally appealing range of guests for their second convention of the year.

First up is well-travelled cichlid enthusiast Ted Judy (USA) who will share details of his collection trips to Gabon in West Africa alongside Dr. Anton Lamboj.

A light lunch may be in order, since the afternoon lecture will be chaired by Dr. Peter Burgess, biologist and consultant for a well-known range of fish foods and aquarium products.

Dr. Burgess will guide attendees through a fascinating and undoubtedly vivid collection of parasites known to infest cichlid species, including the discus, Symphysodon spp., and explain the various options available to eradicate them.

Both events take place at the Lodge Park Social Club, Lodge Pool Drive, Redditch, Worcestershire, B98 7LH, with entry charged at £3.50 for BCA members or £5.50 for non-members.

In addition to the superb presentations there will also be auctions of fish and other items with some rare species and real bargains to be picked up, plus an extensive raffle sponsored by several leading manufacturers.

For further information check out the BCA website or their busy Facebook page.

As well as their standard membership package the BCA also now offer a special online membership offer at a bargain price, click here for more details.

PS: If you’re involved with a club and fancy promoting your activities via SF, please feel free to drop us a line on the forum or by private message.

We’re more than happy to feature both national and local events taking place anywhere in the world, so get in touch and spread the word to other hobbyists in your area and beyond!

© Damiano Luchetti

Two new, closely-related species of zooplanktivorous haplochromine cichlid from southern Lake Victoria, Tanzania have been described in the journal ‘Zookeys’.

Continue reading »

In the first of what we hope will become a regular interview series, we catch up with Malawi cichlid expert, ichthyologist, aquarist and webmaster Michael K. Oliver.

Cyrtocara moorii is one of over 400 cichlid species described from Lake Malawi to date. © Phil Sieradzan

Michael has a long-standing interest in the cichlids of Malawi and both his Masters and Ph.D. research concerned the systematics of African cichlids.

He’s made a number of trips to the lake, diving alongside the fish and collecting specimens for scientific research, and has described some notable species including the rusty cichlid, Iodotropheus sprengerae, and sulphur-headed hap, Otopharynx lithobates.

Since 1997 he’s also been running The Cichlid Fishes of Lake Malawi, Africa, an incomparably comprehensive, open access resource for anyone interested in the lake and its species.

SF: Michael, you’ve been studying the cichlids of Malawi for almost five decades. How did you become interested in these fishes, and how has your passion been maintained?

MKO: Great questions, but they don’t have short answers. Well, first came a general interest in fish, by the time I was ten or twelve. I’ve thought a lot about how that happened – because, why would it? I grew up in Los Angeles, but not near the ocean.

My earliest memories involving fish are of a small aquarium in the office of my childhood pediatrician, and of several family visits to Marineland of the Pacific, an oceanarium (now long gone) with a giant ocean tank, smaller aquariums with fishes from various habitats, performing dolphins and seals – all the usual trappings.

Staring up at the jacks and sharks and bright orange Garibaldis swimming around the huge tank must have made quite an impression on my sub-teen self.

At 14 I got my first aquarium and soon started collecting books about fishes, including scientific books. I joined the Los Angeles Aquarium Society and became active in it, and haunted the local fish stores every few days (sound familiar so far?).

Thumbi Island. East, where Michael first collected cichlids in 1968. This photo was taken from an Air Malawi Britten-Norman Islander just before landing at Monkey Bay in 1971. © Michael K. Oliver

By age 16, I knew I wanted to become a fish taxonomist. I became aware of Malawi cichlids during these teen years. In about 1966 I saw a couple of colorful species of mbuna at my local fish store (US$40 each, I think), and also remember reading an article about them in TFH.

My mother and I traveled quite a bit with my father, a college English professor, during his sabbatical leaves every few years. When I was 17 we were planning a trip that included traveling through Africa from south to north. I successfully lobbied my dad to let me arrange a stop in Malawi; I wanted to see those colorful cichlids in the wild.

I wrote to the Chief Fisheries Officer of Malawi (an actual paper letter in an airmailed envelope – this was the 1960s). It was forwarded to the Senior Fishery Research Officer, Mr. David Eccles, in Monkey Bay. Eccles invited us to visit him there and promised to introduce me to the local fishes.

Talk about a dream come true! David, an excellent naturalist, was a wonderful host during the couple of days my parents and I stayed in Monkey Bay. He organized collecting trips to the rocky shores of Thumbi Island East, and seining on the local sandy beach. We caught dozens of cichlids of many species, as well as cyprinids, catfishes, and even Aplocheilichthys johnstoni. I was like a kid in a … cichlid store.

The sheer diversity of the cichlids was stunning (and only a couple of hundred species were known then). Seeing many of those 40-dollar cichlids at one time, glowing with color, fresh from their rocky habitat, also made an impression. Those two days with Eccles focused my future studies specifically on Malawi cichlids.

Michael described Otopharynx lithobates in 1989. © Unimati.dk

Incidentally, he and I collected specimens of a pretty little rocky shore cichlid with three spots. David wasn’t sure what it was, and my continuing efforts to identify it back in Los Angeles and at the British Museum eventually showed conclusively that it was undescribed. Years later I described it under the name Otopharynx lithobates, and it has justly become a favorite in the aquarium.

With that 1968 visit, I think I was the first American to visit Lake Malawi because of its fishes since the 1929 American Museum of Natural History collectors, R. and L. Boulton, caught a few fishes at Karonga and Deep Bay. I had taken lots of photos, and after returning home I wrote a three-part illustrated article for The Aquarium magazine about my visit to the lake, published in 1970.

At the time, only mbuna were exported to the aquarium trade; my article emphasized that there were a lot of colorful non-mbuna cichlids that would be very suitable for the aquarium.

When I later met Peter and Henny Davies, the first exporters, in Malawi in 1971, they told me that my article had persuaded them to begin exporting non-mbuna “haps,” which did indeed become very popular.

As for how I maintain my interest – how could it be turned off, once started? I think E. O. Wilson’s idea of biophilia – an “urge to affiliate with other forms of life” – has merit, but I believe that some people also feel a more specific attraction to biodiversity.

Certainly I do; in addition to my cichlid interests, I collect beetles, which make cichlids look depauperate!

Apart from Malawi cichlids, Michael’s other great passion is beetles, such as this Neoclytus scutellaris. © Michael K. Oliver

SF: How complete is the taxonomy of cichlids from the lake? Do you think there are still new discoveries to be made?

MKO: Oh, certainly there are many species still to be discovered. About 400 cichlid species have been described to date from Lake Malawi.

Between 450 and 500 additional suspected species have been collected, photographed underwater, or both, but remain undescribed. Just about any visit to a remote rocky shore produces further new discoveries, as do deep-water trawls in unsampled areas.

If recent experience with other fish groups is any indication, there are also likely to be some cryptic species discovered among what we now regard as a single morphologically defined species.

Beyond just recognizing and naming all the species, most of the details of their phylogenetic interrelationships are still unclear; when better understood, this is sure to cause extensive revision of their generic classification.

SF: You recently described a new species which is interesting in a number of ways. Can you tell us about it?

MKO: Hemitaeniochromis brachyrhynchus is the second species of its genus to be described. Compared to the much better known H. urotaenia, the recently named form is anatomically odd.

The head appears scrunched up in front of the eyes because the superficial bone (the lacrimal) between the eye and the upper jaw is extremely narrow, giving the fish an unusually short snout (brachyrhynchus means short snout). The eyes are also unusually large.

Michael recently described Hemitaeniochromis brachyrhynchus which is currently known from just two specimens. © Michael K. Oliver

Circumstantial evidence strongly suggests that H. brachyrhynchus is a paedophage – one of the cichlids that steal and eat eggs or fry being brooded in the mother’s mouth.

But, only two specimens are known; even the life coloration is a mystery. (The description was published as an open-access paper, so if interested to learn more about this cichlid you can freely download it from http://www.mapress.com/zootaxa/2012/f/zt03410p050.pdf.)

SF: Are wild Malawi fish still collected for the aquarium trade and does this practice have any detrimental effect on natural populations?

MKO: Yes, there is at least the large enterprise founded by the late Stuart Grant (see http://lakemalawi.com) at Kambiri Point near Salima, Malawi.

The most significant damage to natural populations by aquarium collectors was probably the intentional transplantation of cichlids between different parts of the lake by the Davies some 40 years ago, simply for their own convenience (for instance, taking Likoma Island endemic cichlids and releasing them at Thumbi Island West, over 200 km away).

This selfish, reprehensible action allowed hybridization and competition between species that would never have encountered each other naturally, and further complicated attempts to understand the already extremely complex natural distribution patterns of the cichlid species.

I’m not aware of any instance where overcollecting for the aquarium trade has seriously threatened a cichlid population in Lake Malawi, but given the small size of the populations of some species, an irresponsible collecting operation could definitely reduce a population (and even some entire species) to a level where it could not sustain itself.

Lake Malawi is home to an enormous diversity of cichlid species, with more than half probably still undesrcibed to science. © Sarah Depper

SF: Do you think that the speciation of Lake Malawi cichlids will ever stop, and is it possible that some extant species may disappear naturally as a result of the ongoing speciation process?

MKO: As long as there are cichlids in Lake Malawi, they will continue to differentiate and speciate, facilitated no doubt by continued fluctuations in lake level which isolate and reconnect stretches of rocky shore, and by other factors such as changes in female preference for male coloration.

Extinction is also a natural process. Many species have surely arisen only to disappear within the lake, and more will do so.

I like to imagine that, a few tens of millions of years from now, as the two tectonic plates between which Lake Malawi lies continue to pull apart – the Rovuma plate on the east, the main Nubian plate on the west – the lake will become connected to the southern Indian Ocean and some future cichlids will adapt and invade the coral reefs.

SF: What do you see as the biggest challenge(s) facing the lake in the future?

MKO: Unfortunately, I think there are several.

1) Continued commercial overfishing in the south (ring-netting, trawling) has already probably driven some cichlid species to extinction and others to its brink. Artisanal fishing by the lakeshore villagers has also become a significant threat in some areas; very long and fine-meshed nets that catch even juvenile cichlids in large numbers are now widely used. See video:

2) Siltation from agricultural runoff into the lake is occurring, notably in the southwest near Salima.

3) Chemical pollution from agriculture could also increase, its effects aggravated by the long turnover time of the lake with its single outflow, the Shire River.

4) Oil spills from planned drilling in the north are a potential concern.

SF: What do you consider the most important items to pack when on a Malawi field trip?

MKO: I’m afraid it’s been a long time since I was there. Some obvious suggestions: For fishwatching, snorkel and mask (fitted with prescription lenses, if needed); a neoprene vest; for SCUBA: your second-stage regulator, dive computer, c-card, dive log, maybe a camera; extra batteries (perhaps rechargeable, with charger) and extra digital storage media.

The friendly and helpful local people always appreciate a few inexpensive gifts such as wristwatches and tee-shirts.

Rhamphochromis are an important food fish and come highly recommended by Michael! © Alexandra Tyers

SF: Finally, perhaps inevitably, fish or chips?

MKO: Both, of course! Keeping to Lake Malawi, grilled tilapia fresh from the lake are hard to beat, but smoked mcheni (Rhamphochromis) are also delicious, if rather bony. Even dried usipa (Engraulicypris sardella) are interesting to taste – once (it probably helps if you like the taste of gallbladder…).

SF: Mmmm, gallbladder! Michael, it’s been a great pleasure having you take part in our first ever SF interview. Thank you very much and all the best for 2013!

MKO: Thank you so much for the honor of being your first interviewee! It has been an entirely positive experience.

We’re going to be running a series of interviews with famous icthyologists and well-known figures from the aquarium world over the coming months, and we want you to ask the questions!

Our first victi…errr…interviewee is Malawi cichlid expert Dr. Michael K. Oliver, who recently described a new species, Hemitaeniochromis brachyrhynchus, from the lake which is intriguing in that no-one knows how it looks when alive with only two preserved specimens known.

Continue reading »

Mike Norén

Biologists from the University of Bonn have discovered that the dwarf cichlid Pelvicachromis taeniatus can perceive light in the near-infrared range, a phenomenon previously considered unlikely at best and which apparently helps them hunt prey in shallow African rivers. Continue reading »

Live specimen photographed at Kerenge Island. © African Diving Ltd.

A new species of the piscivorous cichlid genus Lepidiolamprologus is described in the journal ‘Zootaxa’ this week. Continue reading »

© F. P. Ottoni et al.

A new species of the dwarf cichlid genus Laetacara was described in the journal ‘Vertebrate Zoology’ recently, bringing the total number of species in the grouping to seven.

L. flamannellus (Ottoni et al. 2012) is known only from the coastal floodplains of eastern Amapá state, northern Brazil, where it occurs in highly seasonal habitats. Between July and December permanent water is restricted to the major river channels, but during the wet season from January to July these overflow significantly and flood large tracts of the surrounding grasslands and forest. Continue reading »