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C M Y K Offshore CichlidsofLakeMalawi Offshore of OFFSHORE CICHLIDS Lake Malawi is internationally renowned among evolution- ary biologists and aquarium enthusiasts alike for its immense number of of fishes, which display a dazzling of repertoire of structural diversity, feeding specialisations, and bril- liant colours. Almost all of the attention has focused on the rocky shore fish or , and it has been assumed that the offshore waters and the sandy or muddy shores harbour a relatively low- LAKE MALAWI diversity community dominated by non-cichlid fishes.

This book, the first ever guide to the cichlid fishes of the off- shore waters of Lake Malawi, shows that this habitat harbours a great diversity of little-known cichlid species, which dominate the catches of the commercial and artisanal food fisheries. How they evolved, how they continue to coexist, and how to manage George F. Turner their exploitation for food while conserving these unique spe- cies will be major challenges to scientists and policymakers. Aquarists will be fascinated by the wealth of previously-unknown brilliantly-coloured and bizarrely-shaped species.

Information on distinguishing features, colour, size, distribution and abundance, commercial importance, diet, reproduction, and are given for 199 species, 79 of which are pres- ently undescribed. 186 colour and 44 black and white photo- graphs illustrate all species. Also included are figures showing the main distinguishing features of the fish, maps, a glossary of George F. Turner technical terms, a full bibliography and brief discussion of the evolution, and conservation of these fishes.

The author, Dr George Turner, has worked on African cichlid fishes for 13 years and has spent more than 3 years in Malawi. Most of the information collected in this book was obtained while working on the commercial fisheries of the southern lake for the UN Food and Agriculture Organisation from 1990 to 1992. He is presently lecturer in Ecology at the University of Southampton, England, and continues to carry out research on evolution, ecology, behaviour, taxonomy, conservation, and management of African freshwater fishes.

ISBN 3-928457-33-0

C M Y K 2 Offshore Cichlids of Lake Malawi

1 2 Offshore Cichlids of Lake Malawi

George F. Turner

Department of Biology University of Southampton England, UK

3 Photo cover: 'long-fin yellow'

ISBN 3-928457-33-0

Copyright © 1996. Cichlid Press. All rights reserved. No part of this publication may be repro- duced, stored in a retrieval system, or trans- mitted in any form or by any means—elec- tronic, mechanical, photocopying, record- ing, or otherwise—without prior permission of the author and the publisher.

Cichlid Press, Ahornweg 3, D-31864 Lauenau.

4 Contents

Chapter 1. Introduction ...... 7 Chapter 2. Methods ...... 10 Chapter 3. Habitats, fish communities and their exploitation ...... 12 Chapter 4. Introduction to the cichlid species...... 20 Chapter 5. Tilapiines ...... 27 Chapter 6. Mbuna ...... 32 Chapter 7. Ncheni and Ndunduma ...... 38 Chapter 8. Utaka ...... 66 Chapter 9. and allied genera ...... 76 Chapter 10. and ...... 100 Chapter 11. hennydaviesae complex ...... 113 Chapter 12. Miscellaneous barred species ...... 119 Chapter 13. ...... 129 Chapter 14. and other oblique-striped species ...... 133 Chapter 15. and other spotted species ...... 152 Chapter 16. and other horizontally-striped species ...... 190 Chapter 18. and other blotched species ...... 202 Chapter 19. Discussion ...... 209 Glossary ...... 221 References ...... 231 Index ...... 236

5 6 Chapter 1 Introduction

The purpose of the present book is cles & Trewavas 1989), before the in- to provide an introduction to the off- troduction of offshore research vessels shore cichlids of Lake Malawi and to or commercial trawlers allowed the give some assistance in their identifi- deeper regions of the lake to be ex- cation in the field. plored. Although further ad hoc col- lections and descriptions of a few spe- Since the pioneering work of Fryer cies have been made by Burgess & (1959), the inshore cichlid fishes of Axelrod (1973), Eccles & Lewis (1977, Lake Malawi, particularly the rocky 1978, 1979), Stauffer & McKaye (1986, shore species, have been internation- 1988), Konings (1990a, b, 1993) and ally recognised as an outstanding ex- Turner (1994a, b, c), at least a third and ample of rapid speciation, with the probably more than half of all offshore potential to provide us with critical species are not only lacking a formal insights into the whole evolutionary description, but have never been men- process. Malawian cichlids are also tioned in print. well-known as a tourist attraction at Many of the fishes covered in the the Lake Malawi National Park, and present work are sometimes found in are kept by thousands of aquarists all inshore habitats — often principally in over the world. The excellent publica- these areas — although all occur to tions of Ribbink et al. (1983) and Ko- some extent further offshore in waters nings (1989, 1990), based largely on difficult to survey with SCUBA equip- SCUBA surveys, provide a fairly com- ment, either because of depth or the prehensive coverage of the distribu- turbidity of the water. Such areas are tion, ecology, and distinguishing fea- also those which are easiest to fish with tures of the shallow water cichlids, mass-catching, nonselective gears such particularly those of the rocky shore. as beach seines and trawls. From an Although most inshore species have evolutionary biologist's point of view, still not been formally described, the the offshore cichlids are interesting as great majority can be easily identified they appear to be free (or at least freer and now have fairly well-established than rocky shore fishes) to move informal names. around the lake. Since most theories of Those species which cannot readily speciation assume that populations be collected by SCUBA divers remain have to be separated geographically, it very poorly known. The only substan- is a greater challenge to explain how tial work on the taxonomy and identi- these fishes have evolved into so many fication of the offshore cichlids is species. Consequently, among the largely based on studies of preserved fishes of Lake Malawi, the offshore specimens collected prior to 1930 (Ec- cichlids are (i) the most important for

7 human food, (ii) the most interesting lected several species of to evolutionary biologists, (iii) the spe- during the 1950s, D. S. C. Lewis made cies most likely to be in danger of ex- a large collection of Rhamphochromis in tinction, and (iv) the least known. the 1970s, and D. H. Eccles collected I would like to make it clear that the numerous Lethrinops and other off- present work should on no account be shore species. Even a few preliminary regarded as definitive. There are un- studies on these collections would doubtedly many mistakes. The collec- have been very valuable to me and tions were obtained opportunistically other workers on Lake Malawi. Even and not as part of a systematic survey now, very little of this material has been programme. All the specimens came worked on, many of the specimens from the southern part of the lake. This have been destroyed or their collection book gives a brief outline of what little labels lost. Many other major studies is known of those cichlid species which on Lake Malawi made no taxonomic I encountered during my research on collection at all, such as those funded the fisheries of southern Lake Malawi by FAO in the 1970s and 1980s, ODA and Lake Malombe in 1990-92. Of these in the 1980s and 1990s, and the current species, 83 were undescribed at the World Bank/ ICEIDA project. There- time I carried out the field work, al- fore, each new study of the commer- though 4 have subsequently been for- cial fisheries or offshore fishes has be- mally described by myself or Ad Ko- gun from a basis of limited knowledge nings. Of the undescribed species, I and researchers have either had to re- was able to reliably identify 11 of these peat previous work or more often have on the basis of temporary names as- just avoided trying to identify the signed to them in the collection of the component of the field museum of the Monkey Bay Fish- catch. The great underestimation of the eries Station in Malawi, mostly by D. species richness of the offshore cichlids H. Eccles. I have generally retained the has probably been an important factor temporary 'specific' names, which are in ensuring that little attention has in widespread use within the Ma- been paid to the conservation of these lawian fisheries department, although fishes, while theories about the evolu- I have tried to assign them to the most tion of Malawi cichlids have not really appropriate of the genera described by taken the offshore fishes into consid- Eccles & Trewavas (1989). As far as I eration. I hope that the present work am aware the remaining 68 species represents a step, albeit a small one in have never been identified prior to the the direction of correcting these defi- present study and no information ex- ciencies. ists in any published form whatever. My justification for publishing such During my research on Malawi a preliminary piece of work is that fishes, I have received an immense there is no other field guide for the off- amount of help and encouragement shore cichlids of Lake Malawi. Since from many people. I would like to take the 1930s many collections of offshore the opportunity to thank (in approxi- cichlids have been made: T. D. Iles col- mately chronological order) Tony

8 Pitcher, Eva & Bodo Hert, , Ro Lowe-McConnell, David Eccles, John Tarbit, Stuart Grant, Irv Kornfield, Peter Reinthal, Denis & Sharon Tweddle, Ojinja Mhone, James Whitimani, Lameck Phiri, Jay Stauffer, Ken McKaye, Tony Ribbink, Niek & Carla van Zalinge, Mohammed Seisay, Marina Mdhaili, Steve Alimoso, Nel- son Mwanyama, Moses Banda, Eddie Allison, Ben Ngatunga, Tony Thomp- son, Andy Menz, Dick Beales, John Howarth, Martin Taylor, James Deutsch, Ciro Rico, Gary Carvalho, Rick Shaw, Matt Arnegard, Mairi Knight, Lucy Ambridge, and Cecilia Cruz. I am especially grateful to Ad Konings for the time and effort he has invested in thoroughly reading this manuscript and for many helpful dis- cussions and corrections. My research on Lake Malawi has been supported by the Overseas Development Admin- istration (UK), the Food and Agricul- ture Organisation of the UN, and the Natural Environment Research Coun- cil (UK). My work has been made pos- sible by the kind cooperation of the Chief Fisheries Officer, Boniface Mkoko, the Chief Parks & Wildlife Of- ficer, Matthew Matemba and the Direc- tor of Research at the University of Ma- lawi, Sosten Chiotha. Above all, thanks to Rosanna Robinson, for the tremen- dous support and help she's given me throughout this time — as well getting with her own research on Malawi cichlids.

9 Chapter 2 Methods

Much of the information on distri- Commercial Trawl Sampling. Data bution, diet, maximum size, etc. comes on the abundance and distribution of from previously-published studies. In demersal fishes were taken from analy- addition, I present new data from the sis of 35 samples (17,000 fish) from the following surveys:

1991 & 1992 Surveys. As part of an ODA- funded survey of de- Karonga. mersal fish stocks, 154 samples, comprising al- TANZANIA most 110,000 fish weigh- Chilumba . ing approximately 1,639 kg, were analysed from experimental bottom trawls made in the SE Usisya. Arm (Figure 3). Samples were collected in May, Nkhata Bay. August and November 1991 and February and Likoma Island May 1992. Species com- position by weight and MOZAMBIQUE MALAWI numbers was deter- mined for all samples. Nkhotakota. These samples were col- lected by staff of the Monkey Bay Fisheries Domira Bay Station, and I was not Senga Bay able to accompany the SE Arm research cruises. Conse- SW Arm quently information on Upper Shire River live colours is lacking for Lake Malombe those species not col- lected from other sur- veys. Figure 1. Lake Malawi, showing national boundaries and main collecting locations.

10 commercial 38 mm midwater trawl. Museum Collections. I examined This gear is targeted on chambo (Oreo- preserved material at Natural History chromis spp.), but has a large 'bycatch' Museum (London), the Monkey Bay of smaller haplochromine cichlids. Fisheries Station field museum, some Quantitative figures are given for the specimens collected by the ODA/ percentage composition of this by- SADC pelagic fish resources project catch. Also examined were 13 samples from open waters of the lake, and ma- (8,000 fish) from commercial stern terial loaned from the Berlin Museum trawlers, and 6 samples (2,500 fish) of Natural History. from commercial pair trawlers. All of these samples were taken from the SE Arm. In order to determine the catch composition of artisanal seine fisher- ies which were catching juvenile Oreo- chromis as a bycatch, 15 samples, total- ling more than 9,000 fish were exam- ined. Species composition by weight and numbers were determined. Addi- tionally, the length distribution of all species was obtained from 8 samples. A number of ad hoc observations of the species caught by seines and other gears were made during the course of the project.

Photographic Surveys. As part of the research of the FAO Chambo fisheries project, I participated in a few survey cruises aimed at elucidating the distri- bution of immature spp. I was able to collect and photograph fresh specimens from 15 trawl hauls throughout the southern part of the SE Arm. I was also able to examine a fur- ther 9 hauls taken from the vicinity of Monkey Bay and accompanied the Research Vessel Ethelwynn Trewavas on a two-day sampling cruise around Domira Bay and the Maleri Is. and col- lected a number of specimens from 9 hauls. Quantitative analyses of these catches were not made.

11 Chapter 3 Habitats, fish communities and their exploitation

The fish communities of all habitats During the dry season, most rivers con- in Lake Malawi are dominated, both sist of a series of separate muddy in terms of species richness and total pools, often heavily overgrown with biomass, by endemic haplochromine swamp vegetation. These waters con- cichlids. Moving downstream along tain a fairly uniform fauna dominated the Shire River, the lake's only outlet, by non-cichlid fishes, especially Clarias there is a progressive and gradual at- gariepinus and small Barbus spp. as well tenuation of the endemic cichlid com- as the larger Barbus johnstonii. The non- ponent which dominates the Upper endemic cichlids Oreochromis shiranus, Shire River and Lake Malombe, de- Tilapia rendalli and calli- clines in the middle Shire River, par- ptera are also common. Two further ticularly south of the Liwonde Barrage, cichlid species, Tilapia sparrmanii and and disappears entirely to the south of philander, are occa- the cataracts of the Kapichira Falls. sionally found in such smaller water This represents a major biogeographi- bodies, but have never been recorded cal barrier (Tweddle et al. 1979). Down- from the main lake. Endemic Lake stream of the falls, the fish fauna is Malawi cichlids rarely venture far into Zambezian in origin, and contains the rivers or flood plains. such non-Malawian species as the Thus, we can delineate two clear-cut tigerfish (Hydrocynus), the electric cat- faunal areas which may be referred to fish (Malapterurus), the vundu (Hetero- as the 'Lake Malawi Faunal Area' branchus), and the moonfish (Disti- (Lakes Malawi and Malombe, Upper chodus), as well as occasional exotic Shire River), and the broader 'Ma- species such as the Zambezi shark lawian Faunal Region' which also (Carcharinus leucas) and sawfish (Pristis includes the inflowing rivers and spp.). Cichlids are a relatively trivial streams, various smaller lakes and component of the community, the most pools, as well as the middle Shire River abundant being the tilapias Oreochro- and Lakes Chilwa and Chiuta. It is mis placidus and O. mossambicus. The with the former that the remainder of Zambezian fauna is comprehensively this book is concerned. covered by Skelton (1993). In general, the rivers and streams Lake Malombe and the flowing into Lake Malawi, Malombe Upper Shire River and the Upper and Middle Shire Riv- ers are turbid and highly seasonal, with Lake Malombe is connected to the only a few large permanent rivers. main lake by a short stretch of the slow

12 flowing Upper Shire River. It is shal- and Protomelas similis. All of these spe- low (average depth 4 m in 1992), cies feed on benthic invertebrates, with muddy and highly productive. Mean the exception of P. similis (a macrophyte transparency is around 2.4 m (Secchi feeder) and the two disc) and chlorophyll-a concentration (zooplankton feeders). Other rarer averages 4.9 µg/l (Mwanyama, 1993). included the epi- During the 1940s, the shores were phyte-feeder Hemitilapia oxyrhynchus, mostly forested and reedy (R. H. Lowe- the molluscivore pla- McConnell, pers. comm.). In the 1960s- codon, the benthic macro-invertebrate 70s, trawling was impracticable be- feeders Aulonocara guentheri, Placido- cause of the high concentration of sub- chromis subocularis, Mylochromis labi- merged macrophytes (D. H. Eccles, dodon, an unidentified Mylochromis pers. comm.). By the early 1990s, al- spp., the micro-invertebrate or diatom most all of the shoreline had been de- feeders intermedius, forested and most of the reeds have livingstonii, and Ps. been cleared, except in the small area elegans, the piscivorous Rhamphochro- on the south-eastern side within the mis spp. (generally deep-bodied forms Liwonde National Park. Very little sub- with small heads), Buccochromis noto- merged vegetation remained by the taenia (juveniles), stri- time I worked there in the 1990s. Dur- gatus, D. compressiceps, ing the low water levels of the 19th christyi, Nimbochromis livingstonii, the century, Lake Malombe dried out com- scale-eater taeniatus, and pletely, leaving only a river channel in the generalist . what is now the centre of the lake. Not All 5 tilapiines species were re- surprisingly therefore, there are few, if corded, comprising 3% of the catch any, species endemic to the lake. The weight, but unquestionably caught in lake is intensively fished by artisanal larger quantities by other gears select- fishermen, although larger-scale fish- ing for large fish. Non-cichlid fishes ing, such as trawling, is prohibited by were fairly uncommon: Engraulicypris law. sardella (1%), Clarias gariepinus (1%), During the sampling of artisanal Opsaridium microcephalus (0.5%), with fisheries, 44 taxa were recorded. 30 occasional Bagrus, Labeo cylindricus, species of haplochromine cichlids ac- Barbus arcislongae, Brycinus imberi, counted for almost 95% of the catch (all Aethiomastacembelus, Opsaridium micro- figures given are percentage weight), lepis. but a few species were overwhelm- Overall, this appeared to be a low- ingly dominant: Lethrinops 'pink head' diversity community. It is not clear (33%), (22%), whether this is a natural phenomenon Otopharynx 'argyrosoma red' (21%). or has resulted from intensive fishing. Other species comprising 1-4% of the catch biomass were Otopharynx tetra- Area A stigma, Copadichromis 'chrysonotus black', Lethrinops macrochir, Placido- For the purposes of research and the chromis longimanus, Lethrinops lethrinus, licence of fishing craft, Area A is des-

13 ignated as the southern part of the SE vertebrate feeder Aulonocara 'yellow', Arm of Lake Malawi, south of Boad- the medium-sized chironomid feeders zulu Island. The maximum depth is Lethrinops christyi, L. altus, L. cf. parvi- approximately 50 m, although most of dens, L. longipinnis, L. lethrinus and the bottom is much shallower. The Synodontis njassae, the molluscivore shoreline is almost entirely sandy, usu- , the zooplank- ally with a high mud content. The wa- tivore Copadichromis virginalis, and the ter is relatively turbid, with a Secchi piscivorous Rhamphochromis spp. disc reading around 7.8 m. Chloro- (mostly juveniles). phyll-a concentration averages 1.66 Rarer species recorded were the µg/l, around double that of the lake predatory cyprinid Barbus litamba, the off Monkey Bay (Mwanyama, 1993). epiphyte-feeder Hemitilapia oxyrhyn- The area has long been heavily fished chus, the molluscivores Mylochromis by seines, gill nets, pair trawls and ring anaphyrmus, rhoadesii, and nets. Trematocranus microstoma, the long- The catch of the 1992 trawl surveys snouted chironomid feeders Taenio- was dominated by small haplochro- lethrinops furcicauda and T. praeorbitalis, mine cichlids. The small sediment the benthic feeders Otopharynx 'argyro- feeder Placidochromis 'longimanus soma deep', O. decorus, O. 'argyrosoma Namiasi' comprised 13% of the bio- red', cf leuciscus, Placido- mass, but it now appears that this cat- chromis 'long', Lethrinops polli, Lethri- egory is probably a mixture of at least nops macrochir, Otopharynx auromargina- 2 species. Other numerous small tus, lituris, Ctenopharynx cichlids were the sediment/plankton nitidus, Ct. intermedius, Mylochromis cf. feeders Ps. livingstonii (8.9%), Ps. balteatus, Placidochromis subocularis, elegans (7.6%), and Nyassachromis Lethrinops 'yellow', Placidochromis 'argyrosoma blue' (6.8%) and the mi- 'hennydaviesae III', Lethrinops micrent- cro-invertebrate feeders Trematocranus odon, Aulonocara cf. macrochir, and A. brevirostris (5.4%) and Aulonocara 'or- rostratum, the large piscivores Bucco- ange' (7.8%). All of these species ma- chromis nototaenia, B. rhoadesii, B. lep- ture at very small sizes. Five previously turus, Hemitaeniochromis urotaenia, Stig- unknown small Lethrinops species matochromis woodi, Otopharynx spe- (micrentodon Makokola, dark, blue- ciosus, and Nimbochromis livingstonii, orange, yellowchin, and pinkhead) to- the smaller predators gether comprised almost 15% of the benthicola, Mylochromis gracilis, Sc. sample weight. The sediment feeder psammophilus, 'gut- Lethrinops stridei was the only larger tatus', Taeniochromis holotaenia, Placido- species to be found in reasonable num- chromis 'macrognathus', the scale-eater bers, at around 5% of the catch, but Corematodus taeniatus, the paedophages only relatively small specimens were Hemitaeniochromis spilopterus and taken and these may have been con- liemi, and the zooplank- fused with L. 'micrentodon Makokola'. tivores Nyassachromis cf. eucinostomus, Other species comprising more than N. boadzulu, Copadichromis quadrimacu- 1% of the biomass were the small in- latus, Diplotaxodon limnothrissa, and D.

14 Area B

The area between the latitudes of Boad- zulu Island and Mon- key Bay has a varied Mbenji Island topography and is in some ways intermedi- Domira Bay ate between Areas A and C. The western Makanjila Point Makanjila coast is made up of a Senga Bay . series of alternating rocky headlands and AREA C Maleri Islands muddy bays, while the east side has a less Chipoka . indented, sandy shore Nankhumba Peninsula line. At the southern AREA B SW Arm extreme of the east shore, the Chapola Malembo . shoal is a huge shal- AREA A low sandy bank ex- Malindi . tending out towards .Mangochi Boadzulu Island. The main trawling grounds are the highly productive shelf areas Lake Malombe at 40-80 m depth just Figure 2. Southern Lake Malawi, showing trawling areas in the SE north of Boadzulu Is- Arm and other major collecting sites. land and 10 of the 16 trawl stations were at this depth range. argenteus. Samples were dominated by me- During the early 1990s, commercial dium-sized benthic invertebrate feed- fishing gears made substantial land- ers such as Otopharynx 'argyrosoma ings of Oreochromis spp. (using ring deep' (9.2%), Lethrinops cf. parvidens nets, gill nets and chambo seines) and (7.7%), L. longipinnis (6.5%), and Syno- Engraulicypris, (chirmila and beach dontis njassae (6.8%), along with the seine) as well as a few cyprinids and zooplanktivore Copadichromis virginalis clariid catfish. There was a large pair (7.2%) and the /algal feeder trawl fishery aimed at small haplochro- Pseudotropheus livingstonii (7.4%). mine cichlids at depths of 10-40m. Other species comprising between 1 Some Oreochromis, as well as bagrid and 5% of the sample weight were the and clariid catfish are also caught. medium-sized benthic feeders Oto- pharynx auromarginatus, Lethrinops

15 longimanus, L. microdon, L. gossei, T. 'deep', Aulonocara 'copper', Lethrinops furcicauda, L. altus, and Aulonocara cf. 'deep-water albus', and the small in- macrochir, the small benthic feeders vertebrate feeder Placidochromis Lethrinops 'dark', L. 'oliveri', L. 'grey', 'hennydaviesae II'. L. 'yellow chin', L. 'yellow', Aulonocara The fisheries in this area are domi- 'blue-orange', Nyassachromis 'argyro- nated by the large-scale commercial soma blue', Trematocranus brevirostris, trawlers of the Maldeco fishing com- the small algal/crustacean feeder Ps. pany. The 'midwater trawl', which ac- elegans, the molluscivore Mylochromis tually fishes just off the bottom rather anaphyrmus, the piscivorous Otophar- than in midwater, principally targets ynx speciosus, and various Rhampho- 1-2 year old juvenile Oreochromis spp., chromis species. All of the species found but the 38 mm mesh ensures there is a in Area A were also recorded in this large 'bycatch' which is processed and area, except Placidochromis 'longimanus sold. Most of this consists of the pelagic Namiasi', Lethrinops christyi, and some zooplanktivore Diplotaxodon limno- of the very rare species, such as B. thrissa, along with Rhamphochromis spp. litamba, M. cf. balteatus, C. liemi, N. and some bottom-living haplochro- boadzulu, Hemitilapia oxyrhynchus, N. cf. mines. The demersal stern trawler fleet leuciscus. Rare species found in Area B, operates in somewhat deeper water but not further south included the (50-75m) and is targeted on demersal large molluscivore Lethrinops mylodon, haplochromine stocks. Artisanal fish- the large benthic invertebrate feeders eries are mainly gill nets for Oreochro- laticeps and Lethrinops mis, chirimila (a sort of lift-net) for Co- furcifer, the medium-sized, Lethrinops padichromis and Engraulicypris, and 'loweae', L. 'macrostoma', Mylochromis beach seines for small haplochromines. melanonotus, M. 'double spot', Tramiti- chromis intermedius, Protomelas triaen- Area C odon, Otopharynx 'auromarginatus stripe' and O. 'productus', the preda- The northern part of the SE Arm has tors and Mylo- a very short steeply-shelving rocky chromis formosus, the small benthic shoreline on the western side, while the feeders Lethrinops 'matumbae', Nyassa- eastern shore is mostly open sandy chromis cf. microcephalus, Gephyrochro- beach, although there are rocky reefs mis moorii, Aulonocara 'pyramid', A. at the far north near Makanjila. Most 'green' and A. 'dark stripe', the zoo- of the area is very deep, 90-200 m. planktivores Mylochromis 'chekopae', Chinyankwazi and Chinyamwezi Is- Copadichromis pleurostigma and the fin lands arise from the deep lake bed in eater johnstonii. A number the eastern side of the lake. Water trans- of deep-water species were encoun- parency (12.5 m, Secchi Disc) and pri- tered here occasionally, including the mary production (0.82 µg/l chloro- predators Otopharynx brooksi, Alti- phyll-a) are lower than further south corpus 'geoffreyi', and A. mentale, the (Mwanyama, 1993). medium-sized invertebrate feeders In the standard trawl survey, 6 sta- Alticorpus pectinatum, Aulonocara tions were on the east coast (20-65 m

16 depth), 4 in deep water (90-128 m). A feeder Lethrinops microdon, the preda- further station at the Ilala Gap seemed tors Nimbochromis livingstonii, Alti- to cover a variety of depths from 20-80 corpus 'geoffreyi', and A. mentale, the m in a single haul, perhaps because of zooplanktivores Copadichromis vir- the steeply-shelving shore in this area. ginalis and Diplotaxodon 'macrops', the This station is ignored for the purposes benthic invertebrate feeders Alticorpus of quantitative analysis. Since not one macrocleithrum, Lethrinops 'deep-water cichlid species comprised more than albus', Lethrinops polli, Aulonocara 1% of the catch biomass in both the 'deep', Aulonocara 'yellow', and the in- eastern coast and deep water samples, evitable Synodontis njassae. Many small the two are treated separately. species of Aulonocara, Lethrinops, and Almost half of the fish biomass on the Placidochromis hennydaviesae-com- the eastern coast was comprised of plex, as well as a number of forms of three relatively large benthic inverte- Diplotaxodon and Rhamphochromis, ap- brate feeders, Synodontis njassae pear to be confined to deep waters. (18.8%), Lethrinops longimanus (12.5%), Other distinctive deep water species and Lethrinops longipinnis (12.2%). Also are Otopharynx brooksi, the vertically- common were the zooplanktivore barred Placidochromis 'acuticeps', P. Mylochromis 'chekopae' (7.3%), and the 'platyrhynchos' and P. 'macrognathus', benthic invertebrate feeders Otophar- and the large-mouthed elongate preda- ynx 'argyrosoma deep' (9.7%), another tors Pallidochromis tokolosh and Stigma- species (blue?) of the O. argyrosoma- tochromis 'guttatus'. Large Bathyclarias complex (6.2%) and Lethrinops cf. catfish were conspicuous components parvidens (5.3%). Other important spe- of the catch, and Bargus catfish and cies (forming between 1 and 5% of the mormyrids were occasionally seen. biomass) were Aulonocara cf. macrochir, On the western coast, fishing is Ctenopharynx nitidus, Lethrinops 'yel- largely confined to chirimila, gill nets low', Mylochromis anaphyrmus, Oto- and longlines for surface-living or deep pharynx 'productus', Pseudotropheus water fish, while it seems there is rela- lanisticola, Mylochromis spilostichus, tively low intensity of fishing on the Taeniolethrinops furcicauda and T. eastern shore because of the small hu- laticeps. Rare species not found else- man population and poor road com- where in the 1992 survey include Mylo- munications. A recent aid project by chromis sphaerodon, M. ericotaenia, Nim- the World Bank and Icelandic Govern- bochromis venustus, and Aulonocara ment has opened up a commercial guentheri. trawl fishery in this area, exploiting the The deep-water trawl samples were last largely intact community of off- dominated by the large benthic inver- shore demersal fishes in southern Lake tebrate feeder Lethrinops gossei (24.6%), Malawi. and the smaller benthic-feeders Lethri- nops 'oliveri' (23%), Alticorpus pectin- Domira Bay/ Maleri Islands atum (6.2%), and Lethrinops 'deep-wa- ter altus' (6.6%). Other important spe- No quantitative study of the fish cies (1-5% biomass) were the sediment communities of Domira Bay or the SW

17 similar to Diplotaxodon ecclesi, but with a pale belly. Chilinda It is surprising that demer- sal and pelagic offshore Ngusi cichlids should vary so much over a relatively short Chekopa distance, and it suggests that further offshore species Ilala Gap remain to be discovered in the northern parts of the Chipondo lake which remain unex- Monkey Bay Monkey Bay. plored.

Mazinzi Chirombo Bay The pelagic zone Fowo .Namalaka Mazinzi Bay Nkope The true pelagic zone of Nkhudzi Bay the lake has recently been Ulande studied in detail by the Chapola Shoal ODA/SADC project (Menz, Boadzulu Island 1995). The work of the Mpemba Michesi project focused mainly on the trophic interactions Maldeco. Namiasi within the pelagic waters, Namiasi . . and only a small taxonomic Palm Beach collection was made of the fishes (studied by Turner, Figure 3. SE Arm of Lake Malawi, illustrating trawling stations 1994b), so much is still to be (italics) used in the 1990-92 surveys and other main locali- learned about the distribu- ties. tions of the pelagic species. Fish biomasses were esti- Arm has ever been undertaken. The mated from acoustic surveys alongside Malawi Government fisheries research experimental gill net and trawl sur- programme involves a number of veys. The principal food resources for cruises to determine the total weight open water fish are zooplankton at the of fish caught per unit time at stand- surface mainly (principally ard stations. On one of these trips, I Tropodiaptomus cunningtoni), and pred- collected specimens of a variety of spe- atory fly larvae (Chaoborus edulis) cies. Some of these forms have not been deeper down. The total fish biomass collected from the SE Arm: Lethrinops was estimated at 168,000 tonnes, of 'Domira blue', Mylochromis 'deep', a which approximately 52% (87,000t) blue and yellow form of the O. argyro- was composed of Diplotaxodon limno- soma-complex, Aulonocara 'deep yel- thrissa, a small cichlid which mainly low', a deep-bodied form similar to eats copepods. The deeper waters, 150- Aulonocara 'pyramid', and specimens 200m, were dominated by Diplotaxodon

18 'bigeye', which probably comprises several species, together making up a standing biomass of around 33,000t. These feed mainly on Chaoborus larvae and seem to migrate up near to the surface at night. Other zooplanktivores included the utaka Copadichromis quadrimaculatus (8,700t) and the usipa Engraulicypris sardella, a cyprinid (5,100t), both largely inshore species. Surprisingly large numbers of Syn- odontis catfish (13,400t) were also found, feeding mainly on chaoborid larvae. Rhamphochromis spp. were the principal piscivores. There were around 11,300t of the small R. cf. longiceps (which is partly a zoo- plankton feeder) and 5,500t of the larger species. Predatory cyprinids, Opsaridium microcephalus and O. micro- lepis were much less important (1,300t).

19 Chapter 4 Introduction to the cichlid species

On the following pages, I deal with 2a. Gradual transition from large flank the offshore cichlid species of Lake scales to smaller belly scales. Gener- Malawi which I have actually identi- ally laterally compressed or elongate. fied in the field and, in the case of Background body colour usually sil- Rhamphochromis spp., also those mu- very...... 3 seum specimens which I have exam- ined. It seems premature to attempt to 2b. Abrupt transition from large scales present a formal key, as it seems likely on flank to small ones on belly. Gen- that many more species remain to be erally squat-looking, not laterally discovered, and in my experience in compressed, or elongate. Back- such cases it is tempting to simply as- ground body colour generally dark sign specimens of new species to what- or whitish with broad brown bars. ever taxon they key out as. In general, ...... Mbuna, chapter 6 the species are classified according to their most obvious feature, the mela- 3a. Generally marked with vertical, nin patterns of females and immature oblique or horizontal bars or with males. Thus I present a preliminary key spots. Species lacking spots or stripes to assist the reader in referring to the with small ventrally-placed or pro- appropriate chapter. trusible mouths...... 6

3b. Silvery, faintly countershaded but Key to the offshore cichlids of otherwise lacking a melanin pattern. Lake Malawi Mouth large or upwardly directed...... 4 1a. Lateral line kinked posteriorly, so that there is a single scale row be- 4a. Teeth widely-spaced, large, simple. tween upper and lower lateral lines Some of the anterior jaw teeth of the on the caudal peduncle ...... 2 inner series are enlarged. Generally elongate streamlined, reminiscent of 1b. Two rows of scales between upper a barracuda...... and lower lateral line on caudal pe- ...... Rhamphochromis, chapter 7 duncle. Normally deep-bodied her- bivores attaining large adult size, ju- 4b. Teeth closely spaced, anterior inner veniles with a large black spot on the teeth of lower jaw not markedly en- soft dorsal fin...... larged...... 5 ...... tilapiines, chapter 5

20 5a. Silvery, with strongly upwardly to end behind inner series...... directed mouth and closely packed Lethrinops and allied genera, teeth...... Diplotaxodon, chapter 7 chapter 9

5b. Silvery-white with large mouth and 9b. Lower jaw flattened, slender. Outer widely spaced teeth...... series of lower jaw teeth forms a sin- ...... Pallidochromis, chapter 7. gle straight series posterior to inner series. Eyes large...... 6a. Laterally compressed or slightly Placidochromis hennydaviesae elongate usually with numerous complex, chapter 11 long gillrakers. Mouth small, gener- ally strongly protrusible. Melanin 9c. Lower jaw more robust in appear- pattern simple countershading, ance. Outer series of lower jaw teeth either with a faint horizontal stripe forms a single straight series poste- or 1-3 prominent dark spots...... rior to inner series...... Copadichromis, Nyassachromis, chap- Miscellaneous barred species, ter 8 chapter 12.

6b. Mouth large and/or ventrally placed. Few short gillrakers (in most 10a. Melanin pattern a continuous or species)...... 7 broken oblique stripe...... 11

7a. Melanin pattern comprising mainly 10b. Melanin pattern spots or horizon- vertical bars, which are often faint. tal stripes...... 12 Occasionally a single midlateral blotch superimposed on the barred 11a. Large mouth & head. Large pattern...... 9 widely-spaced teeth...... Buccochromis spp, chapter 13 7b. Melanin pattern consisting of hori- zontal or oblique stripes or spots. .. 11b. Smaller mouth. Generally laterally ...... 10 compressed, but a few species slen- der and elongate. Teeth closely 8a. Cephalic lateral line system ex- packed, small...... panded to form large 'pits' on the un- Mylochromis and other oblique- derside of the lower jaw...... striped species, chapter 14 ... Aulonocara, Alticorpus, chapter 10 12a. Melanin pattern one or two hori- 8b. Pits on underside of lower jaw zontal stripes, sometimes with small or absent...... 9 fainter vertical bars...... Protomelas and other horizontally- 9a. Lower jaw flattened, slender striped species, chapter 16 ('weak'-looking). Outer series of lower jaw teeth curves posteriorly 12b. Melanin pattern consisting of spots or blotches...... 13

21 13a. Melanin pattern consisting of two measured from the tip of the snout to or three small dark grey or black the base of the caudal peduncle (which spots...... can be seen as a 'crease' when the tail Otopharynx and other spotted fin is folded to the side). Head length species, chapter 15 (HL, fig. 4.) is measured as the distance between the tip of the upper jaw and 13b. Melanin pattern consisting of nu- the rear of the operculum and snout merous large black or brown length (SN, fig. 5) from the anterior blotches...... edge of the orbital ring to the tip of the Nimbochromis and other spotted upper jaw. The caudal peduncle length species, chapter 18 is measured from the base of the last dorsal fin ray to the upper part of the 'crease' in the tail. Note that these meas- For each species known from the urements are different from those used field, information is provided on dis- by Eccles & Trewavas, who 'project' tinguishing features, colour, size, diet, their measurements on a fish board reproduction, distribution and abun- and take them parallel to the main axis dance, commercial important and mis- of the fish. I measure directly between cellaneous notes. points using a calliper, which I find DISTINGUISHING FEATURES: An attempt more accurate. In most other measure- is made to provide a list of features ments, I have followed Eccles & Trewa- which can be used in the field to dis- vas- lower jaw (fig. 5) is measured to tinguish each species. I have endeav- the posterior end of the jaw bone and oured as far as possible to use features is not the lower lip and preorbital bone readily seen without making time-con- depth (fig. 5) is measured down the suming counts or dissections. A few midline, not along the anterior edge standardised measurements are given. (which was the method of Green- Standard length (SL on figure 4) is wood). I also use a 'projected' maxi-

CPL

CPD MD HL SL

PEC

Figure 4. Standard measurements of fish. SL= standard length, MD= maximum depth, HL= head length, PEC= pectoral fin length, CPL= caudal peduncle length, CPD= caudal peduncle depth.

22 genera they are small or absent. Lethrinops and the related genera Tra- mitichromis and Taeniolethrinops are col- lectively distinguished by the arrangement of the teeth in the lower jaw. In the typical or 'Haplo- chromis'-type arrange- ment there is a long sin- gle row at the posterior end of the lower jaw. The Figure 5. Head measurements. LJ= lower jaw length, SN= snout length, PO= preorbital bone depth, CD= cheek depth, ED= outer row of the 'Lethri- horizontalye diameter, OW= overall width, IO= interorbital width. nops'-type curves round mum depth measurement (MD, fig. 5) since it is easy to do accurately with callipers. Overall width is measured across the closed opercula (fig. 5) and not over the abdomen, which is prone to swelling in some specimens, espe- cially if poorly preserved. Gillraker counts are generally given as 'lower' or 'ceratobranchial' gillrakers. The gill arch is hinged near the top and rakers on the hinge or above it are not in- cluded in this count (fig. 6). Scale counts follow Eccles & Trewavas- the most frequently used is the lateral line count, for which one counts the pored Figure 6. Diagram of gill arches, with operculum cut away. LGR= lower (ceratobranchial gill- scales in the upper lateral line until it rakers), GF = gill filaments. The mental process ends, then follows the transverse scale is indicated 'MP'. row down and forwards until reach- ing the lower lateral line and starting with the next scale. 'Expanded cephalic sharply to end just behind the inner pits' are an important feature of the rows (fig. 7). Some species show a genera Aulonocara and Alticorpus. rather intermediate form, e.g. Lethri- These are most easily seen by turning nops micrentodon, Ctenopharynx nitidus. the fish over and looking at the under- COLOUR: As it is probably the single side of the head (see Konings 1995b for most important identification feature, excellent illustrations). A number of I have paid particular attention to pro- large depressions will be seen. In other viding colour descriptions. It should be

23 stressed that most of these ioural observation. Other colour notes are based on information comes from specimens collected by unpublished studies by trawls and may not accu- Nelson Mwanyama and rately represent the colours an undergraduate thesis of the live healthy fish. by Karen Grant, both us- However, these are the col- ing material from my col- '' -type ours likely to be seen in lections. Sample sizes are any presently feasible way generally given, but are of sampling most of these small, and generally taken fishes. Males are generally from several fishes all larger and more brightly- from the same sample, coloured than females. which probably means Males may also have that geographic and sea- longer fins, as well as egg- sonal fluctuations in diet Lethrinops-type dummies — in haplochro- are underestimated. mines coloured spots on REPRODUCTION: Very little the anal fin, in Oreochromis Figure 7. Upper view of information is available lower jaw tooth arcades, (Nyasalapia) spp. a long rostral end uppermost. In for most species, espe- elaborate genital tassel. Im- the typical or 'Haplochro- cially those confined to portant features of the col- mis'-type, the outer tooth deeper waters. All known our pattern are shown on row continues posteriorly cichlid species of Lake Ma- as a single row. In the Leth- figure 8. Counts of vertical rinops-type the outer row lawi, with the exception of bars are given as the curves inwards posteriorly Tilapia rendalli, are mater- number under the dorsal to end just behind the in- nal fe- fin. This is because bars on ner rows. males carry the fertilised the caudal peduncle and eggs in their mouths, and forehead are much more prone to fade, in some cases actively defend fry and especially in preserved specimens than retrieve them into the mouth when those under the dorsal fin. threatened. Males never participate in MAXIMUM SIZE: Given as the largest parental care, but often construct large total length reported either from pre- display structures 'nests' or 'bowers' vious literature or (where no author- which are used to attract females, but ity is given) from my collections. not for the rearing of young. It seems DIET: Most information on diet comes safe to assume this breeding system for from previously-published informa- all haplochromine and Oreochromis tion. It should be emphasised that nei- spp. ther Eccles & Trewavas (1989) nor DISTRIBUTION AND ABUNDANCE: Where Konings (1989, 1990a, 1995b) usually no source is given, the data are my own give quantitative information and it and I have endeavoured to distinguish seems possible that some of their state- between positively identified speci- ments are based on inferences from mens later examined in the laboratory morphology, presently uncorroborated and field records, which are less trust- by stomach contents analysis or behav- worthy. Other information comes from

24 lt sub lap

sd dorsal fin     dorsum os na  sps sas cs pos flanks cau ulc

li op belly pa cn da pel es ch peb

Figure 8. Principal areas of colour pattern. Overall body colour is divided into dorsum or upper surface, flanks, and belly (often cited as 'ventrally'). Dorsal fin generally applied to whole of lower part of fin including soft dorsal (sd) unless latter is mentioned separately. Dorsal fin margin is also referred to as the lappets (lap), unless the lappet tips (lt) are differently coloured. Below the lappets may lie a submarginal band (sub), which is usually dark. Vertical bars are usually counted as just those lying under the dorsal fin (1-5). There may be up to three flank spots, the suprapectoral (sps), the supra-anal (sas) and the caudal (cs). The caudal fin (cau) applies to the whole fin, unless upper and lower caudal margins (ulc) are distinguished. Anal fin is often divided into proximal (pa) and distal (da), the latter also being referred to as the anal margin. Eggspots (es) are only taken to be the spots on the anal fin and not any on the dorsal or caudal fins. Pelvic fins (pel) often have a differently coloured anterior border or leading edge (peb). Pectoral fins (pec) are generally hyaline, or colour- less. Chest (ch) refers to the area just in front of the pelvic fins. Gular (not illustrated) is the branchiostegal membrane which lies between the opercula below the chin (cn). The lips (li) may be differently coloured, if not they are included with the chin or snout- the area between the lips and eye. Some species have a dark lachrymal or preorbital stripe (pos). Opercula (op) may have a dark spot (os) at the upper posterior margin. The nape (na) is the upper part of the head in front of the dorsal fin. Blaze refers to an area of bright colour extending from around the front part of the dorsal fin, down the upper surface of the nape, between the eyes and onto the upper surface of the snout.

published studies, and is of varying COMMERCIAL IMPORTANCE: The impor- reliability. Much of Eccles & Trewavas's tance of the species in fishery yields is information comes from trawl survey almost entirely based on my own sur- records collected many years before veys of the trawl fisheries of the SE thorough examination of the types and Arm and the seine fisheries of Lake other collections. In his earlier works, Malombe. Other fisheries have scarcely Konings often uses the phrase 'lake- been studied, but information on the wide distribution' rather freely. In fact artisanal fisheries of the Cape Maclear there are virtually no records of deep- and Nkhata Bay areas was obtained water samples from anywhere be- from the work of Iles (1960) and Smith tween Nkhata Bay and the SE and SW (1993). Importance in the aquarium Arms. trade is taken from Konings (1990) or

25 personal observations. NOTES: Information is provided on possible confusion in identification during my surveys or in previous works, as well as synonymy. Other pieces of biological information, such as growth parameters, are mostly from the work of FAO (1976) and Menz (1995).

26 Chapter 5 Tilapiines

The tilapiine cichlids have long been Oreochromis (Nyasalapia) of importance in food fisheries. The squamipinnis (Günther) three Oreochromis (Nyasalapia) species colour photo on page 49 are endemic to the lake and are very closely related, having almost certainly DISTINGUISHING FEATURES: A typical evolved within the lake (Sodsuk et al. Oreochromis with a broad head, grey- 1995). They are marketed as 'chambo' ish body and several prominent irregu- and command a much higher price lar vertical bars. Large specimens are than most other common species. They distinguished from those of other have been the subject of several stud- chambo by their generally deeper bod- ies aimed at finding out enough about ies, pale grey background colour, their biology and the fisheries targeted prominent vertical bars. They often on them to plan a rational management have a broad band of jaw teeth, with of their exploitation. Most of the early long flexible teeth in the outer row, and work is summarised by Trewavas a clear gap before the array of short (1983) and later work by FAO (1993). deeply embedded inner teeth. Male Over the last 10 years chambo popula- breeding colours are distinctive. tions have collapsed as a result of COLOUR: Grey, whitish ventrally, with overfishing. They are little known to broad irregular dark vertical bars. Ripe aquarists, as their export has been pro- males with a bright blue area (occa- hibited by the Malawian Government. sionally white or green) between the Two other tilapiine species are known eyes. The flanks and belly may be from the lake, neither is endemic and white, dark grey or almost black. each has invaded the lake independ- MAXIMUM SIZE: 37 cm TL. ently. DIET: Stomach contents of fish col- The tilapiine cichlids, along with the lected from Lake Malombe indicated non-endemic haplochromines Astato- that the diet was dominated by the tilapia calliptera and cladoceran Bosmina (43%) and the dia- robustus (both of which are confined to toms Aulacoseira (24%) and Surirella inshore swampy habitats), always (19%). In southern Lake Malawi, the have two clear rows of scales between diatoms Aulacoseira (80-81%) and Suri- the upper and lower lateral lines on the rella (6-11%) dominated (Mwanyama, caudal peduncle, while on the endemic 1993). At Cape Maclear, diets were haplochromines the upper lateral line more varied and comprised a variety kinks ventrally so that there is only of filamentous green and blue-green one row between the lateral lines algae as well as diatoms (Turner et al. posteriorly. 1991a). REPRODUCTION: FAO (1993) collected

27 ripe fish from October to March, Oreochromis (Nyasalapia) karongae although they were most numerous in (Trewavas) December to February. Ripe males colour photo on page 49 were 21-35 cm TL and females 20-37 cm TL. In Lake Malombe, ripe females DISTINGUISHING FEATURES: A typical had up to 1100 ovarian eggs, and the Oreochromis with a broad head, grey- fecundity (F) - length (len) relationship ish body and several prominent irregu- was F = 0.021. len(cm)3.07. Fecundity lar vertical bars. Large specimens are was significantly lower in the SE Arm, distinguished from those of other F = 0.021. len(cm) 2.90, with a maximum chambo by their generally brownish number of ripe eggs of 700. Specimens bodies, and are the only species with I collected near Karonga were mature bright yellow dorsal fin lappets. Gen- at a smaller size than those collected erally more slender than O. squamipin- in the south and a precociously mature nis, with smaller jaws and head than dwarf population was found in Lake O. lidole. They often have broad bands Chiwondo, an isolated lagoon south of of jaw teeth (4-15 rows), with long flex- Karonga. According to Trewavas (1983) ible teeth in all rows. Ripe males are males breed at depths of 16-20m, but I black. have observed territorial males as shal- COLOUR: Grey, whitish ventrally, with low as 2-3m depth. broad irregular dark vertical bars. DISTRIBUTION AND ABUNDANCE: Found Larger fishes are often brownish, with throughout Lakes Malawi and Ma- fainter bars, and bright yellow dorsal lombe and the Upper Shire River. fin lappets. Ripe males black with a COMMERCIAL IMPORTANCE: Substantial. broad white margin to the dorsal and Around half of the 620t of tilapias caudal fins. Anal fin with a white or landed in Lake Malombe in 1990-91 yellow margin. were this species, and around 1150t MAXIMUM SIZE: 38 cm TL. were landed in the SE Arm (FAO 1993). DIET: Stomach contents of fish col- It was the most numerous species in lected from Lake Malombe indicated the gill net (41% of tilapia catch) and that the diet was dominated by the the small pair trawl and demersal stern cladoceran Bosmina (35%) and the dia- trawl catches, but was also taken in toms Aulacoseira (21%) and Surirella substantial numbers by midwater (14%). In southern Lake Malawi, the trawl (26%), pair trawl (21%) and beach diatoms Aulacoseira (79-80%) and Suri- seine (32%). rella (5-13%) dominated (Mwanyama, NOTES: Specimens collected between 1993). At Cape Maclear, diets were Ngara and mouth of the Songwe had more varied and comprised a variety expanded pharyngeal dentition and of filamentous green and blue-green particularly bright male breeding col- algae as well as diatoms (Turner et al. ours. 1991a). REPRODUCTION: FAO (1993) collected ripe adults from July to March. The breeding season has two peaks, around September and February, in the SE

28 Arm. In Lake Malombe, there is a body colour is generally a dark grey. single peak from July to October. Males The lower pharyngeal bone has a long were ripe at 22-37 cm TL, females from blade and a short toothed area. Ripe 20-38 cm TL. Females produced up to males black. 1000 eggs, fecundity = 0.008 length COLOUR: Dark grey with several ir- (cm)3.84. At Cape Maclear, Turner et al. regular dark vertical bars. Ripe males (1991b) observed males defending black with a broad white margin to the nests of 25-190 cm diameter at depths dorsal and caudal fins. Anal fin with a of 0.5 to 28m. white or yellow margin. DISTRIBUTION AND ABUNDANCE: Found MAXIMUM SIZE: 37 cm TL. throughout Lakes Malawi and Ma- DIET: Stomach contents of fish col- lombe, and the Upper Shire River. lected from Lake Malombe indicated COMMERCIAL IMPORTANCE: In 1990-91, that the diet was dominated by the it comprised around half of the 600 cladoceran Bosmina (34%) and the dia- tonnes of tilapiines landed in Lake toms Aulacoseira (16%) and Surirella Malombe, while an estimated 650t (38%). In southern Lake Malawi, the were caught in the SE Arm of Lake diatoms Aulacoseira (47-66%) and Suri- Malawi (FAO, 1993), mostly in gill nets rella (11-14%) dominated, but the (36% of tilapia catch) and seines (35%) Diaptomus (13-29%) was also (Turner, 1995a). important (Mwanyama, 1993). At Cape NOTES: Specimens from muddy ar- Maclear, diets were more varied and eas, such as Lake Malombe, Nkhota- comprised a variety of filamentous kota, and the southern part of the SE green and blue-green algae as well as Arm, generally have few rows of jaw diatoms (Turner et al. 1991a). teeth, deep bodies and a slender pha- REPRODUCTION: Ripe males known ryngeal bone. These were formerly from the SE and SW Arms and near considered to be distinct species, Oreo- Salima, but not recorded from the chromis saka (Lowe). Around Karonga, northern part of the lake, or south of they have more slender bodies and Boadzulu Is., including Lake Malombe expanded pharyngeal bones. At Cape (Trewavas, 1983; FAO, 1993). FAO Maclear and Makanjila, dentition is (1993) collected ripe adults from Sep- variable 4 to 15 rows of jaw teeth and tember to February. Ripe females of 24- slender to broad pharyngeal bones. 37 cm TL produced up 700 eggs. Fe- cundity = 0.016 length (cm)3.52. Brood- ing females recorded from the Shire Oreochromis (Nyasalapia) lidole River and Lake Malombe. Fry are (Trewavas) brooded until at least 50 mm SL. Turner colour photo on page 49 et al. (1991b) recorded males defend- ing nests of 110-308 cm diameter at 17- DISTINGUISHING FEATURES: Large speci- 28m depth. mens have a large head, operculum DISTRIBUTION AND ABUNDANCE: Re- and mouth and a thin band (3-4 rows) corded throughout Lakes Malawi and of short, deeply-embedded teeth, usu- Malombe, and the Upper Shire River. ally set in clearly defined rows. The COMMERCIAL IMPORTANCE: Substantial.

29 In 1990-91, around 1600 tonnes were subspecies by Trewavas (1983) are caught in the SE Arm alone (FAO 1993). found in Lakes Chilwa and Chiuta. It It dominated the midwater trawl (60% is a fish of shallow water, generally in tilapiine catch) and ring net (75%) the vicinity of vegetation. catches, but was also taken in substan- COMMERCIAL IMPORTANCE: Taken in ar- tial numbers by artisanal seine (34%) tisanal gears, especially gill nets and and gill net (23%) fisheries (Turner, seines in shallow sheltered areas. Also 1995a). recorded from catches of pair trawlers. Widely cultured in fish farms, but it is prone to early maturation, like other Oreochromis (Oreochromis) shiranus Oreochromis of the O. mossambicus com- (Trewavas) plex. colour photo on page 49

DISTINGUISHING FEATURES: Deep-bodied Tilapia (Coptodon) rendalli and laterally compressed, golden- (Boulenger) brown with dark horizontal stripes. Differs from all haplochromines and DISTINGUISHING FEATURES: Deep-bodied other Oreochromis in normally having rather greenish fish with faint irregu- four spines in the anal fin. lar vertical bars and a 'tilapia-spot' at COLOUR: Females and immatures, the base of the soft dorsal fin, even in golden brown, darker, almost olive large specimens. Immatures occasion- green, dorsally, with two dark horizon- ally have a slightly reddish chin and tal bars. Ripe males black, with broad belly. red margins to dorsal and caudal fins. COLOUR: Generally whitish with MAXIMUM SIZE: 37 cm TL (Trewavas, green cast and faint vertical or horizon- 1983). tal bars. The anal fin and lower half of DIET: Macrophytes and detritus the tail are usually slightly reddish, (Jackson, 1960), and also benthic and and some immature fish also have a planktonic algae (Trewavas, 1983). reddish chin and belly. Mature adults REPRODUCTION: Ripe males dig simple do not show any breeding or parental crater nests, of 50-90 cm diameter, in colour. shallow water (15 cm-1.5m deep). The MAXIMUM SIZE: 35 cm TL (Konings, breeding season is September to March 1990a), but usually much smaller. or April (Trewavas, 1983). Females of DIET: Macrophytes (Jackson, 1960). 20-21 cm TL lay 520-618 eggs and REPRODUCTION: This is the only cichlid brood the young until they are about found in Lake Malawi which is not a 10 mm long. maternal . Between Sep- DISTRIBUTION AND ABUNDANCE: Re- tember and March, pairs guard eggs corded throughout Lakes Malawi and laid on a rock, or in a crater 40-90 cm Malombe, and the Upper Shire River. wide and 20 cm deep. Occasionally It is also frequently found in pools, and they may dig tunnels as deep as 43 cm, in streams and rivers flowing into the and only 11 cm wide. On one occasion lake. Similar specimens, ranked as a two pairs have been observed to co-

30 operate in guarding a brood of young (Ribbink et al., 1981). DISTRIBUTION AND ABUNDANCE: Re- corded throughout Lakes Malawi and Malombe, and the Upper Shire River. It is also frequently found in pools, and in streams and rivers flowing into the lake. It is widely distributed in Africa. It is a fish of shallow water, generally in the vicinity of vegetation. COMMERCIAL IMPORTANCE: Taken in ar- tisanal seines in shallow sheltered ar- eas. Widely cultured in fish farms.

Figure 9. Tilapia rendalli at Chemwezi Rock. Photo by Ad Konings.

31 Chapter 6 Mbuna

The mbuna, are to many people, the broad bodies, particularly between the quintessential Malawi cichlids, form- eyes. The mbuna dominate the rocky ing the bulk of the ornamental fish shores, in every way, but although the trade and being the most immediately great majority of species are strictly conspicuous fish seen by tourists confined to rocky shores, there are sev- swimming around the clear waters of eral species found over soft substrates, the rocky shores. The most comprehen- some of which are extremely abundant. sive ecological studies have focused These are generally much less studied almost exclusively on mbuna (Fryer, than their relatives on the rocky shores. 1959, Ribbink et al., 1983), leading, as I The Ps. livingstonii group are small have argued elsewhere (Turner, 1994d), brownish vertically barred mbuna to a body of theory about the evolu- which are common, often abundant tion of the Malawi cichlids which is over sandy areas. Ribbink et al. (1983) almost entirely based on circumstances consider them to be members of the Ps. relevant mainly to mbuna. zebra complex. The taxonomy of the The mbuna are a group of closely- group is confused. Ps. elegans is clearly related genera which are formally char- distinguished by all authors (Ribbink acterised by the abrupt transition in et al., 1983; Konings, 1990a), but the sta- size from the large flank scales to the tus of Ps. livingstonii, Ps. lanisticola, Ps. smaller chest scales and by the dark pursus, and Ps. 'livingstonii likoma' is border around the egg-spots on the much less clear. anal fin of males (Eccles & Trewavas, Burgess (1976) described Ps. 1989). An experienced observer can lanisticola, a snail-dwelling species, immediately assign almost any Ma- from Cape Maclear. Both Ribbink et al. lawian haplochromine as mbuna or (1983), and Konings (1990a, 1995b) re- non-mbuna, without using these fea- gard Ps. lanisticola as a junior synonym tures, but it is difficult to be precisely of Ps. livingstonii. I have not examined clear what features are used. Mbuna the types of any of these species, nor generally have strong colours, even have I surveyed the wide range of lo- where the females and juveniles are cations visited by Konings or Ribbink cryptic, they are usually strongly et al. Nevertheless, I consider that the pigmented in browns and blacks, and large drab species of open mud and few exhibit patterns of black or grey sand areas in the SE Arm and Lake on a silvery background which are so Malombe is clearly different from the frequent in non-mbuna (an exception small bright species which inhabits is the female of Ps. ' olive'). snail shells on clear water areas. I ten- Most species have short snouts, a tatively consider them to be distinct rather round-headed appearance and species. The small snail-dweller is cer-

32 tainly conspecific with Ps. lanisticola. or state of preservation of material. Konings (1995b) states that the types However, the matter must be consid- of Ps. livingstonii resemble the large ered unresolved. I am uncertain that southern form, and that this form facultative cleaning behaviour is suf- grades into the smaller brighter form. ficient to warrant specific status, as I think this question requires further Konings (1989, p.107) has illustrated investigation. such behaviour in I am not certain that the form I re- parallelus and I have observed it in M. gard as Ps. livingstonii is associated auratus. In all three cases, non-mbuna with mollusc shells, as has been as- species were cleaned. sumed by other authors. In areas Other mbuna species may occasion- where it dominates trawl catches, snail ally be found away from the rocky shells are rare. I have kept and bred coasts where they may be taken by specimens apparently identical to this seines. I have seen species. In the aquarium, it rarely en- in a seine net catch from the SE Arm ters snail shells, even as a juvenile, in and spp. in a fyke net set in marked contrast to snail-dwelling spe- the Upper Shire River. Some of the va- cies from , such as rieties of the Ps. tropheops complex and Lamprologus multifasciatus. The state- Ps. lucerna and related species are com- ments by Konings (1990a) and others mon on tiny patches of rocks on sandy that juvenile Ps. livingstonii inhabit shores. Mbuna may also be taken by snail shells may be due to confusion Chirimila nets fishing for utaka in the with Ps. lanisticola. water column above rocky areas. Spe- Stauffer (1991) described Ps. pursus, cies which frequently shoal to feed on a facultative cleaner fish, from the Cape plankton are especially likely to be Maclear area. He records it from shal- vulnerable to these gears. Prime can- low rocky areas. He was able to distin- didates would be the spe- guish it from the types of both Ps. cies, Petrotilapia and some of the livingstonii and Ps. lanisticola using Pseudotropheus zebra complex, such as multivariate morphometrics, but does Ps. callainos. In some areas, as around not provide a simple diagnosis suitable Nkhata Bay, they are actively fished for for use in the field. I have not been able using small-meshed gill nets and, in all to unambiguously identify this species areas, children will frequently try to in the samples I have examined. catch them with baited lines. Konings (1995b) considers this species also to be a junior synonym of Ps. Pseudotropheus livingstonii livingstonii. This is possible, since (Boulenger) multivariate techniques are very sen- colour photo on page 50 sitive to differences in body shape, but do not, of themselves prove specific DISTINGUISHING FEATURES: A small, status, since such differences could squat tan-coloured barred mbuna dis- easily be due to geographic variation tinguished from similar species by its or morphological changes associated generally drab colours and the lack of with size, age, reproductive condition a black margin to the anal fin.

33 COLOUR: Females and immatures tan, it made up almost 9% of the total sam- with 5 darker vertical bars under the ple biomass and frequently repre- dorsal fin and another on the caudal sented a higher proportion of the catch peduncle. Males generally more yel- weight than any other species. North lowish with blue iridescence on the of Boadzulu Is., it was often a domi- cheeks and operculum and several yel- nant species at 18-35m, and was occa- low egg-spots on the anal fin. sionally recorded as deep as 60m. It MAXIMUM SIZE: 14 cm TL (Konings was not found north of Monkey Bay, 1990a). where it was replaced by Ps. lanisticola. DIET: Konings (1990a) states that it I recorded it in Lake Malombe. feeds from the sediment, probably on COMMERCIAL IMPORTANCE: A significant diatoms. Specimens dissected by Grant component of shallow water trawl contained diatoms, cladocera, and catches in the SE Arm (3.4% of pair ephemeroptera, sometimes with a trawl catches). Also taken by seines in large amount of sand and detritus Lake Malombe and the SE Arm of Lake Grant & Turner (unpubl.). Malawi. A popular aquarium species, REPRODUCTION: Konings (1990a) re- which is easily bred in captivity. ports males and females leave the open NOTES: For discussion of taxonomic sandy areas and abandon their refuges uncertainties, see above. Fryer's (1959) in snail shells to migrate to rocky habi- reports of Ps. livingstonii apparently tats to breed. Konings (1995b), citing refer to Ps. 'zebra gold' (Ribbink et al. Van Duinen, notes that the species has 1983), a species of deep silt-covered a very short brooding period for an rocky habitats in the north of the lake. mbuna around 16 days. In my experi- ence, the fry are the smallest of any mbuna I have bred in captivity. Ko- Pseudotropheus elegans Trewavas nings (1995b) suggests that this may colour photo on page 50 be a result of the fry's sheltering in the snail shell occupied by the female, al- DISTINGUISHING FEATURES: A small, though it is not known if they actually squat tan-coloured barred mbuna dis- do this. tinguished from Ps. livingstonii by the DISTRIBUTION AND ABUNDANCE: Konings black lower margin of the anal fin and (1990a) states that Ps. livingstonii is from Ps. lanisticola by its fainter bar- found throughout Lake Malawi and ring, larger size, more elongate fins and Malombe, generally at around 25m absence of marked caudal fin stripes. depth, except in the SE Arm, where COLOUR: Females and immatures tan, they are found as shallow as 5m. with 5 darker vertical bars under the Ribbink et al. (1983) did not record the dorsal fin and another on the caudal species from farther north than the peduncle. The anal fin has a black Maleri Islands, although they report a lower margin. Males generally silvery similar form from Likoma Island. In with blue iridescence on the head, and the 1992 survey, it was the only spe- yellowish fins. Pelvic and anal fin mar- cies to be recorded from every sample gin black. Vertical bars faint or absent. from South of Boadzulu Island, where MAXIMUM SIZE: 16 cm TL (Konings

34 1990a). males. Fryer (1959) and Jackson (1961) DIET: Konings (1990a) states that it did not record this species at all. feeds from the sediment, picking out invertebrates. Ribbink et al. (1983) record stomach contents dominated by Pseudotropheus lanisticola Burgess benthic invertebrates, but also contain- colour photo on page 50 ing phytoplankton, zooplankton and loose epilithic algae. Specimens exam- DISTINGUISHING FEATURES: A small, ined by Grant & Turner (unpubl.) con- squat tan-coloured barred mbuna dis- tained diatoms, cladocera, copepods, tinguished from Ps. livingstonii by the cyanophytes, detritus and sand. black (in some populations orange) REPRODUCTION: Konings (1990a) re- lower margin of the anal fin and from ports males and females may migrate Ps. elegans by its stronger barring, to rocky habitats to breed. much smaller maximum size, shorter DISTRIBUTION AND ABUNDANCE: Konings fins and bright caudal fin stripes. (1990a) states that Ps. elegans is found COLOUR: Females and immatures tan, throughout Lake Malawi, generally at with 5 darker vertical bars under the depths greater than 10m. He illustrates dorsal fin and another on the caudal specimens from Senga Bay, Kande Is- peduncle. The anal fin has a black land (Konings 1990a) and Msuli Point lower margin. Caudal fin marked with (Konings 1995b). Ribbink et al. (1983) bold orange stripes on a whitish back- record the species from six locations ground. Males similar, but with gen- within the SE Arm, but also state that erally blue iridescence on the head. it is known from trawl samples in the Pelvics and anal fin margin black. One SW Arm, Nkhotakota and Bandawe or more prominent egg-spots on the and that the type is from Chilumba. In anal fin. the 1992 trawl survey it was abundant MAXIMUM SIZE: 8.5 cm TL. south of Monkey Bay at depths be- DIET: Specimens examined by Grant tween 18 and 35m. It was occasionally & Turner (unpubl.) contained diatoms, found as deep as 60m. It was rare north cladocera, copepods, and cyanophytes. of Monkey Bay. No sand or detritus recorded. COMMERCIAL IMPORTANCE: A significant REPRODUCTION: Konings (1990a) re- component of shallow water trawl ports males and females may migrate catches in the SE Arm (2.7% of pair to rocky habitats to breed. trawl catches). Also taken by seines in DISTRIBUTION AND ABUNDANCE: The Lake Malombe and the SE Arm of Lake type was collected from the Cape Malawi. Maclear areas. Konings' (1990) records NOTES: Neither Ribbink et al. (1983), of Ps. livingstonii from north of the SE nor Konings (1990a) have any record Arm may refer to this species. In the of females of this species. Konings 1992 trawl survey it was abundant (1995b) suggests that perhaps only north of Monkey Bay at 20-44m depth. males live out over the open sand. I It was frequently seen in trawl catches have found that females commonly in which large numbers of snail shells occur in the same trawl samples as (Lanistes) were collected and was often

35 found inside the shells. numerous. COMMERCIAL IMPORTANCE: Probably not COMMERCIAL IMPORTANCE: Perhaps a an important trawl species, as commer- minor component of trawl and seine cial trawlers tend to avoid areas with catches. high concentrations of snail shells NOTES: The type was labelled as hav- which may damage their nets and the ing been collected in Lake Tanganyika. fish caught. The species was subsequently de- NOTES: for discussion of taxonomic scribed as Christyella nyasana by Trewa- uncertainties, see above. vas.

Gephyrochromis moorii Boulenger Cyathochromis obliquidens colour photo on page 50 Trewavas colour photo on page 50 DISTINGUISHING FEATURES: A slender purple and orange species with elon- DISTINGUISHING FEATURES: A deep-bod- gated fins and a strongly forked tail. ied brownish species with a prominent The northern species Pseudotropheus sp. black submarginal band in the dorsal 'acei' is similar, but is either bluish with fin of males. Females have more, thin- a yellow tail or black with a white tail. ner bars than species of the Ps. living- COLOUR: Females and immatures or- stonii group. The mouth is not so ange-brown, white ventrally. Mature ventrally positioned as that of the Ps. males a rich purple-blue on the flanks tropheops complex, which consequently with one or two whitish-yellow egg- have a more steeply-angled snout pro- spots on the anal fin. file. MAXIMUM SIZE: 13 cm TL (Konings COLOUR: Females and immatures 1990a). pale brownish with 10-11 thin dark DIET: Feeds on sediment (Konings brown vertical bars, 1-2 horizontal 1990a). bars. Males greenish-brown, with pur- REPRODUCTION: Konings (1990a) re- plish iridescence. Yellowish dorsally ports males do not hold fixed territo- and on chest. Dorsal fin yellow with a ries. prominent black submarginal band DISTRIBUTION AND ABUNDANCE: Konings and 2-3 large yellow egg-spots on the (1990a) states that it is found in the anal fin. southern half of the lake, and is com- MAXIMUM SIZE: 13 cm TL (Konings mon at Senga Bay. He records it from 1990a). 6-25m depth. In the 1992 trawl survey DIET: Feeds by brushing algae from it was recorded from two stations at rock surfaces or leaves of macrophytes 20-28m on the east coast of the SE Arm, (Fryer, 1959). Ribbink et al. report stom- at approximately the same latitudes as ach contents containing algae, plank- Monkey Bay to Zimbawe Rocks. I also ton, insect larvae, and benthic crusta- collected the species from trawls at ceans. This species is commonly seen 25m off the Maleri Islands and 37m in feeding from the leaves of Potamogeton, Domira Bay. It was occasionally quite in small groups in the company of

36 Protomelas spp. and Hemitilapia oxy- rhynchus and immature Oreochromis. REPRODUCTION: Konings (1990a) reports males dig pits among vegeta- tion or under rocks. DISTRIBUTION AND ABUNDANCE: A wide- spread and abundant species of shal- low vegetated areas. Jackson reports that it is found in areas of mixed sand and rock, where it is often the most abundant fish. Ribbink et al. (1983) re- corded the species at most of their sur- vey areas and also report it from Lake Malombe and the Upper Shire. Ko- nings (1990a) states that it is found throughout Lake Malawi. I did not record it in Lake Malombe, nor in the 1992 trawl survey, but it was frequent in experimental fyke net catches in the Upper Shire and occasionally taken in trawls in the SE Arm and Domira Bay. COMMERCIAL IMPORTANCE: A minor component of seine catches.

37 Chapter 7 Ncheni and Ndunduma

Ncheni and Ndunduma are silvery Diplotaxodon, and Copadichromis) al- fishes which feed on zooplankton or though smaller fishes often take zoo- fish. Rhamphochromis (Ncheni) are plankton (Allison et al., 1995). Most of streamlined pursuit hunters found in the described species seem to be absent all habitats, from shallow swampy ar- from the trawlable areas and also from eas, such as Lake Malombe, to the the pelagic zone. Different species ap- pelagic zone in the middle of the lake. pear to have different habitat prefer- They are found in bottom trawl catches ences, some preferring inshore waters, over the full range of depths fished to while others inhabit the surface of open date. Diplotaxodon (Ndunduma) are waters, and some appear to be largely principally deep-water piscivores and demersal. Large numbers of small zooplanktivores with upwardly-an- Rhamphochromis are caught in shallow gled mouths. Diplotaxodon limnothrissa waters by kambuzi seines in Lake Ma- and its relatives are truly pelagic zoo- lawi. It is not known if all of the off- plankton feeders, which take the place shore or deep water species have in- of the utaka (Copadichromis) in offshore shore-living young or if these represent waters. Pallidochromis (monotypic to different species. Aquarium observa- date) is a deep-water benthic-feeding tions indicate that some species are piscivore. capable of spawning in midwater These genera, comprising mainly (Konings, pers. comm.). deep-water species were very poorly Rhamphochromis are very important represented in the earlier collections, commercially, and are caught by hand and their full diversity could not be ap- lines, chirimila and gill nets as well as preciated until the advent of midwater comprising a substantial proportion of trawling in the 1970s. The Overseas all trawl catches; 12.4% of midwater Development Administration (UK) has trawl bycatch, 8.3% of demersal stern recently approved a project to study trawl, and 15.1% of pair trawl catches, the biology of these species, so I will in 1990-91. They are known as ncheni, present only a brief account of those but have not been independently re- species which can be readily distin- corded in the Malawi Government tra- guished or which are likely to be found ditional fisheries catch assessment in the demersal zone in the southern forms. Apart from large adults caught lake. by the midwater trawl, they are not The large number of Rhamphochromis recorded separately on commercial re- species (12-20 spp.) are elongate turns. Some of the smaller species midwater-hunting predators, which make good aquarium fishes. feed principally on small midwater Diplotaxodon may account for a larger fishes (such as Engraulicypris sardella, proportion of the fish biomass in Lake

38 Malawi than any other . In addi- was described from a single specimen tion to the four described species, there collected at a depth of 90m off the are at least another 9 undescribed. Be- Nankumba Peninsula, at the north of fore the 1990s, only 3 species were for- SE Arm. It is similar in body propor- mally described and none of the pre- tions to D. argenteus. It is apparently a vious field studies adequately distin- ripe male, of 14cm SL, which is black guished between the described and with a broad white dorsal fin margin. undescribed forms. All species are sil- Individuals with similar morphology very, with steeply-angled mouths; were collected from 60m at Domira Bay many are pelagic or part of the deep and 68m near Maleri Island. I have water benthic community. Diplotaxodon examined several other specimens species are marketed as 'ndunduma', which could not be unambiguously but in trawl catch returns they are ei- assigned to any of the above species, ther not distinguished from chisa- collected by Iles from unknown loca- wasawa (Lethrinops and other benthic tions or by the ODA/SADC project species) or are recorded as utaka (Co- from near Nkhata Bay. padichromis spp.). Around the Nan- I described Pallidochromis tokolosh kumba peninsula, where they are from deep water trawls in the south of taken by hand line and chirimila, they the lake (Turner, 1995c). Konings (1995) are known as 'Jamesons'. In addition has photographed a similar form to the well-defined species from the which may be a second species in this trawlable areas of the SE Arm, a num- genus. ber of other Diplotaxodon have been found in deep waters. A large-eyed elongate species (D. 'big-eye offshore' Rhamphochromis esox (Boulenger) Turner, 1994b) reaching 15 cm or more colour photo on page 50 is common in deep waters in the mid- Rhamphochromis leptosoma Regan dle of the lake. This may be the species known by Iles as D. 'micrentodon'. The These two species are fairly easily ODA/ SADC project collected another distinguished from other Rhamphochro- large-eyed, deep-bodied species. Ripe mis by their elongate rather cylindri- males were dark, with a coppery sheen cal body form. They generally have a (D. 'big-eye copper' Turner, 1994b). horizontal dark stripe and rather dark Diplotaxodon ecclesi Burgess & Axelrod greyish flanks. R. esox has a shorter

Figure 10. Rhamphochromis leptosoma. Holotype.

39 decurved snout and a more prominent other Rhamphochromis and their mor- premaxillary pedicel that R. leptosoma, phology suggests that they are benthic but it is not clear if this represents in- forms. R. brevis is deeper bodied, and dividual variability or a species-spe- R. macrophthalmus has a relatively cific trait. Specimens of R. esox and R. larger eye than R. woodi. The types of leptosoma are known from demersal R. brevis were collected from some- trawls in the SE Arm and from experi- where between Mangochi and Fort mental midwater trawls near Mbenji Maguire. Neither R. brevis, nor R. Is. and R. esox has also been trawled macrophthalmus was found in trawl from just south of Karonga. R. esox at- catches in the SE Arm in 1990-92, but a tains at least 36cm TL and R. leptosoma few specimens similar to R. woodi were 33cm SL. collected from the commercial mid- water trawl in the SE Arm and from Rhamphochromis macrophthalmus experimental trawls in shallow water Regan near Boadzulu Is. and were recorded Rhamphochromis woodi Trewavas under the name Rhamphochromis 'big- Rhamphochromis brevis Regan tooth brown'. These have shorter pre- Rhamphochromis 'bigtooth brown' maxillary pedicels (20-24% HL) than R. woodi (26-28% HL). These species are These four species share the same probably confined to inshore waters. general body shape, having a relatively The largest known specimens are R. arched back and flat ventral profile. macrophthalmus: 24cm TL; R. woodi: 22 They are less streamlined than most cm TL; R. brevis: 42 cm TL. Other speci-

Figure 11. Rhamphochromis macrophthalmus. Lectotype.

Figure 12. Rhamphochromis woodi. Lectotype.

40 Figure 13. Rhamphochromis brevis. Lectotype.

Figure 14. Rhamphochromis 'big-tooth brown'. mens, which could not be assigned to ceratobranchial gillrakers. Ripe males any of these species, but with a gener- up to 44 cm TL, single mature female ally similar body shape include a 43 28 cm TL. Collected from demersal cm TL specimen gillnetted at 100m trawls at 50m off Chirombo Bay in the depth off Chisumulu Is. by the ODA/ SE Arm, and experimental midwater SADC project, a 33cm TL specimen fishing at 30m depth over a lake bed collected from Vua by Christy and er- 104m deep near Domwe, and at 80m roneously figured by Eccles & Trewa- over a 104m deep bottom at Chipoka vas as R. ferox, and a very laterally com- in the SW Arm. pressed specimen of 22 cm TL collected by Fülleborn from the far north of the Tanzanian coast of the lake. Rhamphochromis 'big mouth'

A large species similar to the above, Rhamphochromis 'long snout' but with a less ventrally-angled snout colour photo on page 50 and 15-16 ceratobranchial gillrakers. They have very long premaxillary A large species with a long, ven- pedicels (26-29% HL). This species is trally-angled snout, large mouth and known only from the pelagic zone, teeth. Body shape similar to previous having been trawled or gillnetted at species, but with more rounded ven- depths of 30-56m over lake bottoms of tral profile. Generally brownish. 11-14 78-400m at Monkey Bay, Domwe,

41 Figure 15. Rhamphochromis 'big-mouth'.

Makanjila Point, Chipoka, Sungu (near cm TL. Two very similar specimens of Nkhotakota) and Chisi (near Nkhata around 32 cm TL were collected by the Bay). The largest known specimens are ODA/SADC project from a midwater around 40 cm TL. Other specimens trawl over a lake-bed depth of 99m from the same locations have similar near Domwe Is. and from a trawl near body proportions, but have much Chisi Point. These appear to be ripe shorter premaxillary pedicels (22-23% males, and are uniformly dark metal- HL). lic grey, with a slight orange tint on the pelvic and anal fins. Most of the other specimens included in the description Rhamphochromis ferox Regan of R. ferox by Eccles and Trewavas seem not to be conspecific with either of the This species was described by Regan types. from two co-types, which are not con- specific. One specimen (to be desig- nated the lectotype) has a slender snout Rhamphochromis 'shire ferox' and short jaws, thin lips and long, slen- colour photo on page 51 der widely-spaced teeth. A half-skin found in the same jar as the type of R. The other type of R. ferox was col- longiceps is considered to be conspe- lected from the Upper Shire River. It is cific. The largest known specimen is 20 around 23.5 cm TL. It is a heavily-built

Figure 16. Rhamphochromis ferox. Lectotype.

42 fish with thick lips and short, deeply- est known specimen is 44 cm TL. It embedded teeth. No other specimens seems to be quite common in artisanal have been collected, but similar-look- catches and midwater trawl catches ing fishes have been seen in seine from deep water in the south of the catches in Lake Malombe. It is almost lake. certainly an inshore species. Rhamphochromis 'short-tooth brown'

This is a small, streamlined brown- ish species known from commercial

Figure 17. Rhamphochromis lucius. Lectotype.

Figure 18. Rhamphochromis l'short-tooth brown'. (Paralectotype of R. lucius.)

Rhamphochromis lucius Ahl midwater trawl catches in the SE Arm. The second type specimen of R. lucius One of the types (to be designated appears to be this species. The largest as the lectotype) is a large (35 cm TL) known individual is 20 cm TL, but all streamlined specimen with a huge known specimens were sexually im- head. Similar specimens were collected mature. It has a rather cylindrical body, by Christy and Trewavas from un- an upwardly-angled mouth and rela- known locations, and others were tively short straight teeth. taken from commercial midwater trawls in the SE Arm in 1991. The larg-

43 Figure 19. Rhamphochromis longiceps. This specimen is figured in Eccles & Trewavas 1989.

Rhamphochromis longiceps Rhamphochromis 'long fin yellow' (Günther) colour photo on page 51

This species is unusual among Rham- Known from 4 ripe males from phochromis in its closely packed small depths of 24-40m at Chilinda and Palm teeth. It has a small mouth, large oper- Beach, and 2 from near Chitande Is. in culum and deep cheek. The type, a the north of lake, the largest 25 cm TL. half-skin, was collected from Songwe. It is a relatively deep-bodied species Some specimens in the Christy collec- with a large mouth, prominent pre- tion are certainly conspecific, includ- maxillary pedicel and large widely- ing the specimen figured by Eccles & spaced teeth. Ripe males are brownish Trewavas which is 23.5 cm TL and is a dorsally, but the whole of the lower half brooding female. of the body is bright yellow, including the anal fin and the very long pelvic fins. Females and immature males Rhamphochromis cf. longiceps have not been identified. colour photo on page 50

Small specimens collected offshore Rhamphochromis 'kolowilo' off Karonga, Charo, and Nkhotakota colour photo on page 51 by the ODA/SADC project were ini- tially thought to be R. longiceps, but A slender form with large mouth, they have a more slender snout, shal- large, widely-spaced teeth and a lower cheek and smaller operculum. prominent premaxillary pedicel. Im- This form is also known from demer- matures and females are brownish and sal trawls at 5-74m in the SE Arm, attain at least 27.5 cm TL. Some large, Maleris and Domira Bay. The largest deeper-bodied ripe males (up to 35 cm known specimens are 21.5 cm TL. TL), which may be this species have been taken from commercial midwater trawls. They are yellowish-brown dorsally, bright yellow on the belly, pelvic and anal fins.

44 Diplotaxodon limnothrissa Turner seasonal peak. Thompson et al. (1995a) colour photo on page 51 recorded low breeding activity be- tween May and September. Size at DISTINGUISHING FEATURES: Silvery with maturity is around 14 cm (Table 1, see no dark markings. More elongate body also Thompson et al. 1995a). Ripe and (maximum depth 26-30% SL) than D. brooding adults taken from deep wa- 'intermediate' and Copadichromis vir- ter (50-100m), and they may spawn in ginalis. Smaller head, eye and mouth midwater. Average fecundity is around than D. argenteus and relatives. Less 15 eggs (Thompson et al., 1995a). The pointed head and more gill rakers (20- fry are brooded to a large size — about 26) than Nyassachromis eucinostomus 3 cm. complex. DISTRIBUTION AND ABUNDANCE: This COLOUR: Females and non-breeding species probably has a far higher bio- males silvery, darker dorsally, with mass than any other cichlid in Lake purplish sheen. Ripe males dark grey, Malawi. Menz (1995) estimated a total silvery ventrally. Dorsal fin white or biomass of 87,000t in the pelagic zone. yellow dorsal fin, with a blaze of white Assuming an average adult weight of of yellow on dorsal surface of head in- 60g, this is approximately 1.45 billion cluding snout. Pelvics and anal white fish, excluding juveniles. In the 1992 or yellow. 1-2 large pale yellow egg- survey, it was found at depths of 24- spots on anal fin. 102m throughout the SE Arm. Absent MAXIMUM SIZE: 19 cm TL. from shallow inshore waters of 20m or DIET: Plankton, mainly less and never seen by SCUBA divers. (Tropodiaptomus, Mesocyclops, Diaphano- Thompson et al. (1995b) recorded the soma), but occasionally also chaoborid species throughout the pelagic zone of larvae and pupae, Engraulicypris fry the lake from the surface to a depth of and filamentous diatoms (Aulacoseira) at least 220m. Offshore, most individu- (Allison et al., 1995). als were either small fry or larger than REPRODUCTION: Seems to be a continu- 11 cm TL, but intermediate sizes are ous breeder, although there may be a known from trawls in the SE Arm. COMMERCIAL IMPOR- TANCE: In 1990-91, more than 700 tonnes Total Immature Active Ripe/Spent Active Ripe Length Unsexed Females Females Males Males were caught annually in the SE Arm. It com- 8-11.9 7 3 — 2 — prised 53% of the 12-12.9 13 3 — 2 — 38mm midwater trawl 13-13.9 8 5 — 2 — bycatch, 18% of the 14-14.9 1 1 — 3 1 deep-water bottom 15-15.9 1 13 1 7 1 trawl catch, and 5% of 16-16.9 2 11 1 6 2 the pair trawl catch. It 17-17.9 — 2 1 — — has also been seen in catches of chirimila Table 1: Size at maturity, Diplotaxodon limnothrissa.

45 Figure 20. Diplotaxodon 'intermediate'; South-East Arm. nets. It is probably the most commer- Diplotaxodon argenteus Trewavas cially important haplochromine cichlid colour photo on page 51 in the SE Arm. DISTINGUISHING FEATURES: A silvery species with no dark spots or stripes. Diplotaxodon 'intermediate' Eye shorter than snout. It has a larger mouth (lower jaw 14-16% SL) than D. DISTINGUISHING FEATURES: Similar to D. limnothrissa and D. 'intermediate' and limnothrissa, but deeper-bodied (maxi- a more slender body than D. 'deep', D. mum depth 30-36% SL). 'brevimaxillaris', and D. greenwoodi. It COLOUR: Countershaded and silvery. differs from D. 'similis' in its larger eye, Generally darker than D. limnothrissa. wider interorbital, higher number of Sexually active males (ripe?) darker, gill rakers (21-27 ceratobranchials), with dark fins. shorter pectoral fin (not reaching base MAXIMUM SIZE: 20 cm TL. of first anal spine), more upwardly-di- DISTRIBUTION AND ABUNDANCE: Rare. rected mouth, and male breeding col- Positively identified only from the SE ours. Arm, north of Boadzulu Island. COLOUR: Females and immature COMMERCIAL IMPORTANCE: Occasion- males silvery, darker dorsally. Ripe ally taken by midwater trawl. males silvery, with dusky dorsum and NOTES: Ripe males, which may be this head. Black eye, snout, and dorsal, species, collected by T. D. Iles in the anal, caudal, and pelvic fins. 1-2 large 1950s were found in the Monkey Bay yellow egg-spots on anal. Fisheries Research Station collection MAXIMUM SIZE: 20 cm TL. under the name 'Diplotaxodon holo- REPRODUCTION: Appears to be a mid- chromis'. They were not described and water spawner, with a possible breed- no collecting location is given. They ing peak in the cold season. have a uniformly dark colour (black?) DISTRIBUTION AND ABUNDANCE: Com- with a pale dorsal fin (white?). mon. In the 1992 survey, taken from trawls at 34-114m depth throughout

46 the SE Arm. Ad hoc trawls at the sur- sardella, (Allison et al., 1995). face also caught this species. Not posi- DISTRIBUTION AND ABUNDANCE: In the tively identified from outside the SE 1992 trawl survey, taken at 90-128m. Arm. There appears to be no inshore Occasionally collected in much shal- phase in its life cycle, but it is not a com- lower water and also caught in open mon species in the true pelagic zone. water by the ODA/SADC Project COMMERCIAL IMPORTANCE: 4% of mid- (Turner, 1994b; Menz, 1995). water trawl bycatch. Also taken in COMMERCIAL IMPORTANCE: Occasion- deep-water bottom trawls, chirimila ally found in midwater trawl and and hand lines. deep-water bottom trawl catches. Also NOTES: The ODA/SADC project taken by hand lines. collected a number of small specimens NOTES: Most of the 'D. argenteus' with a similar male breeding colour taken by the ODA/SADC project's off- and similar meristics. These may be shore surveys were this species conspecific. (Turner, 1994b).

Diplotaxodon 'similis' Diplotaxodon 'deep' colour photo on page 51 colour photo on page 52

DISTINGUISHING FEATURES: A silvery DISTINGUISHING FEATURES: A silvery species with no dark spots or stripes. species lacking dark spots or stripes. Eye shorter than snout. It has a large Eye shorter than snout, at least in large mouth (lower jaw 15-17% SL), promi- specimens (>130 cm SL). Deep-bodied nent teeth and a less upwardly-di- (depth 31-37% SL). The mouth is much rected gape than other Diplotaxodon. It less steeply-angled than those of D. has a more slender body (max. depth 'brevimaxillaris' or D. greenwoodi. 31-33% SL) than D. 'deep', D. 'brevi- COLOUR: Silvery, countershaded. maxillaris' and D. greenwoodi. It has Generally more yellowish-brown than fewer gillrakers (16-19, occasionally 20- other Diplotaxodon species. Male col- 21) than D. argenteus, as well as a larger ours not known. eye, narrower interorbital, and longer MAXIMUM SIZE: 20 cm TL. pectoral fin (reaching beyond first anal DISTRIBUTION AND ABUNDANCE: Rare. spine base). The two also differ in male Mostly taken from deep waters of 50m breeding colours. It has a smaller or more. Not seen in samples taken by mouth and teeth than Pallidochromis the ODA/SADC project, and probably tokolosh. not a pelagic species. COLOUR: Females and immature COMMERCIAL IMPORTANCE: Often taken males silvery, countershaded. Ripe in small numbers in deep-water bot- males silvery, with a dark grey back tom trawls. and head. The dorsal fin is white and the pelvics black. One or two large egg- Diplotaxodon 'brevimaxillaris' spots on anal fin. colour photo on page 52 MAXIMUM SIZE: 25 cm TL. DIET: Juvenile usipa, Engraulicypris DISTINGUISHING FEATURES: Silvery with

47 no dark spots or stripes. Deeper-bod- REPRODUCTION: Several large males in ied than most congeneric species, with breeding dress were taken in a bottom a very steeply-angled mouth. D. green- trawl catch at a depth of 100m off woodi has a larger mouth and more Domwe Island, suggesting that the strongly projecting lower jaw. species is a colonial demersal spawner. COLOUR: Silvery, countershaded. DISTRIBUTION AND ABUNDANCE: Found MAXIMUM SIZE: 25 cm TL. in a single sample from 102m depth in DISTRIBUTION AND ABUNDANCE: Uncom- the 1992 trawl survey, but often taken mon and not taken in the 1992 trawl in small numbers at 50m depth or more survey. Often seen in other trawl in the SE Arm. Also occurs near Nkhata catches from 50-128m depth, but never Bay. in large numbers. COMMERCIAL IMPORTANCE: Frequently COMMERCIAL IMPORTANCE: Frequently taken in small numbers by deep-wa- taken in small numbers by deep-wa- ter bottom trawl fishery, and to a lesser ter bottom trawl fishery, and to a lesser extent the midwater trawl fishery. Also extent the midwater trawl fishery. Pos- exploited by artisanal fisheries, possi- sibly also exploited by hand line and bly using hand lines and deep-set gill deep-set gill net. nets.

Diplotaxodon greenwoodi Diplotaxodon 'macrops' Stauffer & McKaye colour photo on page 52 colour photo on page 52 DISTINGUISHING FEATURES: A silvery DISTINGUISHING FEATURES: A silvery species with small, closely-packed species lacking dark spots or stripes, teeth and lacking dark spots or stripes. with closely-packed teeth. It is the larg- Small, with eye diameter greater than est Diplotaxodon species and is charac- snout length. Distinguished from D. terised by the deep, laterally-com- ''white belly' by smaller lower jaw (37- pressed body, relatively small eye, the 41% HL). Cheek depth (7-9% SL) less large upwardly-angled mouth and than D. 'copper'. Both of these species strongly projecting lower jaw. have different male breeding colours. COLOUR: Females and immature COLOUR: Ripe males are black, with a males are silvery, and countershaded. broad white margin to the dorsal fin. Ripe males are silvery on the flanks, Females and immature males are sil- but have a dark head, belly, dorsum very, and countershaded. and fins. They have a single row of MAXIMUM SIZE: 12 cm TL. around 5 egg-spots on the anal fin. DIET: Zooplankton. MAXIMUM SIZE: 30 cm TL. DISTRIBUTION AND ABUNDANCE: Abun- DIET: Piscivore. Possibly feeds on lar- dant at depths of 90m or more in the vae and eggs from the mouths of SE Arm. brooding females (Stauffer & McKaye, COMMERCIAL IMPORTANCE: Not known 1986), but known to take smaller to be exploited. cichlids (Allison et al., 1995).

48 Ripe male O. karongae (top) and O. lidole (two lower fishes), Thumbi Island West.

From top to bottom: , O. karongae, and O. lidole (immatures) from Cape Maclear. O. karongae, xanthochromatic form, SE Arm.

O. squamipinnis, male, Karonga (photo by Dennis Tweddle).

Oreochromis shiranus, female, South-East Arm. O. karongae underwater at Thumbi West Island (photo by Eva Hert).

49 Pseudotropheus livingstonii, Lake Malombe. Pseudotropheus elegans, male, South-East Arm.

Pseudotropheus lanisticola, Domira Bay.

Cyathochromis obliquidens, South-East Arm.

Gephyrochromis moori, male, Domira Bay. Rhamphochromis 'long snout', South-East Arm.

Rhamphochromis esox (top) and Rhamphochromis cf. longiceps, both South-East Arm.

50 Rhamphochromis 'shire ferox', Lake Malombe.

Rhamphochromis 'long fin yellow', South-East Arm.

Rhamphochromis 'kolowilo', Maleri Island. Diplotaxodon limnothrissa, male, SE Arm.

Diplotaxodon argenteus, South-East Arm. Diplotaxodon 'similis', South-East Arm.

51 Diplotaxodon 'deep', South-East Arm.

Diplotaxodon 'brevimaxillaris', South-East Arm.

Diplotaxodon greenwoodi, male, SE Arm.

Diplotaxodon 'macrops', male, SE Arm. Diplotaxodon 'white belly', males, South-East Arm.

Pallidochromis tokolosh, Narungu, eastern shore.

52 Above: Copadichromis virginalis, male, long- finned form, Domira Bay.

Copadichromis virginalis, short-finned form, male (top) and female, off Monkey Bay. Copadichromis 'chrysonotus black', male (top) and female, Lake Malombe.

Copadichromis quadrimaculatus, male, off Chirombo Bay. C. quadrimaculatus, female, off Chirombo Bay.

53 Copadichromis pleurostigma, female, Maleri Is. Nyassachromis microcephalus, male, South-East Arm.

Nyassachromis 'eucinostomus black', male, South-East Arm.

Nyassachromis 'eucinostomus yellow', male, SE Arm.

Nyassachromis 'argyrosoma blue', male (top) and female, off Maleri Island. N. 'argyrosoma blue', male (ripening), South-East Arm.

54 Lethrinops 'micrentodon Makokola', male, South-East Arm.

Lethrinops 'yellow chin', male, South-East Lethrinops microdon, male (top) and female, South-East Arm. Arm.

Lethrinops 'oliveri', male South-East Arm. Lethrinops stridei, male, South-East Arm.

Lethrinops 'dark', male, South-East Arm. Lethrinops 'yellow tail', male, South-East Arm.

55 Lethrinops 'yellow', male, SE Arm.

Lethrinops 'black chin', male, South-East Arm.

Lethrinops 'deep water altus', male, South-East Arm.

Lethrinops altus, male, South-East Arm.

Lethrinops 'matumbae', male (top) and female, SE Arm.

Lethrinops 'blue-orange', male, South-East Arm. Lethrinops christyi, male, South-East Arm.

56 Lethrinops longipinnis, male, Domira Bay at 37m depth. Lethrinops polli, male, Domira Bay.

Lethrinops longipinnis, female,SE Arm.

Lethrinops gossei, male (top) and female, South-East Arm. L. 'deep-water albus', female, SE Arm.

Lethrinops 'deep-water albus', male, South-East Arm.

57 Lethrinops longipinnis, male, South-East Arm from a depth of 90 m.

Lethrinops mylodon, female, Domira Bay.

Lethrinops mylodon, male, South-East Arm.

Lethrinops macracanthus, South-East Arm. Lethrinops longimanus, males, Domira Bay.

58 Lethrinops macrochir, male, Lake Malombe.

L. macrochir, female, Lake Malombe.

Lethrinops cf. parvidens, male (top) and female, SE Arm. Lethrinops 'macrostoma', female, SE Arm.

Lethrinops 'pink head', male, South-East Arm. Lethrinops cf. furcifer, male, South-East Arm.

59 Tramitichromis lituris, male, South-East Arm.

Taeniolethrinops praeorbitalis, female, SE Arm. Tramitichromis lituris, female, South-East Arm.

Taeniolethrinops praeorbitalis, male, South-East Arm.

60 Alticorpus mentale, male, South-East Arm.

Alticorpus mentale, female, South-East Arm.

Alticorpus macrocleithrum, male, Domira Bay.

Alticorpus pectinatum (top), A. 'deep' (centre), Taeniolethrinops furcicauda, female, SE Arm. and A. 'geoffreyi', males, South-East Arm.

61 Aulonocara rostratum, female, South-East Arm. Aulonocara rostratum, male, South-East Arm.

Aulonocara cf. macrochir, male, South-East Arm. Aulonocara guentheri, female, Monkey Bay.

Aulonocara 'gold', male, off Monkey Bay.

62 Aulonocara 'copper', male, Domira Bay.

Aulonocara 'deep', male (top) and female, off Monkey Aulonocara 'blue-orange', male, SE Arm. Bay.

Aulonocara 'pyramid', male, South-East Arm.

Aulonocara 'orange', male, South-East Arm. Aulonocara 'orange', female, South-East Arm.

63 Aulonocara 'yellow', male, South-East Arm. Aulonocara 'dark stripe', male, South-East Arm.

Aulonocara 'deep yellow', male, off Maleri Island. Aulonocara 'green', male, South-East Arm.

Aulonocara 'minutus', male, South-East Arm.

Aulonocara 'stonemani', male, South-East Arm.

64 Diplotaxodon 'white belly' COLOUR: Silvery, pale with little evi- colour photo on page 52 dence of countershading. Ripe male colours unknown, although many DISTINGUISHING FEATURES: A silvery specimens have been examined. species with small, closely-packed MAXIMUM SIZE: 35 cm TL. teeth and lacking dark spots or stripes. DIET: Benthic cichlids: Aulonocara, Small, with eye diameter greater than Lethrinops, etc. snout length. Distinguished from D. DISTRIBUTION AND ABUNDANCE: Fre- 'macrops' by the larger mouth (lower quent, but in small numbers. Found at jaw 42-46% HL) and from D. 'copper' depths of 60-126m in the 1992 survey, by the longer pectoral fin (34-41% SL). but also known from as shallow as Both of these species have different 50m. male breeding colours. COMMERCIAL IMPORTANCE: Occasion- COLOUR: Ripe males dark dorsally, sil- ally taken by deep-water bottom ver on flank and belly. Dorsal fin with trawls. a broad white border. Pelvic and anal NOTES: Recently described by the fins dark. 1-2 large pale egg-spots on author (Turner, 1994c), but previously anal. Females and immature males are illustrated by Smith (1991) as 'Diplo- silvery, and countershaded. taxodon big head'. In unpublished field MAXIMUM SIZE: 13 cm TL. notes, known by Iles as 'Rhamphochro- DIET: Zooplankton. mis pallidus' and by Eccles as 'Haplo- DISTRIBUTION AND ABUNDANCE: Abun- chromis leucogaster'. dant at depths of 100m and deeper in the SE Arm. COMMERCIAL IMPORTANCE: Not known to be exploited.

Pallidochromis tokolosh Turner colour photo on page 52

DISTINGUISHING FEATURES: This looks like a deep-sea fish. It is rather flaccid, non-streamlined, pale, big-eyed, and big-mouthed. It is large, fairly elongate and has a prominent mental process. The teeth are larger and more widely- spaced than those of Diplotaxodon spp. but it lacks the enlarged teeth in the second row of the lower jaw of Rham- phochromis. It looks rather similar to Stigmatochromis 'guttatus', but has larger mouth and lacks spots and ver- tical bars.

65 Chapter 8 Utaka

'Utaka' is the name applied by Ma- ratio which Eccles & Trewavas (1989) lawian fishermen to the small shoaling also used in the diagnosis of Copadi- fish caught by day by chirimila nets chromis and Nyassachromis. Konings, operating near the surface of the wa- like Eccles & Trewavas, allows himself ter on steep rocky coasts or on sub- exceptions to his own rules Copadi- merged rock reefs ('chirundu', plural chromis likomae constructs a sand-cas- 'virundu'). Eccles & Trewavas assigned tle, but has flank spots and is retained those species with protrusible mouths in Copadichromis, as is C. virginalis, to the genus Copadichromis, but also which lacks spots (at least as an adult), included C. pleurostigma, which does but does not build a nest on open sand. not have a protrusible mouth, because After some resistance, I have decided of its resemblance to other deep-bod- to adopt Konings' classification, al- ied, spotted species in the genus. Some though I think it is likely that some re- of the species they included in the ge- vision will be required, for example, C. nus are slender-bodied, horizontally- virginalis might be best placed in a new striped or plain coloured sand-dwell- genus. ing species. A number of very similar Most utaka feed largely on zoo- species without protrusible mouths plankton, although Engraulicypris fry were placed in Nyassachromis along are also taken. Although sometimes with deeper-bodied horizontally- believed to be pelagic, they generally striped species. seem to be closely associated with the Konings (1990b, 1995b) has proposed bottom, feeding in large shoals one or an alternative scheme, ignoring the cri- two metres above the substrate. All terion of the protrusible mouth entirely species are strongly sexually dimor- and regarding all species which build phic during the breeding season: males sand-castle bowers as Nyassachromis. are much larger and more colourful. This includes most of the horizontally- Fry are released in shallow water. Al- striped or unmarked species, such as though generally termed utaka there N. boadzulu, N. eucinostomus, and N. are a variety of names for different spe- prostoma. Most of these species have a cies caught by chirimila fishermen. relatively benthic lifestyle and are Those caught in beach seines or in Lake mostly confined to sandy bottoms al- Malombe are generally recorded as though they often aggregate at virun- kambuzi in fisheries statistics, while du and more up into the water column those caught by pair trawlers are not to feed on plankton blooms. Stauffer regularly distinguished from benthic et al. (1993), rejecting this proposal, chisawasawa species. point out that Konings has not consid- In addition to the species discussed ered the relative intestine: body length in detail below, many others are closely

66 associated with rocky shores: these in- N. conophorus. They differ in size, gill- clude , C. jacksoni, raker count and in the form of bowers C. 'chrysonotus blue', C. azureus, C. built by sexually active males. Stauffer verduyni, C. mbenjii, and a number of et al. consider that N. eucinostomus undescribed forms. These species are should be regarded as a 'museum spe- particularly abundant at areas of up- cies' as it was described from a single welling in the vicinity of islands or juvenile specimen from an unknown submerged rocky reefs (chirundu, plu- locality and therefore cannot be iden- ral = virundu). In these areas they are tified in the field. Konings (1995b) pre- exploited by chirimila nets and hand fers to retain N. eucinostomus, and re- lines. Most of these rock associated gard Stauffer et al.'s names as junior species do not appear to cross soft-bot- synonyms. There are merits to both toms or move in open water, and are courses, I have observed what appears never caught in trawl catches. These to be two species in this group on the species appear to have restricted dis- same arena, but on the other hand, tributions and are likely to be vulner- Stauffer et al.'s description are very dif- able to local overfishing. Breeding ap- ficult to map onto specimens taken pears to be seasonal, with a peak at the from areas other than the Nankumba beginning of the hot season, around Peninsula. September to October. Males generally Nyassachromis spp. are more com- build nests on the sand near to rocks monly caught by beach seines than or on the rock itself. C. 'chrysonotus chirimila nets. blue' spawns in midwater, but males hold territories close to rocky shores. The biology and exploitation of these Copadichromis virginalis (Iles) fishes in northern Lake Malawi was colour photo on page 53 studied by Jackson et al. (1961). In addition to N. 'eucinostomus DISTINGUISHING FEATURES: Females and black' and N. 'eucinostomus yellow' immature males lack spots or stripes. (discussed below), there are a number This species has more gillrakers (21-25 of elongate, more benthic forms which ceratobranchials) than the O. argyro- are seined in large numbers from ex- soma species group (10-14) and N. eu- posed sandy beaches, and are some- cinostomus and relatives (15-17). It has times taken in chirimila catches. The a deeper body than C. mloto and Diplo- identification of these species is pres- taxodon limnothrissa. ently not properly worked out: Nyas- COLOUR: Females and non-breeding sachromis eucinostomus, N. inornatus males silvery, with generally yellow- and N. prostoma may belong to this ish colour. Ripe males are black. Larger, group. These are all relatively small long-finned individuals have a bright zooplankton feeders. At least three yellow or white dorsal fin 'blaze' which more species of this group are known extends along the dorsal surface of the to be abundant in shallow waters in the head and nape. Smaller short-finned northern part of the SE Arm (Stauffer individuals, such as those found in et al., 1993): N. cyclicos, N. thinos and Lake Malombe, have a black dorsal fin

67 with a broad yellow or white margin. large catches of ripe males are often These probably represent two species. made on sandy beaches at depths of MAXIMUM SIZE: 11-15 cm TL, depend- around 20-50m — although these may ing on population. not yet have settled on their nests. DIET: Zooplankton. In Lake Malom- However, one member of the C. vir- be, the stomachs were full of copepods, ginalis complex breeds in Lake Ma- although plankton samples and the lombe, which lacks any rocky habitat. guts of other fish species taken in same DISTRIBUTION AND ABUNDANCE: Iles haul indicated that the plankton was (1960) described this species from dominated by the cladoceran Bosmina Nkhata Bay, but stated that it was (N.C. Mwanyama, pers. comm.). Some found elsewhere. FAO (1976) found cladocerans, Chaoborus and algae also this (under the name Haplochromis found in the stomachs (17 specimens mloto) to be the most abundant species examined). in their trawl surveys of the SE & SW REPRODUCTION: Breeding season in the Arms and Domira Bay. Eccles & Trewa- north of the lake is March to June (Iles vas stated that this species (under the 1960). Size ranges of ripe females: name C. mloto) was one of the most Nkhata Bay 8-14cm (Jackson et al., abundant in the SE Arm, from 5 to 40 1961), SE. Arm 9-15 cm (under the fathoms (9-74m). Konings (1990a) has name Haplochromis mloto, Tweddle & found members of the C. virginalis Turner, 1977), Lake Malombe 8-10.5 cm complex throughout the lake, includ- (Table 2). Konings (1990a) reports that ing Mozambican and Tanzanian the smaller, large-eyed forms breed at shores. Two putative species often oc- depths of about 30m from August to cur sympatrically (Konings 1991 a, b). November, while the larger, more slen- In the 1992 survey, it was the 8th most der form breeds between 7 and 15m. abundant taxon identified, comprising Males of both forms excavate a spawn- almost 4% of the total sample weight ing pit against the side of a small rock. and occurring in 37 samples from 18 However, this habit of spawning in to 114m depth throughout the SE Arm. rocky areas does not seem to be uni- It often dominated sample catches, on versal in the C. virginalis complex, as one occasion comprising 86% of the sample weight off Mazinzi Bay at 50m depth. In 1990-91, it was Total Immature Active Ripe/Spent Mature also abundant in Lake Malombe Length unsexed Females Females Males and Domira Bay. Ad hoc trawl 5.0-6.9 4 — — — samples south of Monkey Bay 7.0-7.4 4 1 — — indicated that huge numbers 7.5-7.9 3 — — 2 were often found just off the 8.0-8.4 2 10 8 6 8.5-8.9 — 1 15 16 sandy bays at Chirombo, 9.0-9.4 — 1 8 24 Mazinzi and Nkhudzi. 9.5-9.9 — — 9 17 COMMERCIAL IMPORTANCE: This 10.0-10.4 — — 3 9 is very impor- tant in a variety of mechanised Table 2. Size at Maturity, Copadichromis virginalis from Lake Malombe, July 1992. and artisanal food fisheries. In

68 1990-91, it made up 6% of the midwa- all specimens labelled as Haplochromis ter trawl bycatch, and 3.5% of the pair mloto key out as C. virginalis. Thus, I trawl catch in Area A. It is also likely believe that all of the published data to be important in pair trawl catches on abundance, distribution, growth, elsewhere. It was the most abundant fecundity, population biology, and col- species in chirimila catches around the oration reported for H. mloto from the Nankumba Peninsula in 1992-93 southern part of the lake, actually re- (Smith, 1993), although catches were fer to C. virginalis (FAO, 1976; Turner, highly seasonal, peaking in November- 1977; Tweddle & Turner, 1977; Eccles December. It dominated chirimila & Trewavas, 1989). The photographs catches at Nkhata Bay in 1955-56 labelled as C. mloto in Axelrod (1993) (Jackson et al., 1961). The smaller, deep- also appear to be of C. virginalis. The bodied form of this species is one of accounts of Konings (1989) and the three major 'kambuzi' species Axelrod (1993) are still partly confused caught in Lake Malombe, with more by this earlier mis-identification. C. than 1,500 tonnes being caught annu- mloto has rarely been positively iden- ally, mostly by nkatcha seines. Species tified since its original description. I of the C. virginalis complex are increas- have never seen it. It did not occur in ingly popular as aquarium fishes, un- the pelagic surveys of the ODA/SADC der names like 'Haplochromis firecrest project (Menz 1995). Konings (1995b) mloto'. reports a single immature specimen NOTES: Working with specimens from from a sandy beach at Senga Point and the southern trawling grounds, FAO has tentatively assigned a male col- (1976) gave the following parameters: lected from the Tanzanian shore to this Log wt(g)= 2.752 log length (mm) - species.

3.364; K= 0.671; Linf = 184 mm; max. Iles (1960) recognised two forms of size caught = 176 mm, length at matu- C. virginalis at Nkhata Bay — the deep- rity = 103 mm; natural mortality rate = bodied Kaduna form (including the 0.92; log (fecundity)= 1.507 log (mm) - holotype) and the shallow-bodied, 1.585. Iles (1971), working at Nkhata more yellowish Kajose form. I also rec- Bay on chirimila fisheries, gave figures ognised different forms in the south of of K= 0.778; Linf = 121 mm; max. size the lake, and Konings (1990, 1991 a, b) caught = 128 mm; length at maturity = has found a great number of different 106 mm. colour forms, some of which may rep- Subsequent to the work of Jackson resent different species. and Iles, this species, or species group has been consistently confused with C. mloto. I have examined the types of Copadichromis quadrimaculatus both species and also all the remain- (Regan) ing voucher specimens collected by colour photo on page 53 researchers working on the southern trawl fisheries in the 1960s and 1970s DISTINGUISHING FEATURES: Females and (in Monkey Bay Fisheries Research Sta- immature males grey, generally with a tion collection). Apart from the types, single small dark flank spot. The snout

69 is more acutely pointed than that of C. pled occasional large adults from the pleurostigma and C. pleurostigmoides. middle of the lake over deep anoxic COLOUR: Females and non-breeding bottoms (Menz, 1995). Eccles & Trewa- males dark grey with a small supra- vas found this species to be common pectoral spot and sometimes a smaller in trawl catches at 18m, but occasion- spot on the caudal peduncle. Anal and ally occurring as deep as 55m. In the pelvic fins may have a yellowish tint. 1992 survey, it was fairly common at Ripe males dark blue (possibly paler 28-64m, particularly in Area B (be- when alive), sometimes with a patch tween Monkey Bay and Boadzulu Is.) of bright golden iridescent scales be- and was also found at Domira Bay. hind the operculum. Faint vertical bars Smith (1993) found it to be abundant often visible. Dorsal fin white or yel- off Cape Maclear peninsula and Ko- low and a 'blaze' of similar colour usu- nings (1990a) reported a lake-wide dis- ally extends to the nape and top of tribution. head as far as the upper jaw. There are COMMERCIAL IMPORTANCE: In 1990-91, conspicuous egg-spots on the anal fin. it was often caught in midwater trawl MAXIMUM SIZE: 23 cm TL (Konings catches, but not in large numbers (0.5% 1990a). bycatch weight), and rarely in deep- DIET: Zooplankton. Occasionally water bottom trawls. It is probably small fishes (Engraulicypris?) common in pair trawl catches in REPRODUCTION: From April to Septem- Domira Bay. It was occasionally taken ber. Konings (1990a) reports that males in large numbers by chirimila fisher- defend a rock which is used as a men around the Nankumba Peninsula spawning site at 20-50m depth. I have (Smith, 1993) and Jackson et al. (1961) seen large numbers of ripe males reported that this is an important spe- trawled from 20-40m depth, over cies in chirimila and gill net catches in sandy areas in Domira Bay and Chi- the north of the lake. rombo Bay, so they may breed over NOTES: There is considerable varia- pure sand, although it is possible that tion in colour and body form, particu- these males had not yet established larly of females, suggesting that there their territories. Minimum breeding may be more than one species. How- size is 15 cm (Jackson et al., 1961) or 16 ever, the different forms do not appear cm (Iles, 1971). Fecundity averages 52 to be separated in distribution or habi- eggs (Thompson et al., 1995a) . tat. Although Eccles & Trewavas (1989) DISTRIBUTION AND ABUNDANCE: Iles describe this species as having a supra- (1960) recorded the species from anal spot, I have found that this is gen- Nkhata Bay, Likoma, and Chisumulu erally absent or very faint (see also and stated that it was never caught by photographs in Konings, 1990a). Eccles seines, but was confined, while in- & Trewavas included material in their shore, to rocky areas. He believed that description which had previously been it was the only utaka which might excluded from this species by Iles spend part of its life in open waters. (1960). Iles (1971) gives the following

This was confirmed over 30 years later parameters: K= 0.764; Linf = 190 mm, when the ODA/SADC project sam- max. length caught = 188 mm

70 Copadichromis pleurostigma the lake, and Konings (1990a) reported (Trewavas) similar specimens from Chewere. colour photo on page 54 Konings (1995b) reports that it is con- fined to the northern half of the lake. DISTINGUISHING FEATURES: Females and Iles (1960), Eccles & Trewavas (1989), immature males generally with a sin- FAO (1976) and Smith (1993) have no gle large dark flank spot. Anal and information on this species. In 1990-91, pelvic fins bright yellow. It has a more I collected it from pair trawl catches at rounded snout than C. quadrimaculatus, Namiasi in the SE Arm, and from an and a less protrusible mouth and fewer experimental trawl at 46m near Maleri gillrakers (17-18 ceratobranchials) than Island. C. pleurostigmoides (21 or more). COMMERCIAL IMPORTANCE: Possibly a COLOUR: Females and non-breeding minor component of pair trawl, gill net males silvery with a yellowish hue. and chirimila catches. Fins, especially anal and pelvics, bright NOTES: During 1990-91, the similar yellow. There is a large black spot on Copadichromis pleurostigmoides (Iles) middle of flank, sometimes with a was occasionally recorded in trawl sur- smaller, fainter spot on caudal pedun- veys, but all specimens subsequently cle. Ripe male colour pale blue (Ko- re-examined in detail proved to C. nings, pers. comm.). pleurostigma or C. quadrimaculatus. MAXIMUM SIZE: 20 cm TL (Konings 1990a). DIET: Zooplankton. Copadichromis trimaculatus (Iles) DISTRIBUTION AND ABUNDANCE: The type is from Chilumba in the north of DISTINGUISHING FEATURES: Females and

Figure 21. Copadichromis trimaculatus; Taiwan Reef, Chisumulu Is.

71 immature males generally with three Copadichromis 'chrysonotus black' large dark spots. Anal and pelvic fins colour photo on page 53 bright yellow. COLOUR: Females and non-breeding DISTINGUISHING FEATURES: This species males silvery with a yellowish hue. has a larger eye than other Copadi- Fins, especially anal and pelvics, bright chromis species caught by seines and yellow. Three large black spots on trawls. It is the only utaka species with flanks. Ripe males dark blue, with a brownish body colour and three large bright iridescent blue head, bright or- spots. ange spot on each flank scale (Konings COLOUR: Females and non-breeding 1990a). males are brownish, with three large MAXIMUM SIZE: 23 cm TL (Konings black spots. Ripe males are black with 1990a). a yellow and white dorsal margin. The DIET: Phytoplankton, zooplankton yellow colour extends as a blaze down (Iles, 1961; Konings, 1995b). the forehead and onto the upper sur- REPRODUCTION: In August and Sep- face of the snout. tember, males defend a spawning site MAXIMUM SIZE: 13 cm TL. adjacent to a rock, generally at depths DIET: Cladocerans and copepods. of around 30m or greater, but occasion- Stomachs contained a much smaller ally as shallow as 15m (Konings, 1990a; proportion of Diaptomus than those of Konings, 1995b). Females guard their C. virginalis taken in the same seine fry (Konings 1995b). hauls in Lake Malombe. Chironomids, DISTRIBUTION AND ABUNDANCE: Gener- chaoborids and algae also taken (9 ally a shallow water species rarely seen specimens examined all from Lake in trawl catches in the 1992 survey. It Malombe, Mwanyama & Turner, un- was found in a single sample near the publ.). Ilala Gap. Described by Iles (1960) from REPRODUCTION: A female of 95 mm TL, Nkhata Bay and Likoma, Eccles & collected in Lake Malombe had 50 ripe Trewavas also reported it occasionally eggs in the ovaries. from trawls at 18m in the south of the DISTRIBUTION AND ABUNDANCE: Com- lake and recorded a large shoal 'at one mon in Lake Malombe and the Upper of the most northerly stations'. Konings Shire River. Not recorded in the 1992 (1990a) reported that it is found at trawl survey, but occasionally found in Nkhata Bay, Mbenji and the southern shallower water (20m or less) in the SE part of the lake and has subsequently Arm. (pers. comm.) seen it at Taiwan Reef, COMMERCIAL IMPORTANCE: Forms a sub- Jalo Reef, and in Tanzania. Smith (1993) stantial part of Nkatcha seine catches did not distinguish this species from in Lake Malombe (around 150t pa in C. pleurostigmoides. 1990-91). COMMERCIAL IMPORTANCE: Minor. NOTES: Further examination is neces- sary to determine whether this is the fish described as C. chrysonotus (Bou- lenger). Most publications (Iles, 1960; Jackson, 1960; Eccles & Lewis, 1981;

72 Ribbink et al., 1983, Konings, 1990a, DISTRIBUTION AND ABUNDANCE: Re- 1995b) assume that C. chrysonotus is a corded from shallow water (15-30m) large species (16 cm TL) which is con- on open sandy areas in the northern fined to rocky shores and has a male part of the SE Arm. breeding colour of dark blue, with pale COMMERCIAL IMPORTANCE: Not known. blue dorsally. The name 'chrysonotus' May be locally important in pair trawl means 'gold-back'. Eccles & Trewavas and seine catches. state that this refers to the preserved NOTES: Preserved material not reli- colour, but it is possible that when Bou- ably distinguished. lenger coined the name in 1908, the original male colours were still visible. The type is from an unknown location, Nyassachromis 'eucinostomus but it is a ripe male of 13 cm TL. How- yellow' ever Konings (pers. comm.) has re- colour photo on page 54 corded sexually active blue male C. chrysonotus as small as 7cm! DISTINGUISHING FEATURES: This species has a more pointed snout than those of the Otopharynx argyrosoma group Nyassachromis 'eucinostomus and fewer gillrakers (10 cerato- black' branchials) than C. mloto, C. virginalis, colour photo on page 54 N. 'eucinostomus black', and N. in- ornatus. DISTINGUISHING FEATURES: This species COLOUR: Females and non-breeding has a more pointed snout than those males are silvery. Ripe (?) males have of the Otopharynx argyrosoma group. a black snout, bright yellow opercula, Fewer gillrakers (16 ceratobranchials) and a bright yellow margin to the anal than C. mloto and C. virginalis (21-25), fin. and more gillrakers than N. 'eucino- MAXIMUM SIZE: 10cm TL. stomus yellow', N. cyclicos, N. thinos DISTRIBUTION AND ABUNDANCE: Com- and N. conophoros (10-14). It has a rela- mon in shallow water (35m or less) on tively smaller eye than the type of Co- open sandy areas all round the SE Arm. padichromis inornatus. This, and similar species comprised COLOUR: Females and non-breeding 8.3% of the fish biomass in experimen- males are silvery with a sandy cast. tal trawls at depths of 30m or less to Ripe males black with a broad white the south of Monkey Bay, and 2.5% dorsal fin margin and yellow anal fin north of Monkey Bay. margin. COMMERCIAL IMPORTANCE: In 1990-91, MAXIMUM SIZE: 12 cm TL. members of the N. eucinostomus com- DIET: Mainly copepods & clado- plex accounted for less than 0.5% of the cerans. Some insect larvae and plant midwater trawl bycatch, but are prob- material, as well as grit and detritus, ably important in pair trawl catches indicating that it is at least partly a north of Boadzulu Island. Their impor- benthic feeder (1 specimen examined tance in seine fisheries is unknown. by Grant & Turner, unpubl.).

73 Nyassachromis microcephalus Nyassachromis 'argyrosoma blue' (Trewavas) colour photo on page 54 colour photo on page 54 DISTINGUISHING FEATURES: A small elon- DISTINGUISHING FEATURES: A pale sil- gate laterally compressed species. very species with faint horizontal or Lower gillrakers 10. Differs from other vertical bars. More elongate than C. elongate species by its deeper, more virginalis. Deeper-bodied than N. eu- compressed body. Distinguished from cinostomus and C. mloto. Has fewer gill- P. longimanus and O. 'argyrosoma deep' rakers (13-14) than the three other spe- by its less deep body. Differs from O. cies. 'argyrosoma red' in coloration and COLOUR: Females and immatures sil- more pointed snout. very with an overall sandy cast. Faint COLOUR: Generally silvery, pale vertical or horizontal bars occasionally sandy dorsally. Pelvic fins orange-yel- visible. Ripe males bright blue with a low. Ripe males silvery, with 9 faint ver- yellow margin to the anal fin and nu- tical bars. Head darker with metallic merous bright orange spots on the dor- blue snout. Lips bright bluish white. sal and anal fins and flanks. Operculum silver. Chest dusky, belly MAXIMUM SIZE: 15 cm TL (Konings white. Pelvic fins dusky, black 1990a). anteriorly. Anal whitish proximally, DIET: Zooplankton (Konings 1990a). darker distally, with 0-3 large yellow- REPRODUCTION: Konings (1990a) re- ish egg-spots, sometimes with orange ports that the males build sand-castle margin. Dorsal fin dusky, with dull nests and generally aggregate in shal- orange spots posteriorly, white lappets. low water. Caudal dusky with dull orange spots, DISTRIBUTION AND ABUNDANCE: De- darker on upper and lower edges. Pec- scribed from Monkey Bay, but Konings toral fins yellowish. Specimens from (1990a) reports that it is also found at Maleri have a blue head and dorsum Ruarwe in the north of the lake. Occa- and yellow belly. These may represent sionally taken in shallow water trawl a different species or race or else they catches in 1992. may be full reproductive colours. COMMERCIAL IMPORTANCE: May be sea- MAXIMUM SIZE: 12cm TL. sonally important in seine net or DIET: Two specimens taken from chirimila catches. south of Boadzulu Island contained NOTES: Easily confused with small, mostly remains of small crustaceans slender largely colourless species such and filamentous diatoms. A single in- as other Nyassachromis species, and the dividual from Chilinda, in the north species of the O. argyrosoma complex. eastern part of the SE Arm, had in- Together these small species form a gested chironomid larvae and cope- very large proportion of the inshore pods, with some algal material (Grant fish biomass. & Turner, unpubl.). DISTRIBUTION AND ABUNDANCE: Posi- tively identified from shallow water catches (to 20m depth) throughout the

74 SE Arm, and tentatively identified from trawls as deep as 60m. Found in most samples throughout its preferred depth range and often dominant in shallow waters. COMMERCIAL IMPORTANCE: A major spe- cies in pair trawl catches (11%) and more than 1% of midwater trawl catches. Also taken in beach seines. NOTES: This was one of several spe- cies recorded as Otopharynx argyrosoma in the records of the Monkey Bay Fish- eries Station. It does not show a mid- lateral blotch, and consequently does not fit the present definition of the ge- nus Otopharynx. There may be several similar species. Konings (1990: 165) il- lustrates what appears to be a female of this species as Nyassachromis leuciscus and a male is shown by Konings (1995b). The holotypes of N. leuciscus and O. argyrosoma are small colourless juveniles from unknown locations, so it is very unlikely either species can be positively identified in the field. Konings also illustrates a male (p.165) which resembles N. 'argyrosoma blue', but which has a larger number of egg-spots on the anal fin.

75 Chapter 9 Lethrinops and allied genera

The large number of Lethrinops spe- Lethrinops, especially those of the L. cies, and those of the closely related micrentodon / L. stridei / L. microdon genera Taeniolethrinops and Tramitichro- group. This makes it unfortunate that mis, are demersal feeders on sediment Eccles & Trewavas subsequently chose or benthic invertebrates. Many of these H. longimanus the type species of the species have restricted depth distribu- genus Placidochromis. tions and habitat preferences. Most Many species of Lethrinops remain to Lethrinops species are deep-bodied, be described, including some of the vertically-barred, with small ventrally- most important commercial species. In placed mouths. Lethrinops and related addition to those species I discuss be- genera dominate the demersal trawl low, there are a number of other spe- fisheries, and are also important in cies illustrated by Konings (1990, 1991, beach seine and Nkatcha net catches. 1992), mainly from shallow water. In Lethrinops species caught by the mid- addition, a number of the described water or bottom trawl are marketed as species have not been recorded since chisawasawa, as are small individuals their initial collection and, in some taken by pair trawls. Large individu- cases, the types have not been re-ex- als in pair trawl, seine or gill net catches amined since the 1930s. Eccles & Lewis, are recorded as mbaba, and small fish in a series of papers in the late 1970s, from seine catches are kambuzi. re-described some of these species, as These three genera were originally well as describing several new species. united as Lethrinops and were distin- It appears that some of their re-descrip- guished from other haplochromine tions were based on material not con- cichlids by their dentition: the outer specific with the type material, which row of teeth on the lower jaw curves in some cases was not measured. Also, round to end just posterior to the in- some of their voucher specimens were ner rows, whereas in other haplochro- lost before the description was pub- mine genera, it is extended posteriorly lished and others deposited in the field as a long single row. Eccles & Lewis collection at Monkey Bay have long (1977) included the species Haplochro- ago been destroyed. At present, the mis micrentodon in Lethrinops, because state of knowledge of this large and of its resemblance to L. stridei and L. commercially important group can microdon. This species has a dental ar- best be described as fragmentary. cade which is intermediate between the typical Lethrinops form and the 'Haplochromis-style'. Eccles & Lewis (1977) rightly noted the similarity be- tween Haplochromis longimanus and

76 Lethrinops microdon Eccles & Lewis cles & Trewavas (1989) state that it is colour photo on page 55 only known from the SE Arm. In the 1992 survey, it was found between 48 DISTINGUISHING FEATURES: A deep-bod- and 102m and was occasionally quite ied Lethrinops species with a relatively numerous at depths of 50-80m be- small rounded head and short snout, tween Boadzulu Island and Monkey it has a larger number of lower gill- Bay. However, populations have rakers (24-29) than most similar spe- clearly declined in this area. Experi- cies. It can be quickly distinguished mental trawling between 60 and 80m from the superficially similar L. stridei, depth indicated that this species com- L. micrentodon and L. 'micrentodon prised 44% of the cichlid fish biomass Makokola' by the number of bars be- in 1983-85, but only 22% in 1991 and low the dorsal fin (7, compared to 8- 5% in 1992. 9), and from L. micrentodon and L.' yel- COMMERCIAL IMPORTANCE: In 1990-91 low chin' by the absence of a midlateral this species was the most important in spot. It generally has a larger mental the deep-water demersal trawl fishery process than L. 'oliveri'. (28% by weight) and comprised 2% of COLOUR: Females and non-breeding the midwater trawl bycatch. males are silvery with a bluish sheen, NOTES: FAO (1976) did not distin- with 7 faint bars below the dorsal fin. guish this species or L. micrentodon Anal, pelvic and caudal fins yellowish. from Lethrinops stridei. Ripe males are dark bronze, with prominent dark bars. Lips and head with bluish iridescence. Dorsal lappets Lethrinops micrentodon (Regan) white tipped. Anal with 6 dull yellow egg-spots. DISTINGUISHING FEATURES: A deep-bod- MAXIMUM SIZE: 20 cm TL (Lewis & ied Lethrinops species with a relatively Tweddle, 1990). small rounded head and short snout, DIET: Eccles & Lewis (1977) recorded with fewer lower gillrakers (15-19) stomach contents dominated by than L. microdon, L. stridei and L. benthic diatoms principally the fila- 'oliveri'. It has more vertical bars (8-9) mentous Aulacoseira (= Melosira), with than L. 'oliveri' and L. 'yellow chin' (7). occasional crustacean and chironomid It has a less deep body and smaller eye fragments. than L. 'micrentodon Makokola', which REPRODUCTION: A non-seasonal also lacks a midlateral spot and has a breeder. Ripe males 13.5-20 cm TL, ripe different male breeding colour. Placido- females 11.8-17.8 cm. Sizes at 50% ma- chromis 'longimanus Namiasi' has a turity 153 and 137 mm respectively more acutely-pointed snout. (Lewis & Tweddle, 1990). COLOUR: Females and non-breeding DISTRIBUTION AND ABUNDANCE: Eccles & males silvery, with a golden sheen and Lewis (1977) report that a single speci- 7-8 faint bars under the dorsal fin. men was taken from the SW Arm, but There is a faint midlateral spot between was otherwise known only from 20-60 bars 3 and 5. Anal, pelvic and caudal fathoms (37-110m) in the SE Arm. Ec- fins yellowish. Sexually active males

77 Figure 22. Lethrinops micrentodon; South-East Arm. with bluish sheen on head. Lappets low chin' and L. 'micrentodon Ma- white with orange tips (Eccles & Lewis kokola'. This latter species is illustrated 1977). as L. micrentodon in Konings (1990a) MAXIMUM SIZE: 15.5 cm TL (Eccles & and Baasch (1991). FAO (1976) did not Lewis 1977). distinguish this species from L. stridei. DIET: Benthic diatoms, both pinnate and filamentous forms, with occa- sional crustacean remains (Eccles & Lethrinops stridei Eccles & Lewis Lewis, 1977). colour photo on page 55 DISTRIBUTION AND ABUNDANCE: Uncom- mon, and patchily distributed. The lo- DISTINGUISHING FEATURES: A deep-bod- cation of the type specimens is not ied Lethrinops species with a relatively known. Jackson (1961) reported that it small rounded head and short snout, was known only from the south of the with fewer lower gillrakers (19-23) lake. Eccles & Lewis (1977) report it than L. microdon, but more than most from depth of 20-25m in the SE Arm, other similar species. It has a deeper and also from the SW Arms, Domira body and larger number of bars under Bay, and Bandawe. In the 1992 survey, the dorsal fin than L. 'oliveri'. it was positively identified from 18m COLOUR: Females and non-breeding at Fowo, and recorded from two other males silvery with bluish-green or samples taken at 20m off Namiasi and golden sheen, with 9 (occasionally 7 or 35m at Michesi. 8) faint bars below the dorsal fin. Pel- COMMERCIAL IMPORTANCE: Not record- vic fins yellow. Ripe males darker blue- ed in commercial trawl catches, but grey, with strong vertical bars, dark fins perhaps this is due to mis-identifica- and white dorsal lappets. Anal fin with tion. numerous large yellowish egg-spots. NOTES: I did not initially distinguish MAXIMUM SIZE: 13 cm SL (Eccles & this species reliably from Placidochromis Lewis, 1977), approximately 15.5 cm 'longimanus Namiasi', Lethrinops 'yel- TL.

78 DIET: A benthic feeder, some stom- it is distinguished by its low number achs examined by Eccles & Lewis of lower gillrakers (13-17), deep body, (1977) contained mostly calanoid large eye and lack of a midlateral spot. copepods, some were dominated by It usually has numerous stripes (9, oc- Aulacoseira and others by unicellular casionally 7-8 under the dorsal fin). diatoms. COLOUR: Females and non-breeding DISTRIBUTION AND ABUNDANCE: Eccles & males silvery. Ripe males generally Lewis (1977) recorded this species from dark, with 9 (occasionally 7-8) dark the SE and SW Arms, Domira Bay and vertical bars below the dorsal fin. Bandawe in the north. They report that Snout and cheeks dark blue. Bright it is found at 9-25 fathoms (17-46m) in orange from top of head to chest. Body the SE Arm and 16-30 fth (30-55m) in between bars with iridescent blue the SW Arm. It was formerly the most sheen. Dorsal fin dark, with broad abundant demersal species south of white margin, and orange-tipped lap- Boadzulu Island, but was less common pets. in the 1992 survey. Most of the speci- MAXIMUM SIZE: 13 cm TL (Konings mens caught came from the Namiasi 1995b). station at 24m depth, where it was the REPRODUCTION: Aquarium observa- most abundant species in both Febru- tions (Baasch, 1992a) indicate that the ary and May surveys. Elsewhere, it male constructs a large (90 cm diam- occurred in small numbers in 3 sam- eter) pit with a small spawning cone ples: 20m depth at Fowo and at 34 and in the centre. Females of 10cm TL pro- 44m at Mpemba. duce around 50-60 fry. Ripe males are COMMERCIAL IMPORTANCE: In 1990-91, known to build nests at 30-35m depth, it comprised around 3% of the pair near Makokola Reef (Konings, 1990a; trawl catch south of Boadzulu Island. Baasch, 1992a). Reproduction appears It was also seen in catches by chambo to be non-seasonal. seines. DISTRIBUTION AND ABUNDANCE: Locally NOTES: FAO (1976), Turner (1977) and abundant, but apparently restricted to Tweddle & Turner (1977) give length- a strip of coast 22 km long on the west weight relationship, asymptotic coast of the SE Arm, between Maldeco length, natural mortality, and changes and Nkhudzi Bay, at depths of 24-60m. in abundance for Lethrinops stridei, but In the 1992 trawl survey, it was numer- they did not distinguish between this ous in trawl hauls in this area, but was species and L. microdon and L. micrent- only recorded at Mpemba (34-44m), odon (Tweddle, pers. comm.). Ulande (46-48m) and Nkope (54-60m). COMMERCIAL IMPORTANCE: In 1990-91, this species comprised around 1% of Lethrinops 'micrentodon Makokola' pair trawl catch S of Boadzulu Is. and colour photo on page 55 it probably also occurred in midwater trawl catches. DISTINGUISHING FEATURES: A deep-bod- NOTES: Confused with Lethrinops ied Lethrinops species with a relatively micrentodon by Konings (1990a) and small rounded head and a short snout, Baasch (1992a). During examination of

79 specimens on the laboratory, I found TL (Lewis & Tweddle, 1990). that some individuals I had assigned DISTRIBUTION AND ABUNDANCE: In the to this species had 21-25 lower gill- 1992 survey it was found at depth of rakers. They also have generally 64-84m at Fowo, Chiponda, and Ma- smaller eyes, a lower caudal peduncle zinzi, but further north it was only length/depth ratio and a shorter pre- recorded in deeper water — 92-128m maxillary pedicel. It has not yet been at Monkey Bay, Ngusi, and Domwe. It determined whether these are small was often numerous and frequently specimens of Lethrinops stridei or rep- comprised a larger proportion of the resent a further undescribed species. catch weight than any other species. In 1983-85, this species comprised 12% of the biomass taken by demersal trawl- Lethrinops 'oliveri' ing between 60 and 80m, north of colour photo on page 55 Boadzulu Island and south of Monkey Bay, 10% in 1991 and 7% in 1992. DISTINGUISHING FEATURES: A medium- COMMERCIAL IMPORTANCE: In 1990-91, sized Lethrinops species with a small this species comprised around 4% of rounded head, short snout and elon- the demersal stern trawl catch. gated dorsal and anal fins. It has fewer NOTES: Lewis & Tweddle (1990) re- lower gillrakers (17-21), a smaller head corded this species as Lethrinops sp. A and a less prominent mental process (Tweddle, pers. comm.). During field than L. microdon, but more gillrakers work, I did not accurately distinguish than other similar species, excepting L. this species from L. 'yellow chin' and stridei, which has a deeper body and juvenile L. microdon. According to FAO more vertical bars. Small specimens are (1976), log(length)= 2.467 log(weight)- difficult to tell apart from L. 'dark'. The 2.815. latter species generally has a more deeply forked caudal fin, a more pointed snout and a relatively shorter Lethrinops 'grey' preorbital depth (17-20% HL in L. 'dark'; 19.5-24.7% in L. 'oliveri'.). DISTINGUISHING FEATURES: A fairly non- COLOUR: Females and non-breeding descript medium-sized Lethrinops with males are silvery with a bluish sheen, a relatively small rounded head and and 7 faint bars below the dorsal fin. short snout. It has fewer lower gill- Dorsal fin lappets are brownish with rakers (13-15) than L. microdon, L. yellow tips. Pelvic and pectoral fins stridei, L. micrentodon, and L. 'oliveri'. yellowish. Ripe males are dark silvery- It lacks the midlateral blotch of L. 'yel- grey, with prominent bars. Cheeks with low chin' and L. micrentodon and has blue iridescence, top of head yellow- fewer vertical bars (7) than L. micrent- bronze. Dorsal lappets black. odon (8-9) and L. 'micrentodon Mako- MAXIMUM SIZE: 15 cm TL. kola' (8-9, occasionally 7). L. 'micrent- DIET: Benthic filamentous diatoms. odon Makokola' is a deeper-bodied REPRODUCTION: Non-seasonal breeder. species. Size at 50% maturity of males is 12 cm COLOUR: Live coloration not known.

80 Figure 23. Lethrinops 'grey', male, South-East Arm.

Females and non-breeding males sil- low, black anteriorly. It probably rep- very grey with 7 vertical bars under the resents another species. dorsal fin. Ripe males with dark snout, pale lappets and black submarginal bar. Lethrinops 'yellow chin' MAXIMUM SIZE: 11 cm TL. colour photo on page 55 DISTRIBUTION AND ABUNDANCE: First re- corded in the 1992 survey, it was found DISTINGUISHING FEATURES: A typical at depths of 60-90m. It was infrequent, medium-sized Lethrinops species with but occasionally numerous at Mazinzi a relatively small rounded head and (60m) and Chiponda (72m) short snout. It is distinguished from COMMERCIAL IMPORTANCE: May be lo- most similar species by the low cally important in trawl catches, but number of gillrakers (13-17), and the was not recorded during the sampling prominent midlateral spot. It differs of the commercial fisheries, perhaps from L. micrentodon in its smaller head due to mis-identification. and lower number of bars under the NOTES: I did not initially distinguish dorsal fin (6-7). More slender than L. this species from other medium-sized 'micrentodon Makokola' and with a Lethrinops. A similar specimen col- less deep caudal peduncle (L. ''yellow lected at 90m off Monkey Bay (photo chin': 10-12% SL; L. 'micrentodon 17.34, male) had 19 ceratobranchial Makokola': 12-13% SL) and usually gillrakers. It was a mature male of narrower interorbital width (L. 'yellow 10cm TL with 7 dark vertical bars un- chin': 18-23% HL; L. 'micrentodon der the dorsal fin. Dorsal fin with dull Makokola': 21.5-26% HL). The male orange spots, dark lappets with white breeding colours distinguish this spe- tips. Anal fin with 5-6 dull yellow cies from all similar fishes, except spots. Caudal fin dark. Pelvic fins yel- Trematocranus brevirostris which has

81 fewer lower gillrakers and a larger eye. from other medium-sized species be- COLOUR: Females and non-breeding fore the 1992 survey. There are further males silvery with bluish sheen. similar specimens which are probably Darker sandy-colour dorsally, with 6- different species: similar-looking speci- 7 faint bars below the dorsal fin, and a mens from deeper water to the north dark spot between bars 2 and 3. Ripe of Boadzulu Island generally have males silvery with bluish sheen dorsal- deeper bodies and lack the midlateral ly, whitish ventrally and with promi- blotch, a single individual with very nent dark bars and spot. Top of head similar male breeding colour, taken at dark blue, cheeks, opercula and throat 46 m depth off the Maleri Islands bright yellow, becoming more intense (photo 8.9), measures only 64 mm SL ventrally. Black underneath head and (ca 8 cm TL) and has only 9 cerato- throat, extending to pelvic fins. Pelvic branchial gillrakers. fins yellowish posteriorly, black anteriorly. Anal fin dark grey, black lower margin, with 4-10 yellowish egg- Lethrinops 'dark' spots. Caudal fin dark yellow with colour photo on page 55 paler spots, dark distally and on up- per and lower edges. Dorsal fin light DISTINGUISHING FEATURES: Distin- grey, with brownish-yellow spots on guished from other small Lethrinops soft dorsal. Dorsal lappets white, species by the elongated dorsal, anal sometimes with orange-yellow tips, and caudal fins, high number of and a dark submarginal band. Iris yel- gillrakers (19-20), and dark male breed- low-gold, darker dorsally. ing colour. Difficult to distinguish from MAXIMUM SIZE: 11 cm TL. small specimens of L. 'oliveri'. The lat- DISTRIBUTION AND ABUNDANCE: In the ter species generally has a less deeply 1992 trawl survey it was found to have forked caudal fin, a less pointed snout, a very restricted distribution, occurring and a relatively deeper preorbital bone only in samples from 44-54m depth (17-20% HL in L. 'dark'; 19.5-24.7% in around Boadzulu Island, at trawl sta- L. 'oliveri'.). tions Mpemba, Fowo, Ulande, and COLOUR: Ripe males very dark, Nkope. It was often abundant. In other smoky grey, 7 black bars under the surveys, it was occasionally taken in dorsal fin. Unpaired and pelvic fins shallower water in the southern part very dark grey to black. Sometimes a of the SE Arm. greenish tint on snout. Females and COMMERCIAL IMPORTANCE: Although immature males silvery and weakly the distribution of this species is very barred. restricted, it lies in the centre of the MAXIMUM SIZE: 10cm TL. principal trawling grounds and, in DISTRIBUTION AND ABUNDANCE: Often 1990-91, L. 'yellow chin' comprised abundant at depths of 22-64m and, around 1% of the midwater trawl overall, one of the dominant species in bycatch, and 1.5% of the pair trawl the SE Arm. In the 1992 survey it was catch south of Boadzulu Island. especially abundant at the Ulande sta- NOTES: Not reliably distinguished tions, comprising 35% of the catch

82 weight at 48m and 23% at 46m. It was of the biomass. also the dominant species in catches at COMMERCIAL IMPORTANCE: Negligible. 44m at Mpemba (31%) and at 60m at Nkope (25%), the latter being the far- thest north it was recorded. Lethrinops 'black chin' COMMERCIAL IMPORTANCE: One the colour photo on page 56 most significant demersal haplochro- mines in the trawl fisheries. In 1990- DISTINGUISHING FEATURES: Intermediate 91, it comprised around 1% of mid- in body shape between L. 'oliveri', and water trawl bycatch, 5% of deep-wa- L. 'yellow chin' and the 'high-backed' ter bottom trawl catch and 8% of pair L. altus and L. 'matumbae'. This small trawl catch south of Boadzulu Island. deep-water species has a distinctive NOTES: Females and immature fish black and white male breeding colour. were not reliably distinguished from It can be distinguished from most of other small or immature Lethrinops. the other small Lethrinops by its lower number of gillrakers (12 cerato- branchials) and from Lethrinops 'yel- Lethrinops 'yellow tail' low' by its higher gillraker count, colour photo on page 55 deeper body and large number of ver- tical bars. Lethrinops 'blue orange' is DISTINGUISHING FEATURES: Distin- more slender, with smaller eyes and guished from most other small Lethri- has a larger number of vertical bars (8- nops by the lower number of gillrakers 9 under the dorsal fin). The lips are not (10-12 ceratobranchials). Differs from curved, as are those of L. altus. L. 'blue-orange' in the lower number COLOUR: Ripe males silvery-white, of vertical bars, and from L. 'yellow' by darker dorsally with 6-7 vertical bars the thinner vertical bars, and deeper under the dorsal fin, one across the body. Ripe males are paler and lack the nape and two on the caudal peduncle. strong black and white dorsal fin Lips, chin and chest black. Dorsal fin markings. orange brown with a broad white mar- COLOUR: Ripe males grey with 7 dark gin and some white spots and stripes vertical bars under the dorsal fin. Dor- on lower part. Caudal striped. Female sal fin with dull orange spots, white silvery, with sandy cast and fainter lappets with thin black submarginal brown vertical bars. band. Anal fin with several dull yel- MAXIMUM SIZE: 9.5 cm TL. low spots. Caudal fin yellowish. Pel- DISTRIBUTION AND ABUNDANCE: An un- vic fins orange-yellow, black anteriorly. common, apparently local species, not Females and immature males paler. recorded in the 1992 trawl survey. Sev- MAXIMUM SIZE: 8 cm TL. eral specimens were recorded from a DISTRIBUTION AND ABUNDANCE: Uncom- single sample from the photographic mon. In the 1992 survey, it was re- surveys at 90m depth off Monkey Bay. corded only from the Monkey Bay sta- COMMERCIAL IMPORTANCE: Negligible. tion at depths of between 90 and 100m, If a deep water trawl fishery is estab- where it comprised a small proportion lished it is possible that this, and other

83 similar-sized species could dominate Lethrinops 'blue-orange' the fishery. colour photo on page 56 NOTES: Not reliably distinguished in preserved samples, and would not DISTINGUISHING FEATURES: Distin- have been readily noticed except when guished from Lethrinops 'dark', by its exhibiting male breeding colour. lower number of gillrakers (12-13) and the lack of extended dorsal and anal Lethrinops 'yellow' fins. L. 'yellow' has a larger eye and colour photo on page 56 fewer lower gillrakers. COLOUR: Ripe males generally dark, DISTINGUISHING FEATURES: A small with 8-9 dark bars under the dorsal fin. Lethrinops with a large elliptical eye, a Snout metallic blue. Orange on top of steeply-angled head profile reminis- head, nape and on the dorsal surface cent of Pseudotropheus tropheops and a as far back as the midpoint of the dor- low number of gillrakers (9-10 cerato- sal fin. Behind the head, the orange branchials). The ripe male coloration colour extends ventrally to the chest. is distinctive. Fins dark, anal with several large ir- COLOUR: Ripe males pale, silvery, regular yellowish egg-spots. Females with 7 broad bars under the dorsal fin. and immature males pale, with faint The head and nape are bright yellow- bars. orange. Dorsal fin with broad white MAXIMUM SIZE: 8 cm TL. margin, with yellow tips and a black DIET: Copepods and cladocera. Some submarginal band. The anal fin has ostracods, algae and chironomid lar- several irregular yellowish egg-spots. vae. (1 specimen examined by Grant Females and non-breeding males sil- & Turner (unpubl.)). very with faint bars. DISTRIBUTION AND ABUNDANCE: Abun- MAXIMUM SIZE: 7 cm TL. dant. In the 1992 trawl survey, it was DISTRIBUTION AND ABUNDANCE: Com- present in most samples at 22-40m in mon, occasionally dominant between SE Arm, south of Monkey Bay. It was 40 and 75m in the SE Arm. In the 1992 often numerous, comprising 16% of the survey, it was recorded at Mpemba catch weight from a sample from (44m), Ulande (48-52m), Nkope (54- Namiasi at 24m. It was occasionally 60m), Mazinzi (62m), Chilinda (60- taken as deep as 50m in the SE Arm 66m), and Chekopa (72m). In other and reported from 75m in Domira Bay. surveys, it was also taken at Namalaka COMMERCIAL IMPORTANCE: A major and Monkey Bay, over the same depth component of pair trawl catches sam- range. pled in 1990-91, comprising 14% of the COMMERCIAL IMPORTANCE: Minor com- catch weight south of Boadzulu Island. ponent of trawl catches. It is occasionally taken by other trawls. NOTES: Not previously known before NOTES: Not recorded before the the present work, this species may not present work and initially not reliably have been reliably distinguished in all distinguished by the author in surveys material examined in the trawling sur- of commercial fisheries. veys.

84 Lethrinops 'matumbae' cept L. 'deep water altus' by the curved colour photo on page 56 upper jaw. Differs from L 'deep water altus' in its longer, ventrally-angled DISTINGUISHING FEATURES: A deep-bod- snout and different male breeding col- ied species with a long snout and ours. ventrally placed mouth, it is distin- COLOUR: Silvery with an overall guished from other similarly-shaped brownish hue with 6 faint vertical bars species, such as L. altus and L. longi- under the dorsal fin. Ripe males pinnis by the larger number of gill- bronze, shading to orange on nape, rakers (17), and from L. altus by the with dark brown bars, snout with blue straight jaws. iridescence. Several large irregular yel- COLOUR: Silvery with an overall low spots and streaks on the anal fin. brownish hue. 6-7 vertical bars under MAXIMUM SIZE: 15 cm TL. the dorsal fin. Pectoral fin yellow. Ripe DIET: Benthic invertebrates. males more strongly barred with dark DISTRIBUTION AND ABUNDANCE: Eccles & brownish head and fins and a bluish Trewavas report that the species was sheen on snout. described from a single specimen in the MAXIMUM SIZE: 11 cm TL. Christy collection and do not provide DISTRIBUTION AND ABUNDANCE: In the a redescription. They report the spe- 1992 survey it occurred in most sam- cies as being found at 10-70 fathoms ples taken from north of Boadzulu Is., (18-110m), but most common at 10-30 at depths of 18-75m. It was never abun- fth (18-55m). In the 1992 survey it was dant. Also known from Domira Bay. common at depths of 40-60m in the SE COMMERCIAL IMPORTANCE: A few speci- Arm, occasionally as shallow as 20m. mens occasionally taken by semi- The deeper records of Eccles & Trewa- pelagic trawls. vas are probably the result of confu- NOTES: Eccles & Trewavas (1989) state sion with L. 'deep water altus'. that an undescribed species recorded COMMERCIAL IMPORTANCE: Comprises as 'Lethrinops matumbae' has more around 1% of pair trawl and midwater than 30 lower gillrakers. The speci- trawl catches. mens labelled as such at the Monkey NOTES: Initially, I did not reliably dis- Bay fisheries station field museum, tinguish this species from Lethrinops probably collected by Eccles, do not 'deep-water altus' and consequently have 30 lower gillrakers and appear to the distribution and abundance data be conspecific with the material I have are estimated post hoc, from examina- collected. I did not find any Lethrinops tion of the depth and distribution of species which had such a high number preserved specimens collected in the of ceratobranchial rakers. photographic surveys.

Lethrinops altus Trewavas Lethrinops 'deep water altus' colour photo on page 56 colour photo on page 56

DISTINGUISHING FEATURES: Distin- DISTINGUISHING FEATURES: The curved guished from all other Lethrinops, ex- upper jaws serve to distinguish this

85 deep-water species from all other DIET: Chironomid larvae were found Lethrinops, except L. altus. It differs in the stomach of the single specimen from L. altus in having a less ventrally examined (Grant & Turner, unpubl.). placed mouth and different male DISTRIBUTION AND ABUNDANCE: The spe- breeding colours. cies was described from three speci- COLOUR: Silvery with faint vertical mens from Monkey Bay. Eccles & bars. Ripe males are grey with a black Trewavas record it from the SW Arm head, black fins and 6 strong black ver- at 10-20 fths (18-37m) depth. In the tical bars under the dorsal fin. The dor- 1992 survey it occurred in most sam- sal lappets black, with broad white ples from south of Boadzulu Island at submarginal bar. depths of 20-34m. It was usually taken MAXIMUM SIZE: 10cm TL. in small numbers, but was numerous DIET: Benthic invertebrates. off Namiasi at 24-28m. It was not re- DISTRIBUTION AND ABUNDANCE: Abun- corded north of Boadzulu Island. dant. In the 1992 trawl survey this spe- COMMERCIAL IMPORTANCE: Minor com- cies occurred at depths of 60 to at least ponent of shallow water trawl catches. 120m. COMMERCIAL IMPORTANCE: Estimated to comprise around 5% of deep-water Lethrinops polli Burgess & Axelrod bottom trawl catch in 1990-91. colour photo on page 57 NOTES: Not reliably distinguished from L. altus during the earlier part of DISTINGUISHING FEATURES: It can be dis- the study. The distribution and abun- tinguished from all other Lethrinops, dance data were estimated post hoc, except L. christyi, by its long pointed from examination of the depth and dis- snout, and small terminal mouth. More tribution of preserved specimens col- slender than L. christyi. lected in the photographic surveys. COLOUR: Silvery with a pinkish sheen. Ripe males generally brownish with 7 prominent dark bars under dorsal fin. Lethrinops christyi Trewavas Snout with bluish sheen. Dorsal fin colour photo on page 56 whitish blue anteriorly. Anal fin with several large irregular yellowish egg- DISTINGUISHING FEATURES: Distin- spots. guished from all other Lethrinops, ex- MAXIMUM SIZE: 19 cm TL. cept L. polli by its pointed snout, and DISTRIBUTION AND ABUNDANCE: Re- small terminal mouth. Deeper bodied ported by Eccles & Trewavas as com- than L. polli. mon in deeper trawls in the southern COLOUR: Silvery with a yellowish iri- parts of the lake, mostly deeper than descence. Ripe males blue-grey, with 8 40 fths (74m), but occasionally as shal- prominent dark vertical bars. Head low as 20 fths (37m). In the 1992 sur- and fins dark blue. Several large egg- vey it was common, but never numer- spots on anal fin. ous, occurring at depths of 45 to at least MAXIMUM SIZE: 18 cm TL (Konings 128m. It was more common in deeper 1990a). water. Very scarce on the north-eastern

86 part of the SE Arm (Chekopa, Chilinda, nant species at depths of 90m or more. Chiponda stations). I have recently (1996) collected what COMMERCIAL IMPORTANCE: In 1990-91 it appears to be a female L. gossei from a comprised around 1.5% of deep-water gill net catch off Karonga in the far bottom trawl catch, and occasionally north of the lake. taken by the midwater trawl. COMMERCIAL IMPORTANCE: This and the following species, which were not re- liably distinguished, comprised Lethrinops gossei around 5% of the deep-water demer- Burgess & Axelrod sal trawl catch in 1990-91. colour photo on page 57 NOTES: Eccles & Trewavas report that a similar species with fewer gillrakers DISTINGUISHING FEATURES: A distinctive has been found at Nkhotakota at 200m deep-bodied species with a large eye depth. Lethrinops gossei was known as and mouth. The jaws are characteris- 'Lethrinops orbitalis' by Eccles, who tic — the tip of the lower jaw is curved did not publish a description, and un- upwards and fits into a V-shaped cleft der this name FAO (1976) gave a in upper jaw. length-weight relationship of log(wt)= COLOUR: Females and immature 3.629 log (len)-5.235. males silvery with a pinkish sheen. Ripe males dark grey, with 7 dark bars under the dorsal fin. Fins dark. Head Lethrinops 'deep-water albus' with dark purplish or greenish irides- colour photo on page 57 cence. MAXIMUM SIZE: 20 cm TL. DISTINGUISHING FEATURES: This species DIET: Benthic . has a large head and eye and is super- REPRODUCTION: Studied by Lewis & ficially similar to L. gossei, from which Tweddle (1990) in the South East Arm, it can be distinguished by the absence 1983 to 1985. Mature females measured of a V-shaped cleft in the upper jaw and 13-18 cm TL, and males 12.5-20 cm. by the lower number of lower gill- 50% were mature at 147 and 156 mm rakers (9 compared to 18-20). This spe- respectively. Around 20% of females cies has very similar meristics and male above the size of 50% had ripe gonads breeding colours to L. longipinnis but from December to September, with a L. 'deep-water albus' has a shorter peak of 60% ripe in March. No ripe fe- snout and thinner lips. L. albus Regan males were recorded from October to is a more slender fish. November. (Lewis & Tweddle, 1990). COLOUR: Females and immature DISTRIBUTION AND ABUNDANCE: Abun- males silvery. Ripe males dark grey, dant in deep water. Eccles & Trewavas with 7 dark bars under the dorsal fin. report that it dominates the benthic Fins dark. Head dark with greenish community at depth of 50-70 fathoms iridescence. (92-130m). In the 1992 trawl survey, it MAXIMUM SIZE: 15 cm TL. occurred at depths of 46 to at least DISTRIBUTION AND ABUNDANCE: Com- 128m. It was generally one of the domi- mon in deep water. In the 1992 survey

87 it was found at depths of 80 to 128m. It dant over a wide depth range. The probably occurs deeper. holotype was collected from 40 fath- COMMERCIAL IMPORTANCE: see L. gossei oms (74m) off Monkey Bay. The para- above. types were from 12-58 fths (22-107m) NOTES: This species was known as L. in the SE Arm. Eccles & Lewis (1978) 'cf. albus' during trawl surveys under- stated that it was most abundant from taken by the Monkey Bay fisheries sta- 15-40 fths (28-74m), but was occasion- tions since the 1960s. It is much deeper ally found as deep as 75 fths (140m). bodied than the true Lethrinops albus In the 1992 trawl survey, it comprised Regan, which was described from ma- a larger proportion of the sample bio- terial from unknown locations. L. albus mass than any other cichlid (6.5%) and is probably a species of much shal- it was found to be one of the dominant lower waters. Konings (1990a) illus- species at depths of 40-70m. It was not trates a species he identifies as L. albus, found deeper than 95m. On the east which is found in shallow water coast north of Boadzulu Island its around Kande Island. range extends to water as shallow as 18m, while on the west coast it has not been recorded at less than 45m. Lethrinops longipinnis COMMERCIAL IMPORTANCE: Occurs in Eccles & Lewis the catches of all types of trawlers. In colour photos on page 57 and 58 1990-91, it comprised 5% of deep wa- ter bottom trawl catch and 2% of DISTINGUISHING FEATURES: A deep-bod- midwater and pair trawl catches. ied, laterally compressed species with NOTES: According to FAO (1976) and a long steeply sloping snout, and a Tweddle & Turner (1977), log (weight) small ventrally-placed mouth. It differs = 2.644 log (length)- 3.045, K = 0.571, from L. lethrinus in its lack of dark Linf = 201 mm, maximum length caught markings. The snout is longer than that = 214 mm, length at maturity = 166 of L. 'deep-water albus'. mm, natural mortality = 1.69, and log COLOUR: Females and immature (fecundity) = 2.641 log (length, mm) - males silvery with yellowish pectoral 3.779. It is possible that deep-water and fins. In deep water, ripe males are dark, shallow water specimens represent dif- with strong vertical barring, dark fins ferent species. The red nape may also and a bluish iridescence on the head. be species-specific. These have not In shallower water, males are more been investigated in detail. L. argenteus brownish and less strongly barred. Ahl is very similar, but has not been Some specimens have a dark reddish seen since the collection of the types at nape. the northern end of the lake (Eccles & MAXIMUM SIZE: 21 cm TL. Lewis, 1978). DIET: Mainly chironomid larvae (Ec- cles & Lewis, 1978), but also copepods, algae, and detritus (Grant & Turner, unpubl., 1 specimen examined). DISTRIBUTION AND ABUNDANCE: Abun-

88 Lethrinops mylodon Eccles & Lewis Konings (1990a, 1995b) regarded the colour photo on page 58 difference in pharyngeal bone shape as sufficient to elevate these two forms to DISTINGUISHING FEATURES: A very deep- the level of species. Given the known bodied species with a large head and phenotypic plasticity and polymor- eye, it is distinguished from L. gossei phism of pharyngeal structures in by the absence of a V-shaped cleft in cichlids (Kornfield 1991), even sub- the upper jaw and by the lower specific status seems dubious. Eccles number of lower gillrakers (12-14 com- & Lewis's material consisted of twenty pared to 18-20). It has a higher gillraker specimens from Monkey Bay but only count than L. 'deep-water albus' and 9 from Nkhata Bay, 7 of which were differs from all other deep-bodied mislaid after the description. This Lethrinops species in the massively en- number is perhaps insufficient to in- larged molariform pharyngeal teeth. dicate sympatric . The COLOUR: Females and immature Nkhata Bay material was collected in males silvery, with a golden sheen, and 1960-61, and Eccles & Lewis did not 7 faint bars under the dorsal fin. Ripe observe the live colours of the speci- males are golden-bronze with some- mens. Konings has not observed the times prominent vertical bars, a bright northern form in the field, and appar- blue head and numerous large yellow ently nor has anyone else. egg-spots on the anal fin. MAXIMUM SIZE: 25 cm TL (Eccles & Lewis, 1979). Lethrinops macracanthus (Trewavas) DIET: Snails (Eccles & Lewis, 1979). colour photo on page 58 DISTRIBUTION AND ABUNDANCE: Eccles & Lewis (1979) recorded the species from DISTINGUISHING FEATURES: Large head the SE Arm, Nkhotakota, and Nkhata and eye. Deep body. Distinguished Bay, from depths of 40 to 70m in the from L. gossei by the absence of a V- SE Arm, and as deep as 110m off shaped cleft in the upper jaw. Differs Nkhotakota. It was seen between 48 from L. 'deep-water albus', L. longi- and 64m in 1992 surveys, but was rare. manus, and L. mylodon in its larger I also collected this species from number of gillrakers (usually 19-23 Domira Bay. ceratobranchial) and vertical bars (usu- COMMERCIAL IMPORTANCE: Minor. ally 8-9 under the dorsal fin) and from NOTES: Populations in the SE Arm the latter in the less heavily molariform have declined drastically since the pharyngeal dentition. early 1970s. Eccles & Lewis (1979) de- COLOUR: Females and immature scribed a northern subspecies L. m. males silvery, and 8-9 (occasionally 7) borealis on the basis of its lower pha- dark bars under the dorsal fin. Orange- ryngeal bone, which is more elongated brown spots on flank scales and dor- and with a markedly concave lower sal fin. Ripe males coppery with verti- surface. This form was collected from cal bars more prominent and a bluish Nkhata Bay, while L. m. mylodon is re- iridescence on the head. corded as far north as Nkhotakota. MAXIMUM SIZE: 19 cm SL (Eccles &

89 Trewavas), ca 25 cm TL. from L. gossei by the absence of a V- DIET: Chironomids. Also diatoms, shaped cleft in the upper jaw. Differs small snails, ostracods, copepods and from L. 'deep-water albus' in larger annelids (Eccles & Lewis, 1979). number of gill rakers. Distinguished DISTRIBUTION AND ABUNDANCE: Holo- from Lethrinops macracanthus by the type is from Monkey Bay. Eccles & lower number of gillrakers (usually 15- Lewis (1979) examined material from 18) and vertical bars (usually 7). L. SE Arm, Domira Bay, and Nkhata Bay longimanus is generally darker and and stated that this species was wide- more strongly barred. spread in southern Malawi, often COLOUR: Females and immature forming a substantial proportion of the males silvery with pinkish iridescence, catch at 20-35 fathoms (37-65m), but and 7 dark bars under the dorsal fin. was also occasionally found as deep as Ripe males dark grey with bronze iri- 40 fathoms (74m). This depth range has descence, blue head. Vertical bars been heavily exploited by stern trawl- prominent. ers and FAO (1976) reported that in MAXIMUM SIZE: 15 cm SL (Eccles & 1972, L. macracanthus had comprised Trewavas), ca 20 cm TL. 10-15% of the experimental trawl catch DIET: Benthic invertebrates, espe- weight in the SE Arm in 1972, it had cially chironomids (Eccles & Lewis declined rapidly and was virtually ab- 1979). sent by 1974. It did not occur in the DISTRIBUTION AND ABUNDANCE: Holo- 1992 survey. I found a single large type from the SW Arm, but material specimen in a haul from 22-30m south examined by Eccles & Lewis was from of Boadzulu Island made as part of the the SE Arm, and they also recorded it photographic survey. from Nkhotakota and Nkhata Bay, at COMMERCIAL IMPORTANCE: It was for- depths of 20-80m (Eccles & Lewis merly significant in trawl catches, but 1979). Stated to form a substantial part is now too rare to be of commercial of the catch on occasions. In 1992 sur- importance, at least in the SE Arm. veys of the SE Arm it was frequently NOTES: Length-weight relationship is taken and often a dominant species given by FAO (1976) as log (wt)= 2.647 between 50 and 72m. I also collected x log (len) - 3.055. Eccles & Lewis (1979) this species from Domira Bay. reported that some specimens from COMMERCIAL IMPORTANCE: Minor com- Sungu Spit (near Nkhotakota) were ponent of trawl catches. intermediate between L. macracanthus NOTES: According to FAO (1976), log and L. longimanus. Both of these spe- (wt)= 2.935 x log (len) - 3.689, natural cies were found at the site and also to mortality = 1.54, maximum length the north and south. caught = 212 mm. Konings (1990a) il- lustrates and discusses a species with Lethrinops longimanus Trewavas a similar body shape, but fewer gill- colour photo on page 58 rakers, under the name L. sp. 'Longi- manus'. It is known only from Likoma DISTINGUISHING FEATURES: Large head Island and is probably an undescribed and eye. Deep body. Distinguished species.

90 Lethrinops macrochir (Regan) samples, from 18-24m at Fowo, colour photo on page 59 Namiasi, and Michesi. It was not abun- dant. It was also seen in catches south DISTINGUISHING FEATURES: A typical of Boadzulu Island during the photo- Lethrinops in shape. Distinguished graphic survey and in artisanal catches from L. cf parvidens and Tramitichromis from Lake Malombe. lituris by the higher number of gill- COMMERCIAL IMPORTANCE: Taken in rakers (11-14), and from these species fairly small numbers by beach seines and L. 'pink head' by the bright yellow in the SE Arm. It is estimated that in ventral colour. It has a much shorter 1990-91 seine fishermen on Lake Ma- snout than other species with a simi- lombe landed around 150 tonnes of this lar colour, such as Taeniolethrinops prae- species, making it the 6th most impor- orbitalis and T. furcicauda. tant haplochromine species in the fish- COLOUR: Females and immature ery. males silvery with bright yellow head NOTES: According to Eccles & Lewis and ventral surface. Faint narrow ver- (1979), ripe males are similar to fe- tical bars, the upper parts of bars 2-4 males, but with a bluish head (Eccles widened to give the impression of a & Lewis, 1979). This could either rep- suprapectoral spot. Ripe males (several resent geographic variation, or more specimens taken from a seine net catch probably, fish which were not fully in Lake Malombe): generally dark pur- ripe. Trewavas (1931) considered this plish-brown, with 8 vertical bars un- species to be conspecific with L. auritus, der the dorsal fin, and a dark blotch but Eccles & Lewis disagreed, point- between bars 2-4. Snout bluish. Fins ing out the smaller size at maturity of dark, spotted. Dorsal fin margin bright that species. yellow orange. Anal fin with a row of whitish yellow egg-spots on ventral margin, which is tipped bright red. Lethrinops 'loweae' MAXIMUM SIZE: 12cm SL (Eccles & Lewis, 1979), ca 16cm TL. DISTINGUISHING FEATURES: A relatively DIET: Benthic invertebrates, espe- deep-bodied species with a short snout cially chironomids (Eccles & Lewis, and strong vertical barring. It has 11- 1979). A single specimen examined by 12 gillrakers, more than L. cf. parvidens Grant & Turner (unpubl.) contained and Tramitichromis lituris and fewer chironomids, diatoms, and detritus. than L. macracanthus, L. longimanus, L. DISTRIBUTION AND ABUNDANCE: The 'oliveri', L. 'yellow chin', L. 'micrent- types are from unknown locations. odon Makokola', L. micrentodon, L. Eccles & Lewis (1979) recorded the spe- stridei and L. microdon. L. 'pink head' cies from Lake Malombe, SE and SW and L. macrochir lack the strong verti- Arms and Domira Bay, although all the cal bars and species of the L. longipinnis specimens they measured were from complex have a more pointed snout. the SE Arm. Eccles & Lewis report a COLOUR: Pale with 6-7 dark vertical depth range of 5-10 fathoms (9-18m). bars. Fins dark. Mature males darker In the 1992 survey it occurred in four overall. Live colours not known.

91 Figure 24. Lethrinops 'lowae', male, South-East Arm.

MAXIMUM SIZE: 17cm TL. deep-bodied species with a large head DISTRIBUTION AND ABUNDANCE: Re- and mouth. It has more ceratobranchial corded only from the northern part of gillrakers (14) and fewer vertical bars the SE Arm from the stations Chekopa (5-6) than L. cf. parvidens, L. 'pink head', (60-66m), Fowo (64m), Chilinda (66m), Tramitichromis liturus and L. 'bighead'. Monkey Bay (92m) and Ilala Gap (22- COLOUR: Pale sandy with 5-6 brown- 80m). Rare. ish vertical bars under the dorsal fin. COMMERCIAL IMPORTANCE: Negligible. Dorsal fin lappets dark. Ripe males not recorded. Lethrinops 'macrostoma' MAXIMUM SIZE: 11cm TL. colour photo on page 59 DISTRIBUTION AND ABUNDANCE: Rare. Recorded from a single sample at 18m DISTINGUISHING FEATURES: A small, pale off Fowo and taken by the photo sur-

Figure 25. Lethrinops 'bighead', male, South-East Arm.

92 vey from 40-50m off Namalaka. COLOUR: Females and immature COMMERCIAL IMPORTANCE: Minor, al- males silvery. Ripe males with bright though the focus of its distribution is blue snout, pale blue opercula. The heavily fished by trawlers. upper part of the head and the nape are pinkish-orange. There are 8-9 thin dark vertical bars under the dorsal fin, Lethrinops 'bighead' with a faint blotch across the upper part of 3 bars. Dorsal fin with white DISTINGUISHING FEATURES: Relatively lappets and orange tips. elongate body. Large head and mouth. MAXIMUM SIZE: 12cm TL. Fewer gillrakers (11) and more verti- REPRODUCTION: Matures at small sizes cal bars (7) than L. 'macrostoma'. in Lake Malombe: some ripe females Larger head than L. cf. parvidens, L. being less than 6cm TL (Table 3). Ap- 'pink head' and T. lituris. proximately 30 eggs laid. COLOUR: Pale sandy with darker DIET: 14 specimens from Lake brownish vertical bars. Ripe males Malombe examined (Mwanyama & darker with broad white dorsal mar- Turner, unpubl.). Major items were: gin and dark submarginal band, dark cladocerans (dominant in 5 individu- anal and pelvic fins. There is a dark als), copepods (4), odonata (3), chiro- stripe on the preorbital and the chin nomids (1). Algal remains, detritus and and chest are also dark. sand were found in all stomachs. The MAXIMUM SIZE: 10cm TL. stomach of a single specimen from the DISTRIBUTION AND ABUNDANCE: Known SE Arm of Lake Malawi contained from a single preserved sample, for cladocera, diatoms and detritus (Grant which the label was in poor condition, & Turner, unpubl.). but appears to have been from Ulande DISTRIBUTION AND ABUNDANCE: In the at 20m depth. It was quite numerous 1992 trawl survey it was numerous in in this sample. three samples at 20-28m at Namiasi COMMERCIAL IMPORTANCE: May con- and Fowo, but was otherwise not re- tribute to seine and pair trawl catches. corded. In the photographic surveys it as also recorded between 5 and 28m near Namiasi. It is abundant in Lake Lethrinops 'pink head' Malombe. colour photo on page 59 COMMERCIAL IMPORTANCE: More than 2,500 tonnes were taken annually by DISTINGUISHING FEATURES: An average seines in Lake Malombe 1990, where it looking Lethrinops species which is dis- was the most important commercial tinguished from L. cf parvidens and Tra- species. I examined 3,488 fish caught mitichromis lituris by the higher num- by the artisanal seine fishery in 1991- ber of gillrakers (12-13). It lacks the 92. The mean individual fish weight bright yellow ventral colour of L. was 3g. The estimated annual catch, macrochir. Placidochromis 'longimanus from Lake Malombe alone, is therefore Malombe' has a different dental arcade approximately 850 million fish. and male breeding colour. NOTES: It is possible that this species

93 is conspecific with L. auritus, but Ec- slightly deeper body and absence of cles & Lewis (1979) in their redescrip- broad flat first gillraker. tion of this species report a different COLOUR: Females and immature male breeding colour and illustrate a males silvery with sandy cast. Thin ir- different gillraker form. However, at regular bars sometimes present, often least one of the specimens used in the coalescing into a faint spot dorsally. Ripe males with a copper cast, par- ticularly on the nape. Fins dark, spotted. Anal fin with several large Total Immature Active Ripe/Spent Mature pale egg-spots. Length unsexed Females Females Males MAXIMUM SIZE: 18cm TL. 5.0-5.4 17 — — 6 DIET: According to Eccles & 5.5-5.9 25 4 2 5 Lewis (1979), benthic invertebrates, 6.0-6.4 42 9 7 11 especially chironomids, and also 6.5-6.9 28 13 4 12 7.0-7.4 30 10 1 4 diatoms. Occasionally copepods 7.5-7.9 27 3 1 4 and bivalves. The stomach of an in- 8.0-8.4 2 3 7 4 dividual, of 9.5cm SL, collected off 8.5-8.9 — 3 1 — Namiasi in July 1992 contained mostly algae, detritus, and sand Table 3. Size at Maturity, Lethrinops 'pink head', Lake grains. A 10cm specimen taken off Malombe, October 1991 Chilinda had been feeding mainly on ephemeropteran nymphs and description (deposited in the Monkey chironomid larvae (Grant & Turner, Bay field museum) has a gillraker form unpubl.). identical to L. 'pink head'. The male DISTRIBUTION AND ABUNDANCE: Eccles & breeding colours attributed by Eccles Lewis (1979) recorded this species from & Trewavas to Placidochromis longi- all round the SE Arm and northwards manus are identical to those of L. 'pink to Nkhata Bay, and also from the Mo- head' and differ from the P. longimanus zambique coast. They found it was of- I have observed. This is probably due ten abundant at depths of 5-20 fathoms to an error in identification — the two (9-37m), but that it was recorded as species have similar meristics and deep as 35 fathoms (65m). In the 1992 morphometrics. survey, it was taken in most samples throughout the SE Arm and was over- all the fourth most important (in terms Lethrinops cf. parvidens of biomass) cichlid in the survey. It was colour photo on page 59 abundant at 30-45m, and common from 18-60m. Occasional records from DISTINGUISHING FEATURES: An 'average' as deep as 100m may represent mis- looking Lethrinops species, with a rela- identification. tively elongate body. Distinguished COMMERCIAL IMPORTANCE: Around from L. 'pink head' and L. macrochir by 1.5% of midwater trawl catch. Likely the lower number of gillrakers (8-11), to be important in pair trawl catches and from Tramitichromis lituris by the in some areas.

94 NOTES: This species formed the basis MAXIMUM SIZE: 20cm TL. of Eccles & Lewis's (1979) redescription DISTRIBUTION AND ABUNDANCE: Eccles & of Lethrinops parvidens. FAO (1976) give Trewavas report of L. furcifer from a the following parameters: log (wt, g)= single trawl station at 5 fathoms (9m) 3.187 log (len, mm) - 4.269; K= 0.487; depth probably refers to this species.

Linf = 208mm; max. size caught = In the 1992 survey, it occurred at 18- 204mm; length at maturity = 153mm, 28m depth at the Fowo, Ulande, and natural mortality = 1.20; log (fecundity) Chekopa stations in the SE Arm, and = 2.236 log (len, mm) -2.901. It is prob- in photographic survey hauls from the able that this information refers to L. same depth range south of Boadzulu cf. parvidens. I have examined Eccles & Island, as well as from deeper water Lewis's voucher specimens deposited (50m) between Chirombo and at the BMNH and at Monkey Bay Fish- Nkhudzi. eries Station and others I have collected COMMERCIAL IMPORTANCE: Around 1% from the same localities. I have also of midwater trawl catch. Likely to be measured some of the types of L. important in pair trawl catches in some parvidens Trewavas. I believe that the areas. material collected by Eccles & Lewis is NOTES: FAO (1976) give a length not conspecific with the type series of weight relationship for Lethrinops fur- L. parvidens, which itself may include cifer which probably refers to the L. cf. more than one species. There is no des- furcifer I have collected: log (wt)= 2.502 ignated lectotype of L. parvidens and log (len)-2.829. I referred the specimens the collecting location is unknown. I collected to L. furcifer as a result of Some of the types appear to be sexu- comparison with voucher specimens in ally mature males, which differ in pat- the Monkey Bay Fisheries Station col- tern from those of L. cf. parvidens and lection, which probably means they are are also much smaller. They differ in conspecific with those identified by multivariate shape analysis, but sim- Eccles and by FAO. Lethrinops furcifer ple counts and ratios are all overlap- Trewavas is very poorly known. It was ping (Best & Turner, unpubl.). originally described from 12 specimens by Trewavas in 1931. No collecting data or illustration is given and the descrip- Lethrinops cf furcifer tion is very fragmentary, comprising colour photo on page 59 only meristics and two morphometric ratios. Eccles & Trewavas did not re- DISTINGUISHING FEATURES: Very similar examine this species, but gave a diag- to L. cf. parvidens, but attains a larger nosis apparently based on the original size and is slightly more elongate. 8- Trewavas description and provided a 12 lower gillrakers. drawing of a specimen which appears COLOUR: Females and immature to be a mature male. Both Fryer (1959) males silvery with a sandy cast. Ripe and Konings (1990a) provide informa- males coppery particularly on the tion on a small shallow water species nape. Snout and cheeks blue. Anal fin from the north of the lake which they with several large pale egg-spots. state is L. furcifer. Fryer (1959) reports

95 that Lethrinops furcifer is the common- Tramitichromis lituris (Trewavas) est species on sandy shores in the north colour photo on page 60 of the lake, that it attains at least 15.5cm and that it feeds on chironomids. He DISTINGUISHING FEATURES: A typical also reports another species which can Lethrinops shape, very similar to L. cf be distinguished only by its higher gill- parvidens, but differs slightly in head raker count (16-17) and finer pharyn- shape and in the shape of the anterior- geal teeth. Konings (1990a) states that most ceratobranchial gillraker which is L. furcifer is found throughout the lake, long and flat. Distinguished from L. is most abundant at 7m depth, and il- macrochir and L. 'pink head' by the lustrates specimens from Ruarwe, lower number of gillrakers (6-10). The Chitande Island and Senga Bay. He lower pharyngeal bone of this genus states that it is a colonial breeder dur- is distinctive (see Eccles & Trewavas, ing the months of May to September 1989). Differs from other Tramitichromis (and perhaps at other seasons) which spp. in the melanin pattern which lacks constructs sand-castle nests. The speci- horizontal or oblique elements or mens he illustrates do not seem to be spots. conspecific with the L. cf. furcifer I have COLOUR: Females and immature collected. I have not examined the males silvery, with brownish yellow types of L. furcifer, but in the illustra- cast and faint vertical bars, with a faint tion by Eccles & Trewavas, the head blotch joining three bars. Ripe males length is approximately 28% SL (pro- dark blue, with metallic orange spots jection measurement from figure) on flank scales. Nape orange. Dorsal while in Konings' illustrations, the lappets white with orange tips. Throat head length is 33-35% SL and in the orange or red. specimens of L. cf. furcifer I have col- MAXIMUM SIZE: 18cm TL. lected, it is 34-35% SL. Konings' illus- DIET: Chironomid larvae and algae tration of a specimen from Senga Bay were found in the single individual has a strong hint of an oblique bar par- examined (Grant & Turner, unpubl.). tially obscured by the male breeding REPRODUCTION: Digs a pit next to a colour. This is not apparent in L. cf. fur- small rock, in 7-15m depth (Konings cifer, nor the specimen figured by Ec- 1992). cles & Trewavas, nor is it mentioned DISTRIBUTION AND ABUNDANCE: The in Trewavas' description. Trewavas type locality is unknown. The species does, however, mention a dark flank was not known to Jackson, except spot. The specimens illustrated by through museum specimens. Eccles & Konings (1991c) as Taeniolethrinops Trewavas report that the species was 'furcicauda Ntekete' may be conspe- caught in trawl catches at 10 fathoms cific with L. cf. furcifer (see under T. (18m) off Chilinda in the SE Arm and furcicauda, below). near Malembo. It was also abundant in a trawl taken at 5 fathoms (9m) on the western shore at 'the narrowest part of the lake' which probably means near Salima. In the 1992 survey it was

96 often abundant in trawl samples be- Taeniolethrinops furcicauda tween 20 and 32m, but was also re- (Trewavas) corded in shallower water. Not known colour photo on page 61 from Lake Malombe, but similar speci- mens have been collected in the mid- DISTINGUISHING FEATURES: One of a dle Shire River. group of species characterised by a COMMERCIAL IMPORTANCE: Important in long snout and ventral mouth. Distin- pair trawl and seine catches north of guished from T. praeorbitalis and T. Boadzulu Island. laticeps by its shorter snout, and from NOTES: The last taxonomic study of L. cf parvidens and L. furcifer by its this species was by Trewavas in 1931. longer snout and bright yellow ventral The species is poorly defined and may coloration. represent a complex of related species. COLOUR: Females and immature In 1990, Konings stated that T. lituris is males dark grey dorsally, with silvery common in the south of the lake and flanks, marked by 7 faint irregular ver- illustrates specimens from Senga Bay, tical bars. Dorsal and caudal fins dark but in 1992, decided that these repre- with dark grey spots. Lower part of the sented a different species. 'Lethrinops body, pelvic, anal and lower part of red flush', which he suspects might be caudal fins bright yellow. T. variabilis, but the specimens he illus- MAXIMUM SIZE: 21cm TL (Konings trates seem to be conspecific with those 1990a). identified by Eccles & Trewavas, DIET: A benthic feeder. Stomach con- Stauffer (pers. comm.), and by myself tents dominated by chironomid larvae as T. lituris. Eccles & Trewavas state and copepods, along with sand detri- that the types of T. variabilis have a tus and some algal material. Occasion- melanin pattern which either consists ally also other insect larvae and bi- of a partial oblique band or a series of valves. (5 specimens of 7.5-16cm SL spots. Neither pattern is present in the examined by Grant & Turner, unpubl.). specimens I have assigned to T. lituris. DISTRIBUTION AND ABUNDANCE: Eccles & Konings (1992) reports what he regards Trewavas recorded this species from 5- as the true T. lituris at 7-15m depth at 20 fathoms (9-37m) in their trawl sur- Mdoka in the far north of the lake. He veys. In the 1992 survey, it was taken (pers. comm.) has examined preserved in most samples at 18-32m throughout material of both the Mdoka and 'red the SE Arm. It was sometimes the flush' forms and find that the former dominant species in terms of biomass. is more similar to the description pro- Occasionally taken in small numbers vided by Eccles & Trewavas. The male as deep as 55m. I have not seen this is bright blue and lacks the orange species while diving or snorkelling nape seen in southern specimens. It shallow water. was a colonial nester, excavating a pit COMMERCIAL IMPORTANCE: It is caught under a rock or occasionally building by pair and semipelagic trawls, gill a sand platform on top a rock. Further nets, and occasionally by seines. work is required to determine which NOTES: My identification follows ex- (if either) is the true T. lituris. amination of voucher specimens in the

97 Monkey Bay Fisheries Station. FAO furcicauda. The bright yellow ventral (1976), probably following the same colour and lack of a continuous black guide, give a length-weight relation- oblique stripe distinguish this species ship of log (g) = 2.725 log (mm) -3.329. from T. laticeps, at least in the south of The type specimens of this species are the lake. T. laticeps has a more thickset said by Eccles & Trewavas (1989) to build with a broader head. L. christyi have a dark oblique bar and Konings also has a long snout, but the mouth is (1990a) illustrates a barred specimen. less ventrally placed, and it lacks the Like the voucher specimens at Mon- yellow coloration of T. praeorbitalis. key Bay, the specimens from the SE COLOUR: Females and immature Arm which I have tentatively assigned males dark grey dorsally, with silvery to this species do not have an oblique flanks, marked by 7 faint irregular ver- bar. I have examined some preserved tical bars. Dorsal and caudal fins dark specimens from Nkhata Bay which with dark grey spots. Lower part of the show a clear, unbroken bar. There may body, pelvic, anal and lower part of be more than one species, and a re-ex- caudal fins bright yellow. Ripe males amination of the types is required. generally dark blue, flanks with golden Konings (1991c) discusses a species hue ventrally, sometimes with 7-9 faint under the name Taeniolethrinops sp. dark bars visible. The dorsal fin has 'furcicauda Ntekete'. The female illus- white lappets. Anal fin with 20 or more trated resembles T. laticeps in its promi- orange egg spots. nent oblique bar, but the angle of the MAXIMUM SIZE: 30cm TL (Konings photograph makes it difficult to assess 1990a). the snout length. The male illustrated DIET: In shallow water this species resembles males I have assigned to can be observed repeatedly plunging Lethrinops cf furcifer, having a relatively its snout into soft sediment, and sift- large eye, short snout, and a blue head ing material through its opercula. Ac- and orange nape. Males defended a cording to Jackson, it feeds mostly on large (1m diameter) crater nest at 15m Chaoborus larvae. Eccles & Trewavas depth. Konings considered them to be state that chironomids make up most an undescribed species. The collecting of the diet. Stomach contents of 3 speci- locality was near the village of Ntekete mens (12-17cm SL) indicate large between Makanjila Point and the Mo- amounts of sand and detritus are in- zambique border (Konings 1995b). gested, along with diatom material and chironomid larvae (Grant & Turner, unpublished). Taeniolethrinops praeorbitalis REPRODUCTION: Konings (1990a) has (Regan) observed males defending large (2.5m colour photo on page 60 in diameter) sand castle nests at 20m depth during July and August. DISTINGUISHING FEATURES: One of a DISTRIBUTION AND ABUNDANCE: Jackson group of species characterised by a reports that it is found as deep as 40m long snout and ventrally placed off Monkey Bay and that it is common mouth, it has a longer snout than T. in seine net catches off Nkhotakota.

98 Eccles & Trewavas reported that it was onymised Lethrinops macrorhynchus common in trawl catches from 5-20 Regan and L. fasciatus Ahl with this fathoms (9-37m), but also recorded it species. I have examined the type of L. from as deep as 30 fths (55m) off Mbenji macrorhynchus, which has a strong dark Island. Konings (1990a) states that it is oblique band. It is difficult to make common over open sand in shallow accurate morphological comparisons, water throughout the lake. In the 1992 because the specimen has been cut in survey, it was frequently caught in half, but the overall shape resembles trawls between 18 and 32m in the SE T. praeorbitalis more than T. laticeps. I Arm, and I have often observed this have examined some small specimens species while diving or snorkelling in collected from the north of lake (near shallow water in the south of the lake. Nkhata Bay) by the JFRO. These ap- In 1990-91, it was taken in experimen- pear to be very similar to L. macro- tal fyke nets and gill nets in the Upper rhynchus in shape and colour pattern, Shire River, but was not observed in as does a photograph of T. 'laticeps seine catches in Lake Malombe. itungi' from the Tanzanian coast COMMERCIAL IMPORTANCE: In 1990-91 in (Konings 1995b). I have not examined the SE Arm, it was caught in midwa- the types of L. fasciatus. ter trawl (about 1% of the non-tilapia catch), deep water stern trawl (rare), pair trawl, gillnet, and seine. Tramitichromis brevis (Boulenger) NOTES: Length-weight relationship given by FAO (1976) as log (g) = 2.831 See chapter 14, Mylochromis and log (mm)- 3.52, but this information other oblique-striped species. was almost certainly obtained from a mixture of T. praeorbitalis and T. laticeps. Taeniolethrinops laticeps (Trewavas) Eccles & Trewavas (1989) designated this as the type species of the genus See chapter 14, Mylochromis and Taeniolethrinops which was separated other oblique-striped species. from Lethrinops on the basis of the pos- session of an oblique dark bar. How- ever, examination of the syntypes of Lethrinops lethrinus (Günther) this species, indicates no evidence of such a pattern. The SE Arm specimens See chapter 16, Protomelas and other which I have assigned to this species horizontally-striped species. do not generally exhibit an oblique bar although similar individuals observed underwater may develop a faint, bro- Tramitichromis intermedius ken bar according to mood. All speci- (Trewavas) mens I have examined from the south- ern part of the lake which have a con- See chapter 15, Otopharynx and other tinuous black oblique stripe are mor- spotted species. phologically more similar to the type of T. laticeps. Trewavas (1931) syn-

99 Chapter 10 Aulonocara and Alticorpus

Species of these two genera are char- from the north-western rocky shore acterised by a melanin pattern of ver- from Chilumba to Chinteche. Males are tical bars and enlarged cephalic pores strongly barred with a blue head and of the lateral line system. These are a thin white dorsal fin margin. North- most prominent as large pits visible on ern specimens have a patch of orange inspecting the underside of the lower behind the head, while southern speci- jaw. The expansion of these pores ap- mens only have orange on the nape. pears to be associated with a strategy Aulonocara brevinidus Konings, which of feeding on concealed prey, often at attains 11.5cm TL, is found on the east low light levels. Shallow water species coast from Tanzania to Masinje. It has can often be seen around dusk, hover- a bluish head, 7 vertical bars, large ing in a slightly head-down position bright yellow egg-spots, and a dark just above the substrate, occasionally upper and lower part of the caudal fin. darting into the substrate, presumably None of these species were positively to capture small burrowing inverte- identified in my collections and they brates. Many Aulonocara species are may not occur in the SE Arm. confined to rocky shores, some living Stauffer & McKaye (1988) distin- in caves underneath rocks, others guished Alticorpus from Aulonocara on spending most of their time foraging the basis of the prominent mental proc- over sand. ess, but in my view this trait is vari- Subsequent to my field work, Ko- able within as well as between species. nings (1995b) described 3 new species Alticorpus species are confined to deep of Aulonocara from shallow sandy water, and many have large mouths areas. Aulonocara aquilonium Konings and appear to be piscivorous. known only from Mdoka, north of In the 1992 trawl survey, the genera Chilumba, is a small slender species Aulonocara and Alticorpus comprised with 7 vertical bars under the dorsal 6% and 2.6% respectively of the total fin, a bluish head and a dorsal fin with sample biomass. Deep water areas bright yellow spots, black lappets and were under-represented in the survey, a white submarginal band. Aulonocara so it is likely that Alticorpus, which is gertrudae Konings, a medium sized predominantly found in deeper water, species attaining around 13.5cm TL, is actually more significant than this has 9 vertical bars and numerous egg- figure suggests. North of Monkey Bay, spots on the anal fin of females as well they represented 6.5% compared to as males. It is known from the eastern 3.2% for Aulonocara. coast all the way from the north of the Tanzanian coast to Masinje, just north of Makanjila Point. It is also known

100 Alticorpus mentale nus fryeri' in trawling records from the Stauffer & McKaye Monkey Bay Fisheries Station. Under colour photo on page 61 that name FAO (1976) give a length- weight relationship of log (weight) = DISTINGUISHING FEATURES: A large- 3.222 x log (length) -4.322. Four speci- mouthed species, with small eyes and mens of Alticorpus 'dwarf mentale' enlarged cephalic pits and a prominent were collected from Domwe at 128m, mental process. Juveniles are distin- the deepest station in the 1992 survey. guished from most other Alticorpus They resembled A. mentale in body spp. by the low number of gillrakers shape, enlarged cephalic pits, gillraker (9-12). Adults are larger than mature count (8-10 ceratobranchials) and male A. 'geoffreyi' and differ in colour. A. breeding colour (dark bars with white 'geoffreyi' has a relatively larger eye dorsal fin lappets), but ripe males were and smaller mouth. only 11-12.5cm TL. COLOUR: Females and immature males silvery-white. Ripe males gen- erally dark grey to black, silvery on Alticorpus 'geoffreyi' flanks with 6 prominent black vertical colour photo on page 61 bars. Unpaired fins and pelvic fins black. Dorsal fin lappets white. Anal DISTINGUISHING FEATURES: A large- fin with 7 large pale egg-spots. mouthed species, with expanded ce- MAXIMUM SIZE: 20cm SL (Eccles & phalic pits and a prominent mental Trewavas), approximately 25cm TL. process. Distinguished from most DISTRIBUTION AND ABUNDANCE: Eccles & other Alticorpus spp. by the low Trewavas report that the species is number of gillrakers (9-12) and from abundant at depths of 20-70 fathoms A. mentale by its smaller mouth, rela- (37-130m) in the southern part of the tively larger eye, smaller size at matu- lake, and have recorded the species rity, and colour. from 10 fathoms (18m). In the 1992 sur- COLOUR: Females and immature vey, it was common from 60 to 128m, males silver, darker dorsally, with a and recorded twice at 46m in the SE pronounced brownish-yellow cast, and SW Arms. It was never seen in and 7 faint vertical bars. Ripe males shallower water. have silvery flanks and 6-7 prominent COMMERCIAL IMPORTANCE: Occurred in vertical bars. The dorsal fin is dark yel- 42% of the stern bottom trawl catches low-brown. Caudal fin dark brown. sampled in 1990-91, but comprised no Anal fin dark with several very large more than 0.5% of the catch weight. It yellow streaks and spots. Pelvic fins may initially be an important species dark. Snout and cheeks with dark me- should trawling be extended to deeper tallic blue iridescence. water, although its large size at matu- MAXIMUM SIZE: 20cm TL. rity means it is likely to be rapidly DISTRIBUTION AND ABUNDANCE: In the eliminated by heavy fishing. 1992 trawl survey, it was the most nu- NOTES: Eccles & Trewavas report that merous Alticorpus species. Found at 54 the species was known as 'Trematocra- to 128m, but most numerous in the

101 hauls from deeper water and compris- 52m, but included a mixture of species ing on occasion a substantial propor- typical of deep and shallow waters. tion of the catch — 23% by weight in COMMERCIAL IMPORTANCE: Negligible. one haul at 100m depth at the Ngusi NOTES: Previously unknown and not station. distinguished from A. 'geoffreyi' by the COMMERCIAL IMPORTANCE: Contributes author until 1992, it is still not abso- to stern bottom trawl (1.5% by weight), lutely certain that these species are dis- and occasionally to midwater trawl tinct. catches. NOTES: Under the name 'Trematocra- nus geoffreyi', FAO (1976) give the Alticorpus pectinatum length-weight relationship as log Stauffer & McKaye (weight) = 3.114 x log (length) -4.090. colour photo on page 61

DISTINGUISHING FEATURES: A deep-bod- Alticorpus 'deep' ied species with prominent cephalic colour photo on page 61 pits, a mental process and a relatively large number of gillrakers (17-18 DISTINGUISHING FEATURES: A deep-bod- ceratobranchials). Alticorpus peter- ied species with enlarged cephalic pits, daviesi differs in male breeding colour and a prominent mental process. Dis- and has a more pointed snout. Ripe tinguished from A. mentale and A. males are easily identified by their bril- 'geoffreyi' by the deeper body and liant yellow breeding dress. smaller mouth and from A. profundi- COLOUR: Females and immature cola, A. peterdaviesi and A. pectinatum males silver, darker dorsally with by the larger mouth and fewer lower traces of 7-8 faint vertical bars. Ripe gillrakers (10-12). males golden-brown with dark brown COLOUR: Females and immature vertical bars. Head and nape bright males silver, darker dorsally with yellow. Chin and cheeks with metallic traces of faint vertical bars. Ripe male pale blue iridescence. Dorsal fin yel- with 8 dark vertical bars under the low-brown, with white lappets and a dorsal fin, black dorsal fin lappets, a thin black submarginal band. Caudal bluish iridescence on the cheeks and fin yellow-brown. Anal fin dark grey several large whitish-yellow irregular to black with several large yellow- spots on the anal fin. Few specimens white spots and streaks. Pelvic fins or- have been seen, and so this may not be ange brown, with a broad black ante- fully-developed breeding colours. rior margin. Pectoral fins brownish. MAXIMUM SIZE: 16cm TL. MAXIMUM SIZE: 16cm TL. DISTRIBUTION AND ABUNDANCE: A rare, DISTRIBUTION AND ABUNDANCE: In the deep-water species known only from 1992 surveys, this species was taken in the northern part of the SE Arm, from all samples from 92 to 128m (the deep- a sample at 102m at Ngusi, 72m off est haul), often in large numbers. It Monkey Bay and a haul near the Ilala comprised 27% of the catch weight in Gap which was reported to be from one sample from 118m depth at Ngusi

102 station. It occurred sporadically in both are valid species. Without exam- small numbers at depths as shallow as ining the types, I have assumed that 60m. both Trewavas and Stauffer have been COMMERCIAL IMPORTANCE: Minor con- correct in the conclusions they have tribution to stern bottom trawl and drawn from the specimens they exam- midwater trawl catches. ined and have thus tentatively con- NOTES: Stauffer & McKaye (1988) re- cluded that Burgess & Axelrod's pho- garded this species as distinct from tograph of a male with bright yellow Alticorpus (= Trematocranus) peterdaviesi breeding colours is actually of the para- (Burgess & Axelrod) and gave no indi- type of A. peterdaviesi (and thus is A. cation that there might be any confu- pectinatum), and that the other speci- sion between them, but did not pro- men illustrated on p. 208 of Konings vide a diagnosis of the species. Eccles (1990a) is the holotype of A. peter- & Trewavas' discussion clearly implies daviesi. It remains possible that the two that they suspected that A. pectinatum types of A. pectinatum are conspecific. might be a junior synonym of Burgess & Axelrod's species, a view made ex- plicit by Konings (1990a). A. peter- Alticorpus peterdaviesi daviesi was described from 2 speci- (Burgess & Axelrod) mens, one of which is lodged as the holotype at the US National Museum DISTINGUISHING FEATURES: A deep-bod- (USNM) and the other as the paratype ied species with large cephalic pits and at the Natural History Museum in Lon- a prominent mental knob, it can be dis- don (BMNH). Trewavas (pers. comm.) tinguished from most other Alticorpus examined the BMNH specimen and by its smaller mouth and larger concluded that it was conspecific with number of gillrakers (17-18 cerato- A. pectinatum. Stauffer (pers. comm.) branchials). It differs from A. pectin- examined the specimen in the USNM atum in male breeding colour and its and concluded it was distinct from A. more pointed snout. pectinatum. Neither author had exam- COLOUR: Ripe males have a pale back- ined the other specimen at the time of ground colour with black vertical bars, their respective publications. Konings dark head and fins. Dorsal fin lappets (1990a) published photographs of both white. Numerous large egg-spots on freshly collected type specimens of A. the anal fin (see Konings, 1990a). peterdaviesi. The two individuals look MAXIMUM SIZE: 15cm TL (Konings, rather different and may not be con- 1990a). specific. Only one was figured in the DISTRIBUTION AND ABUNDANCE: A rare original description, and this agrees deep water species, tentatively identi- well with the description of A. pectin- fied during the 1992 survey from a sin- atum. If this individual is the holotype, gle trawl catch at 102m depth in the Stauffer & McKaye's species is likely north of the SE Arm and from the types to be invalid. On the other hand, if the which were collected at a similar depth other specimen, not figured in the and location. original description, is the holotype, COMMERCIAL IMPORTANCE: Negligible.

103 NOTES: This species is dubiously dis- MAXIMUM SIZE: 15cm TL (Konings tinct from A. pectinatum see above for 1990a). discussion. DISTRIBUTION AND ABUNDANCE: A rare deep water species, described from Alticorpus profundicola specimens collected at 159 m off Stauffer & McKaye Nkhotakota. Six specimens taken by the 1992 survey key out as this species, DISTINGUISHING FEATURES: A deep-bod- all from 90-100m depth in the north of ied species with large cephalic pits and the SE Arm. a large eye. It is distinguished from COMMERCIAL IMPORTANCE: Negligible. other Alticorpus species by the number NOTES: Eccles & Trewavas state that of ceratobranchial gillrakers (14-16) it was recorded under the name which is more than have A. mentale, A. 'Trematocranus brunneus' from depth 'deep', and A. 'geoffreyi' and fewer of 55-125m in the SE and SW Arms and than A. peterdaviesi and A. pectinatum. as far north as Senga Bay. At the time I It is distinguished from similarly- examined the collection, specimens shaped Aulonocara species by its higher deposited in the Monkey Bay Fisher- gillraker count and more prominent ies Station under that name appeared

Figure 26. Alticorpus profundicola. Paratype from Nkhotakota. mental process, which is, however, to represent several species. They were much less strongly developed than in collected during the time of the trawl other Alticorpus spp. surveys reported by Eccles & Trewa- COLOUR: Live coloration not known. vas. Preserved females and immature males pale, darker dorsally with traces of faint vertical bars.

104 Alticorpus macrocleithrum bluish silver with 7 dark vertical bars (Stauffer & McKaye) under the dorsal fin and one on the colour photo on page 61 caudal peduncle. The head is dark blue. Fins dark. Dorsal fin with white DISTINGUISHING FEATURES: A deep-bod- lappets, tipped with orange. Numer- ied, laterally compressed species, eas- ous irregular pale egg-spots on anal ily recognised by the projecting fin. Konings (1990a, 1995b) and Reitz cleithrum, visible as a prominent bulge (1992) illustrate ripe males with a simi- under the head. lar colour, but one shown by Konings COLOUR: Generally brownish, lighter (1989) is uniformly bright metallic ventrally, with traces of vertical bars. blue. Unpaired and pelvic fins brownish. MAXIMUM SIZE: 20cm TL (Konings Pectoral fins yellowish. Ripe males 1990a). with 6-7 prominent brown vertical REPRODUCTION: Males are colonial bars, darker fins and a pale blue irides- nesters and dig pits. They appear to be cence on the snout and lips. non-seasonal breeders (Konings, MAXIMUM SIZE: 18cm TL. 1995b). In the aquarium, it spawns in DISTRIBUTION AND ABUNDANCE: Eccles & a small depression and produces 50- Trewavas state that it was commonly 150 fry (Reitz, 1992). trawled from depths of 35-125m in the DISTRIBUTION AND ABUNDANCE: The south of the lake. In the 1992 survey, it lectotype is from Vua at the northern was common in most samples from 90 end of the lake. Eccles (in Eccles & to 128m (the maximum depth fished) Trewavas) provided no ecological in- and on occasion comprised up to 22% formation on this species. Konings of the sample weight. A single small (1995b) reported that the species has a specimen was taken at 60m off Nkope lake-wide distribution, and observed and several adults collected at 68m off it on the northern rocky shore between the Maleri Islands, but no specimens Mdoka and Kande Island, in the SE were seen in any haul from less than Arm (West Reef) and on the Tanzanian 60m depth. coast (Hongi Island). He recorded it is COMMERCIAL IMPORTANCE: In was not found at 15-30m depth over sand. In observed in commercial landings in the 1992 survey it was common, espe- 1990-91. cially around 22m depth off Nkope. COMMERCIAL IMPORTANCE: Minor com- ponent of shallow water trawl catches. Aulonocara rostratum Trewavas NOTES: Konings (1990a, 1995b) con- colour photo on page 62 siders A. macrochir to be conspecific with A. rostratum, a possibility previ- DISTINGUISHING FEATURES: A large ously raised by Eccles (in Eccles & deep-bodied species with large ce- Trewavas 1989). I did not find it easy phalic pores. It has a longer snout than to distinguish this species from the fol- any similar species. lowing one. COLOUR: Generally sandy with a pinkish cast. Ripe males are generally

105 Aulonocara cf macrochir Aulonocara guentheri Eccles colour photo on page 62 colour photo on page 62

DISTINGUISHING FEATURES: A large DISTINGUISHING FEATURES: A relatively deep-bodied species with enlarged ce- deep-bodied species with enlarged ce- phalic pores and no mental process. It phalic pits and no mental process. It is has a larger eye than any other large characterised by its small terminal Aulonocara. It can be distinguished mouth, small eye (28-32% head length) from A. rostratum by its shorter snout, and bright yellow ventral colour. A. larger eye, greyish rather than pinkish 'copper', which has a similar overall cast, and the darker colour of the rip- body shape, is a deep-water species ening males, which also have fewer with prominent broad vertical bars. egg-spots on the anal fin. COLOUR: Generally silver grey, darker COLOUR: Generally silver-grey, with dorsally, with nine thin dark bars un- 7 faint vertical bars under the dorsal der dorsal fin and 2-3 on caudal pedun- fin. Pelvic and pectoral fins yellowish. cle. Snout, lips, throat, belly, and pel- Ripe males have darker vertical bars vic and anal fins bright orange-yellow. and fins. The snout and lips are dark Pectoral fins yellowish. Caudal and blue, sometimes metallic green in life. dorsal fins dusky, with dark spotting. Dorsal fin without white lappets. 4-5 Dorsal lappets dark. Sexually inactive large pale egg-spots on anal fin. males with numerous (up to 13) orange MAXIMUM SIZE: 18cm TL. egg-spots on anal fin. A male, probably DISTRIBUTION AND ABUNDANCE: In the not fully ripe, from Kambiri Point is 1992 survey, I recorded this species illustrated by Konings (1995b). It is from depths of 18 to 75m in the SE Arm. generally orange-brown, particularly It was quite numerous, particularly at on the nape, with bright orange egg- the Ulande East trawl stations, where spots, small orange spots in the dorsal it comprised 6-19% of the catch weight fin and with a broad white margin on at 50-52m. It was much less common the spinous dorsal fin, with a thin or- south of Boadzulu Island. ange margin posteriorly, which contin- COMMERCIAL IMPORTANCE: This species ues onto the soft-rayed part. occurred in around half of the mid- MAXIMUM SIZE: 14cm TL. water trawl samples and one-third of DIET: Stomach contents of a single the deep-water stern trawl catches specimen examined, indicating a diet which I sampled, but comprised less of benthic invertebrates, mostly chi- than 1% of the total annual catch ronomids and ephemeroptera (Grant weight. It also occurred in pair trawl & Turner, unpubl.). catches. DISTRIBUTION AND ABUNDANCE: A shal- NOTES: This species does not appear low water species, rarely taken in trawl to be conspecific with A. macrochir samples, but I have observed it, either Trewavas, which is probably a junior while diving or through examination synonym of A. rostratum. of artisanal catches, at Monkey Bay, Chirombo Bay, Namiasi, and Lake Malombe, all sheltered muddy areas.

106 Konings (1995b) has recorded it from larger eyes and wider heads. These are Monkey Bay, Kadango, Makanjila mostly found in shallower water (50- Point, and Senga Bay. 80m) at Nkope to Ilala Gap and may COMMERCIAL IMPORTANCE: A small represent a distinct species. component of seine net catches in Lake Malombe and SE Arm. Aulonocara 'gold' NOTES: This species was described in colour photo on page 62 1989 from two of the three syntypes Regan. These speci- DISTINGUISHING FEATURES: A deep-bod- mens were collected in 1920 from un- ied species characterised by vertical known localities. My identification is bars, a short snout, and a large eye. It largely based on the live colour re- has a longer snout and deeper cheek ported by Konings (1995b) who has than A. 'deep'. examined the types and made further COLOUR: Generally sandy-coloured, collections. darker dorsally, with six faint brown- ish bars under dorsal fin. Ripe males Aulonocara 'deep' are bright golden yellow with brown- colour photo on page 63 ish vertical bars. Snout and cheek bright metallic pale blue. Caudal fin DISTINGUISHING FEATURES: A deep-bod- dark, especially on upper and lower ied species with 6 dark vertical bars borders. Pelvic fins black anteriorly, under the dorsal fin. It has a short yellowish posteriorly. Anal fin dark on snout and a large eye. anterior ventral margin, with several COLOUR: Generally silver grey, darker pale blue white spots and streaks. Dor- dorsally, with seven broad faint brown- sal fin with dark lappets and a pale ish bars under dorsal fin and 2-3 on bluish submarginal band, breaking caudal peduncle. Ripe males with dark into several rows of spots posteriorly. dorsal, anal, caudal, and pelvic fins, MAXIMUM SIZE: 15cm TL. dark head and dark vertical bars. 4 DISTRIBUTION AND ABUNDANCE: A rare large pale yellowish egg-spots on anal species, recorded from 40-90m depth fin. in the northern part of the SE Arm. MAXIMUM SIZE: 14cm TL. COMMERCIAL IMPORTANCE: Minor. DISTRIBUTION AND ABUNDANCE: A deep NOTES: In the field, this species was water species, which is common at not reliably distinguished from A. depths of 90-120m. 'deep' until a male in breeding colours COMMERCIAL IMPORTANCE: A small had been collected. component of deep water trawl catches. Aulonocara 'copper' NOTES: Some male individuals have colour photo on page 63 orange heads. Not reliably distin- guished from other similar species DISTINGUISHING FEATURES: A deep- during field work. There are similar bodied, prominently-striped species, specimens with 7 vertical bars under distinguished from other deep-bodied the dorsal fin, some of which have Aulonocara spp. by its very laterally

107 compressed body and small terminal COMMERCIAL IMPORTANCE: Minor. mouth. NOTES: At 46m depth off the Maleri COLOUR: Generally brownish, with a Islands, I collected a ripe male (photo coppery cast. Head and unpaired fins 8.8) which was a similar colour to those dark brown, with 7-9 dark brown ver- of A. 'pyramid', but had a slightly tical bars under dorsal fin, and a fur- deeper body and a less flat lower pro- ther 1-2 on caudal peduncle. Pectoral file. This may represent another spe- fins brownish, pelvic fins brown with cies. black anterior margin. Ripe males darker, generally more greyish. Metal- lic blue iridescence on cheek. Dorsal fin Aulonocara 'blue-orange' lappets, gular membrane and pelvic colour photo on page 63 fins black. 4 large pale yellowish egg- spots. DISTINGUISHING FEATURES: A relatively MAXIMUM SIZE: 15cm TL. small, fairly deep-bodied species, DISTRIBUTION AND ABUNDANCE: A deep which can be distinguished from A. water species, frequently encountered 'orange' only by male breeding colours. in small numbers at depths of 60-120m COLOUR: Generally silvery, brownish in the SE Arm and Domira Bay. dorsally with traces of 7-8 vertical bars. COMMERCIAL IMPORTANCE: A minor Ripe males generally darker with component of deep water trawl darker vertical bars under dorsal fin catches. and 1-2 on caudal peduncle. Fins dark, spotted. Dorsal fin lappets white, with thin black proximal edge anteriorly. Aulonocara 'pyramid' Pelvics dark, black anteriorly, with a colour photo on page 63 white anterior edge to the elongate fila- ment of the first ray. The dark orange DISTINGUISHING FEATURES: An elongate colour of the head extends posteriorly species distinguished by its very large including the nape and chest. Snout elliptical eye and flat lower profile. and lips with metallic blue iridescence. COLOUR: Generally silvery, sandy MAXIMUM SIZE: 11cm TL. dorsally with traces of vertical bars. DIET: Chironomid larvae. Some algae Ripe males generally darker with 7 and detritus also ingested (1 individual brownish bars under dorsal fin and 2- examined by Grant & Turner, unpub- 3 on caudal peduncle. Fins yellowish, lished). pelvics with black anterior margin. DISTRIBUTION AND ABUNDANCE: Often Lips, snout and cheeks metallic blue- numerous in shallow waters of less green. than 30m, north of Boadzulu Island in MAXIMUM SIZE: 12cm TL. the SE Arm. Occasionally the dominant DISTRIBUTION AND ABUNDANCE: Rare, species in catches in this area, as in the occasionally encountered at 30 to 72m 1992 survey where it comprised 27% depth between Boadzulu Island and of the catch weight at 28m at Chiponda Monkey Bay in the SE Arm of Lake and 9.6% at Fowo at a depth of 18m. Malawi. Not recorded north of Namalaka or

108 south of Boadzulu Is. been made of all small Aulonocara from COMMERCIAL IMPORTANCE: In 1990-91 it Lake Malawi. was of little commercial importance, but it might be significant if pair trawls were to fish this area more frequently. Aulonocara 'orange' NOTES: Very much smaller, more colour photo on page 63 lightly-built individuals with the same male colour have been observed in DISTINGUISHING FEATURES: A small spe- beach seine catches south of Boadzulu cies mainly recognised by its male Island. These have not been collected breeding colours and the female's or- and may prove to be distinct. Eccles (in ange fins. Eccles & Trewavas, 1989) described a COLOUR: Generally silvery, sandy similar colour pattern for a species dorsally with traces of 7 vertical bars. trawled in shallow waters south of Pelvic and anal fins pale orange. Ripe Boadzulu Island and suggested that males with purplish cast and darker this may be Aulonocara nyassae Regan. vertical bars. Fins very dark, with black Konings (1995b) has collected males tips to dorsal lappets. Head and nape with an identical colour from around dark orange. Mazinzi Reef, well to the north of MAXIMUM SIZE: 9.5cm TL. Boadzulu Island. He has based a rede- DIET: 2 specimens examined con- scription of A. nyassae on these speci- tained mostly copepods, along with mens. Konings' illustrations and Ec- some algae and detritus (Grant & cles' description differ from A. 'blue- Turner, unpubl.). orange': (i) the dorsal fin lappets are DISTRIBUTION AND ABUNDANCE: Abun- dark; (ii) the orange colour on the nape dant. In the 1992 trawl survey, it was does not extend to the belly or onto the the dominant species at 28-34m at opercula. Mpemba (16-33% catch weight) and Although Eccles' and Konings' re- was also abundant at 26-34m at descriptions both state the A. nyassae Michesi (13-21%). Not positively iden- has 8-9 vertical bars under the dorsal tified from north of Monkey Bay nor fin, Eccles' records that live males have deeper than 40m. Records from deeper 8 bars — the same number as the speci- water, made from preserved material, mens illustrated by Konings. The holo- may represent different species. type of A. nyassae has 9 bars. Konings COMMERCIAL IMPORTANCE: An impor- (1995b) illustrates live specimens of tant component of pair trawl catches Aulonocara gertrudae Konings which (8%), and also taken in bottom trawl have 9 bars under the dorsal fin. Given and midwater trawl catches (1-1.5%). the large number of morphologically similar small Aulonocara species, and the unknown collecting locality of the Aulonocara 'yellow' type, these re-descriptions of A. nyassae colour photo on page 64 seem premature and the name is prob- ably best applied only to the type speci- DISTINGUISHING FEATURES: A small spe- men until detailed examinations have cies, mainly recognised by its male

109 breeding colours. It has a more slen- only from the Maleri Islands. der head and body than A. 'orange', A. COMMERCIAL IMPORTANCE: Not known, 'blue-orange', and the A. 'dark stripe' as commercial fisheries from the SW group, and differs from the similarly- Arm and Domira Bay have yet to be shaped A. 'green' in the mixture of bi- studied. cuspid and unicuspid jaw teeth. COLOUR: Females and immature males silvery, sandy dorsally with Aulonocara 'green' traces of 7 vertical bars under dorsal colour photo on page 64 fin. Pelvic and anal fins hyaline. Ripe males silvery-yellow, with darker ver- DISTINGUISHING FEATURES: A small slen- tical bars. Fins dusky, anal fin with der species distinguished from the dark margin. Caudal darker dorsally similar A. 'yellow' by male breeding and ventrally, spotted in centre. Pelvic colour and the jaw teeth, which are all fin black anteriorly. Dorsal fin lappets unicuspid. white with broad black submarginal COLOUR: Dorsal fin dark orange- band. Row of dark brownish spots be- brown, with bright orange lappets. low submarginal band. Head dark Caudal fin yellowish. Anal fin dark grey dorsally, lower part of head pale with numerous large bright yellow lemon yellow to orange. egg-spots. Dark stripe through eye ex- MAXIMUM SIZE: 8cm TL tending to preorbital bone. Cheeks and DISTRIBUTION AND ABUNDANCE: Com- snout with metallic green tint. mon, occasionally dominant at depths MAXIMUM SIZE: 9cm TL. of 40-95m, particularly at 44m at DISTRIBUTION AND ABUNDANCE: Rare. Mpemba (7-16% catch weight). Positively identified in the SE Arm COMMERCIAL IMPORTANCE: A significant from a single trawl sample from 35- component of deep water bottom trawl 40m, to the NW of Boadzulu Island. catches (1.5%), also taken in midwater COMMERCIAL IMPORTANCE: Minor. trawl catches. NOTES: Preserved material and fe- males not reliably distinguished from related A. 'yellow' and the A. 'dark Aulonocara 'deep yellow' stripe' complex. colour photo on page 64

DISTINGUISHING FEATURES: A small Aulonocara 'dark stripe' Aulonocara, with a higher gillraker colour photo on page 64 count than any other known Aulono- cara species (12-13 ceratobranchials). DISTINGUISHING FEATURES: A small spe- COLOUR: Ripe males silvery with pur- cies with a large eye, ventral mouth plish cast and 7 dark vertical bars un- and flattened lower profile, it is distin- der the dorsal fins. Fins dusky. Snout guished from A. 'orange' by male and cheek bright yellow. breeding colour and shallower head MAXIMUM SIZE: 9cm TL. depth. DISTRIBUTION AND ABUNDANCE: Known COLOUR: Males (ripe?) silvery-white,

110 darker dorsally with 7 vertical bars cal bars, dark fins and a green irides- under dorsal fin and a prominent pre- cence on the snout. These may be a dif- orbital stripe. ferent species. MAXIMUM SIZE: 8cm TL. DISTRIBUTION AND ABUNDANCE: Rare. Positively identified from a single Aulonocara 'stonemani' trawl sample from 35-40m, to the NW colour photo on page 64 of Boadzulu Island. COMMERCIAL IMPORTANCE: Unknown. DISTINGUISHING FEATURES: A small slen- NOTES: Preserved material not reli- der species, distinguished from simi- ably distinguished from related spe- lar species by its ventrally positioned cies. Similar specimens from 72m at mouth, flat lower profile and by male Chekopa have 6 vertical bars under the breeding colours. dorsal fin and differ in multivariate COLOUR: Ripe males silvery, with 5-6 shape. Other 7-striped specimens from dark vertical bars under the dorsal fin. 54m at Nkope also differ in shape. The dorsal fin with large yellowish These may represent further species, spots, and white lappets, tipped with but these are known only from pre- black. The anterior spine is black. The served material and so male coloration caudal fin is yellowish, the pelvic fins is unknown. black, and the anal fin has a broad black margin and whitish-yellow egg- spots. There is a black stripe through Aulonocara 'minutus' the eye, which extends onto the colour photo on page 64 preorbital bone. MAXIMUM SIZE: 5-7cm TL. DISTINGUISHING FEATURES: A small slen- DISTRIBUTION AND ABUNDANCE: Rare. der species, with a small terminal Positively identified from a single mouth and long filaments on the soft trawl sample from 90m, off Monkey parts of the dorsal and anal fins. Bay. COLOUR: Females and immature COMMERCIAL IMPORTANCE: Negligible. males silvery, darker dorsally with NOTES: This species may be conspe- traces of 6 vertical bars under dorsal cific with Haplochromis stonemani Bur- fin. Ripe males silvery, with darker gess & Axelrod which was described vertical bars. Fins darker. Head orange. from a single specimen from 43 fath- MAXIMUM SIZE: 7cm TL. oms (79m) depth off Monkey Bay. Bur- DISTRIBUTION AND ABUNDANCE: Appar- gess & Axelrod provide a photograph ently confined to deep water. Positively of the type, which has a very similar identified from a single trawl sample morphology to this Aulonocara. The from 90m, off Monkey Bay. colour is similar but more silvery, sug- COMMERCIAL IMPORTANCE: Negligible. gesting that it might be a male which Occasionally seen in demersal stern is not fully ripe. I have not examined trawl catches. the type, but Oliver (1984) mentions NOTES: I have observed similarly- that it has expanded cephalic pits, shaped males with broad dark verti- which implies that it should be as-

111 Figure 27. Aulonocara 'long', male, South-East Arm. signed to the genus Aulonocara. How- pores take the form of small, deep pits. ever, Eccles & Trewavas made no men- Colours: Live colours are not known. tion of expanded sensory pits in their Females and non-breeding males are redescription, in which they assigned pale, with 7 dark vertical bars under the species to Placidochromis. However, the dorsal fin. they did not actually re-examine the MAXIMUM SIZE: It attains at least 10cm type, although Eccles had seen it in TL. 1973. Their redescription is based on DISTRIBUTION AND ABUNDANCE: Uncom- other material, which may not be con- mon. Known only from 90-126m depth specific. in the north of the SE Arm.

Aulonocara 'long'

DISTINGUISHING FEATURES: A medium- sized elongate species. It has a small terminal mouth and resembles a slen- der version of A. 'copper'. The cephalic

112 Chapter 11 Placidochromis hennydaviesae complex

This group is one of the least known ange-brown spots in dorsal and cau- among Malawi cichlids. A single spe- dal fins. Dorsal has a broad white sub- cies was described by Burgess & Axel- marginal band with red lappets. rod on the basis of one specimen. I MAXIMUM SIZE: 8.5cm TL. presently assign seven species to this DIET: Copepods and chironomids group, but P. 'platyrhynchos' also bears (Eccles & Trewavas, 1989). some resemblance to some of the spe- REPRODUCTION: Ripe males collected cies assigned to Sciaenochromis. All spe- in April and May, at depths of 84-128m. cies are characterised by a melanin DISTRIBUTION AND ABUNDANCE: Rare. pattern of vertical bars, a large eye, Known from the holotype, 8 specimens pointed snout and a lower jaw which collected by Eccles (see Eccles & Trewa- is flat in cross section. They are mainly vas) and 7 specimens from the present deep water species, although one spe- survey. Material examined by Eccles & cies (III) is recorded from water as shal- Trewavas was collected from the north low as 45m. All are small. of the SE and SW Arms (Eccles & Trewavas 1989) at 43 to 70 fathoms (80- 130m). I collected specimens from the Placidochromis hennydaviesae SE Arm at 84-128m depth. The trawl- (Burgess & Axelrod) ing gear did not permit fishing in colour photo on page 161 deeper water. COMMERCIAL IMPORTANCE: Minor. DISTINGUISHING FEATURES: A relatively NOTES: I have identified this species deep-bodied species with a smaller eye from the redescription provided by than others in this group. Distin- Eccles & Trewavas. It should be stres- guished from similar species by the sed that these authors did not exam- low number of gill rakers (9-12), which ine the holotype while working on are deeply forked, and the presence of their redescription, although Eccles six vertical bars under the dorsal fin. saw the type at the time of its collec- COLOUR: Silver grey, with six dark tion in 1973. vertical bars under the dorsal fin, and generally a further two on the caudal peduncle. Ripe males with black chin, Placidochromis 'hennydaviesae II' belly, anterior of pelvic fins, and anal fin margin. Upper and lower borders DISTINGUISHING FEATURES: A relatively of caudal fin also black. Six or more deep-bodied barred species. Distin- orange egg-spots on anal fin and or- guished from similar species by the

113 Figure 28. Placidochromis hennydaviesae, male, South-East Arm.

Figure 29. Placidochromis 'hennydaviesae II', male, South-East Arm.

Figure 30. Placidochromis 'hennydaviesae III', male, South-East Arm.

114 Figure 31. Placidochromis 'hennydaviesae IV', male, South-East Arm.

Figure 32. Placidochromis 'hennydaviesae V', female, South-East Arm.

Figure 33. Placidochromis 'hennydaviesae VI', male, South-East Arm.

115 high number of long pointed gill rakers Placidochromis 'hennydaviesae III' (25 or more), and the presence of 5 ver- colour photo on page 161 tical bars under the dorsal fin (6-7 for other species). DISTINGUISHING FEATURES: A relatively COLOUR: Silver grey, with five dark deep-bodied species with a small eye. vertical bars under the dorsal fin, and Distinguished from species I, IV and generally a further two on the caudal VI by the higher number of gill rakers peduncle. (20-23), and from species II by the MAXIMUM SIZE: 9cm TL. lower number. The mouth is larger and REPRODUCTION: Males in breeding the lower jaw more flattened than in dress recorded from January to May species VI, and the mouth and eye are (no other samples known) from depths smaller than in species V and II. of 90-100m. COLOUR: Silver grey, with 6-7 dark DISTRIBUTION AND ABUNDANCE: Uncom- vertical bars under the dorsal fin, and mon. Confined to deep water (85- generally a further two on the caudal 130m). Known only from the north of peduncle. the SE Arm. MAXIMUM SIZE: 9cm TL. COMMERCIAL IMPORTANCE: Minor. REPRODUCTION: Males in breeding NOTES: At around 90m depth off dress caught in February, May, Septem- Monkey Bay there appears to be a hy- ber and December, suggesting that this brid zone between this and sp. III. In- species is a non-seasonal breeder. dividuals with 6 bars and 26-29 gill- These were collected from depths of 46- rakers are found together with 5 barred 86m, and throughout the recorded fish with 25-28 gillrakers. Alternatively range, suggesting that there is no mi- it could be that the two species coexist gration to particular breeding grounds. in this area, and perhaps are variable DISTRIBUTION AND ABUNDANCE: Fre- in gillraker counts. Further studies are quently caught in trawls, but never in necessary to clarify the status of this large numbers. Recorded from 45-90m population. from Nkope in the SE Arm to the Maleri Islands. COMMERCIAL IMPORTANCE: Occasion- ally taken in bottom trawls, but in small numbers. NOTES: See under previous species. Water depth Number of bars 44-68m 80-92m 100-120m Placidochromis 'hennydaviesae IV' 4 01 0 5 4 17 54 656128 DISTINGUISHING FEATURES: Some- 7911 what more elongate than other simi- lar species, but deeper bodied than P. 'platyrhynchos'. Distinguished Frequency distributions of vertical bars in P. 'hennydaviesae II' & 'III' from most other similar species of

116 this group by the presence of 7, rather Placidochromis 'hennydaviesae VI' than 5-6, dark vertical bars under the dorsal fin. Differs from most similar DISTINGUISHING FEATURES: A relatively species in the lower number of gillrak- deep-bodied species with a large eye. ers (10-12), and from P. hennydaviesae Distinguished from species I, and IV in the form of the gillrakers which are by the higher number of gill rakers (15- short and not forked. 16), and from species II by the lower COLOUR: Silver grey, with seven dark number. The mouth is smaller and the vertical bars under the dorsal fin, and lower jaw less flattened than in species generally a further two on the caudal IV and V. peduncle. Leading edge of pelvic fins COLOUR: Silver grey, with 6-7 dark black. vertical bars under the dorsal fin, and MAXIMUM SIZE: 11cm TL. generally a further two on the caudal REPRODUCTION: Sexually active males peduncle. taken in May. MAXIMUM SIZE: 9cm TL. DISTRIBUTION AND ABUNDANCE: Fre- REPRODUCTION: Males with some quently present in samples from wa- breeding colours taken in July and ter deeper than 90m. Known only from August. the north of the SE Arm. DISTRIBUTION AND ABUNDANCE: Known COMMERCIAL IMPORTANCE: None. from 35 specimens taken between 92 and 126m in the north of the SE Arm. COMMERCIAL IMPORTANCE: None. Placidochromis 'hennydaviesae V'

DISTINGUISHING FEATURES: A relatively deep-bodied species with a very large Placidochromis 'platyrhynchos' eye and mouth. Distinguished from all other species, except P. hennydaviesae DISTINGUISHING FEATURES: Distin- (sp. I) by the deeply forked gillrakers. guished from other species of this Differs from sp. I in the larger mouth group by the much more elongate and eye and larger number of gill- body. 10-11 lower gill rakers. Distin- rakers (15-19). guished from Sciaenochromis species by COLOUR: Silver grey, with 5-6 dark the more pointed snout, larger eye and vertical bars under the dorsal fin, and less steeply-angled gape. generally a further two on the caudal COLOUR: Pale, sandy with six or seven peduncle. brownish vertical bars under the dor- MAXIMUM SIZE: 12.5cm TL. sal fin, and generally a further two on DISTRIBUTION AND ABUNDANCE: Rare. the caudal peduncle. Ripe males with Known only from 4 specimens taken yellow or orange dorsal lappets and a between 118 and 128m in the northern broad white submarginal band. part of the SE Arm. MAXIMUM SIZE: 13cm TL. COMMERCIAL IMPORTANCE: None. DISTRIBUTION AND ABUNDANCE: Uncom- mon. Recorded from 118-126m in north of the SE Arm.

117 Figure 34. Placidochromis 'platyrhynchos', male, South-East Arm.

COMMERCIAL IMPORTANCE: Occasion- ally taken in bottom trawls, but in small numbers.

118 Chapter 12 Miscellaneous barred species

This chapter deals with a mixed bag DISTRIBUTION AND ABUNDANCE: Rare. of species. Most are characterised by Four specimens taken from a haul off dark vertical bars, at least in the fe- Monkey Bay at a depth of 84-94m and males and immatures. In some cases a single individual from 80m at the males are vertically barred, but fe- Domira Bay. males are plain or their colour is un- COMMERCIAL IMPORTANCE: Negligible. known. There is no reason to think that NOTES: Females and immatures could this is a monophyletic group and it is easily have been overlooked in trawl likely that some species have affinities catches. with the Lethrinops species, and per- haps with other differently-marked forms. Placidochromis 'macrognathus' colour photo on page 161

Placidochromis 'carnivore' DISTINGUISHING FEATURES: A deep-bod- colour photo on page 161 ied species with broad vertical bars and a large mouth. Differs from Lethrinops DISTINGUISHING FEATURES: a small fairly polli and P. 'acuticeps' in the larger nondescript deep-water species with 9- mouth, stronger lower jaw and less 10 lower gillrakers and 6 thin vertical pointed snout. bars under the dorsal fin. The mouth COLOUR: Generally silvery, with a is fairly large and the outer teeth small brownish cast, with 5-6 broad brown and unicuspid. Body shape is similar vertical bars under the dorsal fin, and to Otopharynx brooksi, Nimbochromis 1-2 on the caudal peduncle. Ripe males polystigma, and N. livingstonii, but it darker, with dark fins, and several lacks the spotted pattern of these spe- large whitish yellow streaks on anal cies. fin. COLOUR: Ripe male silvery with 6 thin MAXIMUM SIZE: 13cm TL. dark vertical bars under the dorsal fin. DIET: Not known, but morphology Dorsal, caudal and anal fins dark. Dor- suggests predatory habits. sal with white margin, anal with nu- DISTRIBUTION AND ABUNDANCE: Voucher merous white egg-spots. Pelvics black. specimens of confirmed identity from Pectorals brownish. Head dark, with 40-75m throughout the SE Arm. Re- blue iridescence on snout. Specimen corded from depths of 25-100m from from Domira Bay had a yellowish the SE Arm and Domira Bay. Fre- chest. Female and immatures un- quently found, but never in large num- known. bers. MAXIMUM SIZE: 9.5cm TL. COMMERCIAL IMPORTANCE: Minor.

119 Taken by the bottom trawl and mid- key Bay at depth of 70-125m. water trawl fisheries. COMMERCIAL IMPORTANCE: Minor. NOTES: Specimens labelled 'Haplo- NOTES: Specimens were found in the chromis macrognathus' were found at Monkey Bay field collection under the Monkey Bay field collection, but had name 'Haplochromis acuticeps', but not been described. these had not been described.

Placidochromis 'acuticeps' Sciaenochromis 'deep water' colour photo on page 161 colour photo on page 161

DISTINGUISHING FEATURES: A relatively DISTINGUISHING FEATURES: A slender elongate species with indistinct broad barred species with a large mouth and vertical bars, a flat lower jaw, and a eye. It has fewer vertical bars than sharply pointed snout. Differs from other Sciaenochromis species. It has a Lethrinops polli in larger mouth and more prominent premaxillary pedicel from P. 'macrognathus' in the smaller than Sc. psammophilus and the poste- mouth, weaker lower jaw and more rior angle of the lower jaw is more pro- pointed snout. Deeper bodied than truding. The interorbital width is Placidochromis 'long' or P. 'platy- broader (16.4-17.2% HL) than in Sc. rhynchos'. psammophilus (13-14.7% HL). Sc. ahli, COLOUR: Generally silvery, with a which I have not observed in the field, pinkish-brown cast, with 6 broad in- has a deeper body (34-36% SL com- distinct dark vertical bars under the pared to 28-30% SL), shorter premax- dorsal fin. Ripe males darker, with illary pedicel (19-20% v 22-24% HL), dark fins, and several large whitish and shorter preorbital bone (16-17% v yellow streaks on anal fin. 18-20% HL), and more vertical bars. It MAXIMUM SIZE: 15cm TL. is more slender than P. 'macrognathus' DISTRIBUTION AND ABUNDANCE: Rare. and P. 'acuticeps', and is more laterally Known only from the SE Arm off Mon- compressed and has a more steeply an-

Figure 35. Sciaenochromis 'deep water', female, South-East Arm.

120 gled gape than P. 'platyrhynchos'. dal fins spotted and dorsal margin COLOUR: Females and immature white with orange lappets. Numerous males sandy with 6 brownish vertical large yellow-orange egg-spots on anal bars under the dorsal fin and two on fin. the caudal peduncle. MAXIMUM SIZE: 14cm TL (Konings MAXIMUM SIZE: 12 cm TL. 1993a). DISTRIBUTION AND ABUNDANCE: Known DIET: Generally swims about 30cm only from deep water (100-128m) in the above the sand, taking small fishes and north of the SE Arm around Domwe probably invertebrates from the bot- and Chinyankwazi Islands. tom (Konings 1993a). A single speci- COMMERCIAL IMPORTANCE: Minor. men I collected contained sand and NOTES: Konings' revision of Sciaeno- remains of chironomid larvae (Grant chromis was published in 1993, after my & Turner unpubl.). field work. I did not distinguish this REPRODUCTION: Konings (1993a) notes species from Sc. psammophilus and re- that territorial males excavate a small corded both as 'Sc. cf. ahli'. I did not nest in the sand, often partly under a find any specimens of Sc. ahli in the small rock. Reproduction at Kande Is- material I collected and found that Sc. land is apparently all year round. 'deep water' and Sc. psammophilus had DISTRIBUTION AND ABUNDANCE: Konings clearly separate depth distributions. Sc. (1993) records this species from around fryeri (Konings 1993), which was for- Kande Island, and has tentatively ob- merly confused with Sc. ahli (e.g. Ec- served it from Likoma, Masinje, Senga cles & Trewavas, 1989), is a dark brown Bay and Mdoka. He records it as un- species confined to rocky shores. common and found on sandy sub- strates at 5-30m depth. It was posi- tively identified from 19-50m depth Sciaenochromis psammophilus and tentatively recorded from 22-60 m Konings in the 1992 survey of the SE Arm, al- colour photo on page 162 though more frequently encountered in shallower waters. It never repre- DISTINGUISHING FEATURES: A slender sented a large proportion of the catch. sandy-coloured species with faint ver- COMMERCIAL IMPORTANCE: Minor. tical bars and a large mouth and eye, NOTES: This species was described and a short snout. The interorbital subsequent to the 1992 survey and was width is narrower (13-14.7% HL) than not reliably identified in the field. It in other Sciaenochromis species (17.5- was confused with the preceding spe- 20.8% HL). cies. COLOUR: Females and immature males sandy with 8 or more sandy col- Sciaenochromis benthicola Konings oured vertical bars under the dorsal colour photo on page 162 fin. Ripe males grey-blue, yellowish on flanks with dark vertical bars, dark DISTINGUISHING FEATURES: A relatively pelvic and anal fins and a dark lower elongate species with numerous verti- edge to the caudal fin. Dorsal and cau- cal bars and three faint spots. Gener-

121 ally similar in shape to M. spilostichus, rence in one sample at 20m. Sometimes from which it differs in colour pattern quite abundant. and in its deeper body. The rounded COMMERCIAL IMPORTANCE: Recorded head, large eye and prominent premax- from 42% of semipelagic trawl catches illary pedicel distinguish this species and 92% of deep-water demersal trawl from most others with vertical bars and catches, but comprised a relatively spots. small proportion of the total biomass COLOUR: Females and immature (0.5 and 1.5% respectively). May also males brownish, paler ventrally with be caught by hook and line. 10 brown vertical bars under the dor- sal fin. Faint blotches on the third and ninth bars and another at the posterior Placidochromis 'green stripe' of the caudal peduncle. Ripe males are blue dorsally with a blue head, blue DISTINGUISHING FEATURES: A very small, flanks and bright yellow belly. Orange elongate, laterally compressed species spots on dorsal and caudal fins and with several vertical bars. upper part of body. Dorsal, anal and COLOUR: Generally sandy with 9 ver-

Figure 36. Placidochromis 'green stripe', male, South-East Arm. pelvic fins with white margins. Dorsal tical bars under the dorsal fin. Ripe fin lappets tipped with orange. males with darker bars, snout metallic MAXIMUM SIZE: 17.5cm TL. blue, iris greenish in centre, darker DISTRIBUTION AND ABUNDANCE: Type outside, operculum yellow, faint dark from Kaporo in the far north of the bar through eye to jaw angle. Con- lake. Konings (1995b) also recorded it spicuous horizontal band of iridescent from Mdoka and Ntekete. In the 1992 green along midline, between vertical survey, it was common at 44-100m bars. Dorsal fin yellowish, with bright depth throughout the SE Arm. There yellow lappets and black submarginal is an unconfirmed record of its occur- band. Caudal and anal fins yellowish,

122 anal dark distally. Pelvics clear with NOTES: This species has never been black anterior margin. previously illustrated or reported, al- MAXIMUM SIZE: 6.5cm TL. though it is fairly distinctive and com- DISTRIBUTION AND ABUNDANCE: Rare, mon in trawl catches in the well-stud- recorded only from 72-114m depth off ied SE Arm. Monkey Bay. COMMERCIAL IMPORTANCE: Minor. Placidochromis 'longimanus NOTES: There appear to be several Malombe' other small elongate deep water spe- cies which are poorly known. DISTINGUISHING FEATURES: A deep-bod- ied species with faint narrow vertical bars. 12-14 Gillrakers. Differs from P. Placidochromis 'long' 'longimanus Namiasi' in male breed- colour photo on page 162 ing colours, and from Lethrinops 'pink head' in male colour and dentition. DISTINGUISHING FEATURES: A small elon- Deeper bodied than N. 'argyrosoma gate species, distinguished from other blue'. Dental arcade is of the 'haplo- similar species by its acutely pointed chromis' form. Generally three tooth snout and broad vertical bars. rows in lower jaw. Lethrinops micrent- COLOUR: Generally pale sandy with odon has a less pointed snout. 6-7 vertical bars under the dorsal fin COLOUR: Females and immature and 1-2 on caudal peduncle. Ripe males generally silvery, sandy dorsally, males pale silvery grey, with darker occasionally with very faint traces of vertical bars. Darker dorsally and with vertical bars. Ripe males bright pow- dark snout and lips. Snout with me- der blue, or turquoise. 7-8 darker ver- tallic blue iridescence. Unpaired fins tical bars under dorsal fin occasionally and pelvics dusky. Pelvics black visible. Dorsal fin with white lappets, anteriorly, and anal distally. Anal fin often with red tips. Caudal fin with with 2 large yellowish egg-spots. Dor- orange spots. Anal fin with yellow or sal darker distally, with white lappets. red margin and 3-7 white egg-spots. Eye silvery, dark dorsally. Pelvic fins with dark anterior margin. MAXIMUM SIZE: 12cm TL. MAXIMUM SIZE: 12.5 cm TL. DISTRIBUTION AND ABUNDANCE: Found DISTRIBUTION AND ABUNDANCE: Known in most trawl samples taken between only from Lake Malombe where ripe 45 and 55m in the northern part of the males are frequently seen during the SE Arm, usually in small numbers, hot dry season. occasionally numerous. Occasionally COMMERCIAL IMPORTANCE: Probably a taken at other depths, the total range significant component of seine catches recorded in the 1992 survey being 22- in Lake Malombe. 92m. NOTES: During my examination of COMMERCIAL IMPORTANCE: Common in catches from Lake Malombe in 1991- all types of commercial trawl catches, 92, I did not reliably distinguish fe- but comprising a fairly small propor- males and immature fish from those of tion of the biomass. the more numerous Lethrinops 'pink

123 Figure 37. Placidochromis 'longimanus Malombe', male, Lake Malombe. head'. It would be surprising if this vas draw attention to the similarity species was really restricted to Lake between this species and Lethrinops Malombe, as this lake was dry during micrentodon. Trewavas (1935) included the 19th century. Although it keys out the latter in the genus Haplochromis, but as Placidochromis longimanus Trewavas, Eccles & Lewis (1977) re-assigned it to it is not certain that they are conspe- Lethrinops on the basis of the form of cific. The type material of P. longimanus the dental arcade. Ctenopharynx inter- is from the SE Arm, but Eccles & Trewa- medius and Ct. nitidus share this dental vas also report it from Lake Malombe arcade form, as do the species they and the SW Arm, stating that it is re- have removed to Tramitichromis and stricted to beds of rootless macro- Taeniolethrinops. Eccles & Trewavas phytes (Ceratophyllum and Najas) at (1989) state that they believe that such depths of 5-15m. However, Eccles & feeding structures may represent con- Trewavas' colour notes for the ripe vergence or parallelism and that male of P. longimanus are identical to shared melanin pattern may be more those I have made for Lethrinops 'pink reliable indicators of phylogeny. On head'. The two are similar in morphol- this basis, P. longimanus should prob- ogy and meristics and occur in the ably also be included in Lethrinops and same habitat in Lake Malombe. As I it seems an odd choice to be the type have collected numerous voucher species of Placidochromis. Inclusion of specimens and colour photographs of P. longimanus in Lethrinops would cause labelled material, I am inclined to be- nomenclatural problems, as there is lieve that these species were confused already a Lethrinops longimanus, which by Eccles & Trewavas and so any eco- would have priority to the name, as it logical information in that work must was described previous to Haplochro- be regarded as suspect. Eccles & Trewa- mis longimanus.

124 Placidochromis 'longimanus of Boadzulu Island (11%). Namiasi' NOTES: The distribution, colour pat- colour photo on page 163 tern and gillraker counts are similar to those attributed to Lethrinops micrent- DISTINGUISHING FEATURES: A deep-bod- odon by Eccles & Lewis (1977). Eccles ied species with faint narrow vertical & Lewis report that L. micrentodon has bars. 15-17 gillrakers. Differs from P. dentition which is sometimes similar 'longimanus Malombe' in male breed- to other Lethrinops species but some- ing colours, and from Lethrinops 'pink times there is a single row of up to 4 head' in male colour and dentition. teeth at the back of the lower jaw, Deeper bodied than N. 'argyrosoma which is more similar to the 'Haplo- blue'. Dental arcade is of the 'Haplo- chromis-style' and would not be eas- chromis'-form. Generally two tooth ily distinguished from P. 'longimanus rows in lower jaw, but a third row in Namiasi'. The specimen figured by Ec- the anterior part of the jaw. Lethrinops cles & Lewis as L. micrentodon (which micrentodon has a less pointed snout. was not included in the material on COLOUR: Females and immature which they based their redescription) males generally silvery, sandy dorsally, has a more clearly decurved snout and occasionally with very faint traces of much deeper body than the specimen vertical bars. Ripe males bright royal figured by Eccles & Trewavas. Both of blue. 7-9 darker vertical bars under these have a much less pointed snout dorsal fin usually visible. Dorsal fin than P. 'longimanus Namiasi'. Eccles & with white lappets. Dorsal and caudal Lewis state that the pectoral fins of L. fins with orange spots. Anal fin dark micrentodon extend past the insertion with several large pale egg-spots. Pel- of the first anal spine, often as far as vic fins with dark anterior margin. the third anal ray. Those of P. MAXIMUM SIZE: 12cm TL. 'longimanus Namiasi' just reach the DIET: Mainly sedimented filamentous first anal spine. Eccles & Lewis state diatoms, ingested with large amounts that the dorsal fin of L. micrentodon of sand and detritus. Some copepods reaches about midway along the cau- also taken. One individual contained dal peduncle, while that of P. an adult beetle about 2mm long. Two 'longimanus Namiasi' extends much individuals 8-8.5cm SL examined further, in the case of breeding males, (Grant & Turner, unpubl.). well past the end of the caudal pedun- DISTRIBUTION AND ABUNDANCE: Posi- cle. Eccles & Lewis did not, however, tively identified specimens are all from measure mature males, although their the SE Arm, to the South of Boadzulu figure is of a 15.5cm male individual, Island at depths of 5-30m. In 1992 trawl larger than their quoted maximum size surveys, this and similar species, com- for the species. I have only made a pre- prised 13% of the biomass of the fish liminary examination of the types of taken south of Boadzulu Is. but it was L. micrentodon and the material col- not recorded elsewhere. lected by Eccles & Lewis, but regard COMMERCIAL IMPORTANCE: A major their redescription with caution. component of pair trawl catches south

125 There are a number of similar forms P. subocularis has a greater number of some of which may deserve specific thinner vertical bars. status. COLOUR: Females and immature males pale straw-gold with 4-5 broad Placidochromis 'longimanus Maleri' dark vertical bars under the dorsal fin colour photo on page 163 and 2 on the caudal peduncle. Vertical bars each with round dark superim- Similar to P. 'longimanusNamiasi', posed spot following the line of an but the ripe males have a yellow dor- oblique bar, as in Mylochromis spp. sal fin margin. Known from one sam- Usually an oblique dark bar through ple at 25m off Maleri Island. the eye. Red dorsal fin lappets. Ripe males with dark blue snout, cheeks, Placidochromis 'longimanus and operculum. There is a patch of red- Chirombo' edged scales behind the operculum. colour photo on page 163 Dark red spots on dorsal and caudal fins. Dorsal fin lappets red-orange. Similar to P. 'longimanus Namiasi', Anterior of pelvic fins and most of anal but with only 13-14 ceratobranchial fin black. gillrakers on the first arch. Known from MAXIMUM SIZE: 16cm TL (Konings, depths of 20-50m south of Boadzulu 1990a). Island and also off Chirombo Bay. DIET: Konings (1990a) states that it feeds on benthic invertebrates includ- Placidochromis 'longimanus ing molluscs. The stomachs of 4 speci- slender' mens of 8.5-12cm SL (Grant & Turner, unpubl.) contained mainly large A slender rather drab species inter- benthic invertebrates: ephemeropteran mediate in form between P. 'longi- nymphs, molluscs, chironomid larvae. manus Namiasi' and the sympatric Also some cladocera, copepods and Nyassachromis 'argyrosoma blue'. It has algal material. 10-12 lower gillrakers and is known REPRODUCTION: A colonial sand-castle from 19-35m depth between Namiasi nester (Konings, 1995b). and Malindi and also off Chirombo DISTRIBUTION AND ABUNDANCE: Eccles & Bay. Trewavas examined material assigned to P. subocularis from the SE and SW Arms and unknown localities, and re- Placidochromis cf. subocularis port it from many trawl hauls at 15- colour photo on page 163 18m depth. Konings (1990a) reports that it is only found in the southern DISTINGUISHING FEATURES: A species part of Lake Malawi, as well as in Lake with prominent broad stripes. Distin- Malombe. In the 1992 trawl survey, it guished from P. johnstoni by its smaller was found in small numbers in most mouth, and from Mylochromis spp. by trawl samples taken at depths of 18 to the vertical bars being more prominent 40m in the SE Arm. Occasionally nu- than the oblique bar. The holotype of merous. Probably common in shal-

126 lower water. I occasionally observed requires further investigation. Konings small specimens in artisanal catches (1995b) considers that Placidochromis from in Lake Malombe. subocularis may belong in Mylochromis COMMERCIAL IMPORTANCE: Taken in and it certainly bears some resem- small quantities in pair and midwater blance to M. ericotaenia and M. trawl catches. Probably also a signifi- labidodon. cant component of seine and gill net catches. NOTES: During my field work in Ma- Placidochromis johnstoni (Günther) lawi from 1990-92, I regarded this spe- colour photo on page 161 cies as P. subocularis, following the specimens in the Monkey Bay field DISTINGUISHING FEATURES: A species museum, collected and labelled dur- with prominent broad bars. Distin- ing the 1970s, when DH Eccles was guished from P. subocularis by its larger working there. Konings (1995b), how- mouth, lack of oblique row of spots ever, illustrates as P. subocularis, speci- and broader more tapering stripes. mens with thin faint vertical bars and COLOUR: Pale straw-gold with 5 rather small jaws. All come from the broad dark vertical bars under the dor- north (Chesese in Malawi, Lundu in sal fin and 1 on the caudal peduncle. Tanzania, and Londo in Mozambique). Pelvic, pectoral and anal fins golden. He points out that the strain well-es- Dorsal and caudal fins with numerous tablished in the aquarium trade has dark reddish brown spots. According strong vertical bars, large molariform to Eccles & Trewavas (1989), ripe males pharyngeal teeth and comes from Lake have a dark blue head, bars partly ob- Malombe. His illustration of this strain, scured. Fins with orange-brown spots. which resembles the specimens I have Dorsal fin with a white submarginal collected from the south of Lake Ma- band and orange lappets. Anal fin lawi, has only 4 vertical bars under the dark. dorsal fin, while the specimen from MAXIMUM SIZE: 17cm TL (Konings Londo has 9 bars. It seems likely that 1990a). these represent two species. Eccles & DIET: Insects and small fish (Eccles & Trewavas's illustration of the type Trewavas, 1989). When snorkelling, shows 6 bars. The holotype is from an this species is often seen hunting in unknown location. According to small packs, moving rapidly just above Konings the type does not have promi- the bottom, often in company with nent vertical bars, and these are not Nimbochromis polystigma. mentioned by Günther, Regan, or DISTRIBUTION AND ABUNDANCE: Con- Trewavas. Konings (1990a) also illus- fined to shallow waters. Not taken in trates an intermediate-looking form trawl catches in the 1992 survey, al- from Eccles Reef in the SE Arm, which though Eccles & Trewavas record it he (Konings pers. comm.) considers to from two hauls at around 18m depth. be most similar to the holotype. This Ribbink et al. (1983) recorded this spe- has five faint thin vertical bars under cies from most of their study sites. the dorsal fin. The status of these forms Jackson (1961) states that the species is

127 strongly associated with Vallisneria is morphologically almost identical, beds. Konings (1990a) records a lake- but is easily distinguished by its ob- wide distribution and material exam- lique-striped pattern. ined by Eccles & Trewavas came from COLOUR: Females and immature the SE Arm, SW Arm, Karonga, and males silvery-grey with four or five Vua. According to Eccles & Trewavas broad grey vertical bars under the dor- (1989) and Konings (1990a) this species sal fin. Lowe (1952) reports that breed- is found in Lake Malombe (Konings ing males are blue and gold. recorded it in 1988-89). Konings MAXIMUM SIZE: 20cm TL (Konings (1990a) comments on the larger 1990a). amount of red pigment in males from DIET: It feeds on the scales of tilapi- Malombe, suggesting that he has cer- ine cichlids. Its resemblance to these tainly observed it in person. I did not fishes is presumably an adaptation to see this species there in 1991-92, and it allow it to approach them undetected may have been exterminated by (Eccles & Trewavas, 1989). seining. DISTRIBUTION AND ABUNDANCE: A rare COMMERCIAL IMPORTANCE: Frequently species only found in the company of seen in seine net catches. Occasionally its host, the chambo, Oreochromis exported as an aquarium fish. (Nyasalapia) spp. Jackson (1961) reports NOTES: Eccles & Trewavas consider that C. shiranus is confined to the south Haplochromis sexfasciatus Regan to be a of the lake, but Eccles & Trewavas state junior synonym. Konings (1990a) that it is found throughout Lakes Ma- records specimens from Chisumulu lawi and Malombe and the Upper Island under the name Placidochromis Shire River, but it was not recorded in 'Johnstoni Gold', but later decided it is their trawl survey, nor in the 1992 sur- probably a local race of the species vey. The low power of the vessel used (Konings 1995b). He also observed for both surveys meant that it did not another deeper bodied species which make substantial catches of the large, he called P. 'johnstoni solo'. In body fast-swimming tilapiines with which shape and behaviour this strongly re- C. shiranus lives. From 1990 to 1992, sembles the species of the Protomelas despite sorting tens of thousands of taeniolatus complex. Oreochromis species for length-fre- quency analysis, I rarely found this species, recording fewer than 20 speci- Boulenger mens in the SE Arm trawl fisheries and colour photo on page 163 none in Lake Malombe or the Upper Shire River. DISTINGUISHING FEATURES: Silvery-grey COMMERCIAL IMPORTANCE: Probably colour and broad grey vertical bars are caught by all gears (beach seines, gill very similar to immature Chambo, nets, semipelagic trawl) taking large Oreochromis (Nyasalapia) spp. Distin- numbers of Oreochromis, but never guished from all other vertically- likely to make up a substantial propor- barred species by the large flat areas tion of the catch. of closely packed jaw teeth. C. taeniatus

128 Chapter 13 Buccochromis

A genus of large, heavily-built pisci- Buccochromis nototaenia vores. All have large, deep heads and (Boulenger) a prominent oblique stripe. Eccles & colour photo on page 164 Trewavas's key provides no characters which unambiguously distinguish this DISTINGUISHING FEATURES: A large heav- genus from Mylochromis (= Maravi- ily-built species with a large mouth chromis), but the overall appearance and head, and a prominent oblique does differ, Buccochromis tending to stripe. Deeper bodied than B. lepturus, have a heavy-headed, predatory look. B. spectabilis, and B. rhoadesii. Distin- Buccochromis species are well-repre- guished from B. atritaeniatus (which sented in the material examined by has not been identified in the field) by Eccles & Trewavas — no less than 170 its longer premaxillary pedicels. The specimens from the 7 nominal species shallow preorbital and different posi- — but none of the material postdates tion of the oblique stripe distinguishes the Christy collection of the 1920s and B. heterotaenia, which is generally a the genus has not been seriously col- darker colour and usually associated lected or studied since. I have never with rocky shores. According to Eccles positively identified B. atritaeniatus, B. & Trewavas, specimens above 16cm SL oculatus or B. spectabilis in the field. can be distinguished from those of B. Konings (1990a, 1995b), who has more oculatus by the shorter preorbital bone experience of shallow water and north- (25-29.5% of head length, compared to ern areas has not seen the latter two 21-25%), but this character does not species either, but provides illustra- hold for smaller specimens. tions purporting to show B. atritaenia- COLOUR: Females and immature tus. Measurements I have taken from males silvery-yellow, darker dorsally. the illustrations of the ratio of the pre- Generally there is a prominent black maxillary pedicel to head length (given oblique stripe. Pelvic and anal fins and as diagnostic by Eccles & Trewavas) the lower half of the caudal fin are indicate that these specimens are not bright orange. Ripe males are bright B. atritaeniatus, but could be either B. blue with golden flanks. Some of the nototaenia, B. heterotaenia or some other scales behind the operculum have a species. Similarly Axelrod & Burgess's bright red spot. Dorsal fin margin or- (1977) photograph also appears to be ange or red, sometimes whitish proxi- of B. nototaenia. B. heterotaenia is largely mally. Anal fin dark, with a broad or- a rocky shore species. See Eccles & ange margin and numerous egg-spots. Trewavas for identifying features and MAXIMUM SIZE: 37cm TL (Eccles & Konings (1995b) for an accurate ac- Trewavas). count of its biology and live colours. DIET: Morphology suggests a pisci-

129 vorous diet. NOTES: Juveniles are common in REPRODUCTION: Occasionally 8-10 heavily fished areas south of Boadzulu large ripe males have been taken to- Island and in Lake Malombe. Some gether in trawl hauls at a depth of 10- small specimens lacking the orange 20m off clean sandy beaches, such as pigment were collected in the 1992 sur- the Chapola Shoal. This suggests lek vey. Since there is presently no way of breeding. Konings (1995b, quoting distinguishing small specimens of this Mwale) states that males build sand species from those of B. oculatus, it is castle nests, occasionally among rocks, not known whether this indicates that but usually on sand. Females guard fry the orange colour is not always shown for up to 4 weeks, usually in the vicin- by this species. ity of rocks (Konings 1995b). DISTRIBUTION AND ABUNDANCE: In the 1992 survey, this was by far the most Buccochromis cf. oculatus numerous species of the genus and one (Trewavas) of the most common large predators colour photo on page 164 in the SE Arm. It was the only species of the genus positively recorded from DISTINGUISHING FEATURES: A large heav- Lake Malombe. Also collected from ily-built species with a large mouth Domira Bay. Recorded from 14 out of and head, and a prominent oblique 25 stations at depths of 18-44 m south stripe. Deeper-bodied than B. rhoadesii, of Monkey Bay, but in only two of the B. spectabilis, and B. lepturus. It has a hauls in SE Arm north of Monkey Bay. larger mouth than B. atritaeniatus and Konings records a distribution encom- a shallower preorbital than B. noto- passing all of Lakes Malawi and Ma- taenia. lombe. Eccles & Trewavas examined COLOUR: Live colours not known, al- material from the SE and SW arms and though a specimen tentatively as- from Karonga and Mwaya in the north. signed to this species was generally They did not record the species in their greyish with an oblique dark stripe. trawl surveys, because of the difficulty MAXIMUM SIZE: 27 cm TL. in distinguishing it from congeners. It DIET: Morphology suggests a pisci- seems likely that their distribution and vorous diet. abundance records from B. atritaenia- DISTRIBUTION AND ABUNDANCE: The tus actually refer to this species. lectotype of B. oculatus was collected Jackson (1961) considered that this spe- from Monkey Bay. All of the para- cies was common and well-known, fre- lectotypes are from the south-eastern quented sheltered sandy areas and re- arm or from unknown locations. Eccles placed B. oculatus in the southern parts & Trewavas (1989) report the species of the lake. from a single trawl haul at 18m depth COMMERCIAL IMPORTANCE: Frequently at 'the narrowest part of the lake'. taken in small numbers in the seine, Konings (1990a) does not record this pair trawl and midwater trawl catches. species. I have tentatively assigned to Likely to be locally important in hand- the species a single specimen from 35m line and gill net catches. depth off Chirombo Bay.

130 COMMERCIAL IMPORTANCE: Probably ticularly on the head, with numerous minor. yellow-orange spots on the flanks and NOTES: Eccles & Trewavas's descrip- spots and stripes on the dorsal and tion of the species lists 20 types, of caudal fins. There are numerous egg- which 18 are known to have come from spots on the anal fin. the SE Arm, all collected by Christy in MAXIMUM SIZE: 34 cm TL (Eccles & the 1920s. The area is both the most Trewavas). heavily-fished and the best-studied DIET: Morphology suggests a pisci- part of the lake. The rarity or absence vorous diet. of the species in subsequent surveys REPRODUCTION: Konings (1995b) re- suggests that (i) B. oculatus has not been ports that males are not territorial and adequately distinguished from conge- single sexually active males are often neric species in the field; (ii) it is syn- seen in groups of normally-coloured onymous with B. nototaenia; or (iii) individuals. In the aquarium, they do populations have declined sharply not build nests but spawn among rocks during the last 70 years. (Konings 1990a). DISTRIBUTION AND ABUNDANCE: A wide- ly distributed, but uncommon fish of Buccochromis rhoadesii (Boulenger) fairly shallow sandy areas. The collect- colour photo on page 164 ing locality of the holotype is un- known. Although Jackson (1961) con- DISTINGUISHING FEATURES: A large heav- sidered this to be a species of the south- ily-built species with a large mouth ern part of the lake, material examined and head, and a faint oblique stripe. by Eccles & Trewavas came from the The lower half of the body and lower SE Arm and the far north of the lake fins are bright yellow. It is more slen- between Vua and Mwaya. Konings der than B. nototaenia and deeper-bod- (1990a) states that it is found through- ied and with a longer snout and larger out the lake, and illustrates specimens mouth than B. lepturus. from Chisumulu Island. He records it COLOUR: Females and immature as living in small groups at 15-30m males silvery, darker dorsally. The ob- (1995b). Eccles & Trewavas report the lique bar is prominent, black and con- species from 18m depth in the SE Arm tinuous in smaller fish, but is faint grey and in shallower non-systematic and often obscured by vertical barring trawls, but were uncertain of their abil- in larger specimens. The lower jaws, ity to identify the species in the field. chest, lower part of the operculum, In the 1992 survey, it was recorded on pelvic and anal fins and lower part of single occasions from the Namiasi, the caudal fin are bright yellow orange. Michesi, Nkope, and Mazinzi stations In larger specimens, the yellow colour and was abundant at the latter. All of extends further dorsally. Individuals the samples came from trawls at from Chisumulu Island (illustrated by depths of 20-26 m. Konings, 1990a) may be entirely yel- COMMERCIAL IMPORTANCE: A minor low. Territorial males (Konings, 1990a, component of the shallow water trawl Spreinat, 1992a) are bright blue, par- and seine fisheries. Exported, under

131 the names 'Haplochromis lepturus' or tained fish remains, plant material, and 'Yellow Lepturus', in fairly small num- fatty tissues which may be gastropod bers for the ornamental fish trade, remains (Grant & Turner, unpub- mainly from Likoma Island (Spreinat, lished). 1992a). REPRODUCTION: Konings (1990a) re- ports that in the aquarium they do not build nests but spawn among rocks. Buccochromis lepturus (Regan) Konings (1995b) reports that males are colour photo on page 164 not territorial and single sexually ac- tive males are often seen in groups of DISTINGUISHING FEATURES: A large heav- normally-coloured individuals. ily-built species with a large mouth DISTRIBUTION AND ABUNDANCE: A shal- and head, and a prominent oblique low water species, uncommon in trawl stripe. More slender than most other catches. Jackson (1961) reports that this congeneric species. Distinguished from species is characteristic of open sandy B. spectabilis (which has not been iden- beaches and is more common north of tified in the field) and Mylochromis 'tor- Nkhotakota. Reported by Konings pedo' by its larger mouth (lower jaw (1990a) to occur throughout Lakes 43.5-50% HL in B. lepturus; 40-41.7% in Malawi and Malombe, although I did B. spectabilis; 36% in M. 'torpedo'.). not observe it in the latter. Konings spp. and the elongate (1995b) has observed small groups at Mylochromis (M. formosus, M. gracilis, 15-30m depth. Frequently taken in M. spilostichus) are more slender with small numbers in the 1992 survey at smaller head and jaws. depths of 20-26m. Material examined COLOUR: Females and immature by Eccles & Trewavas came from the males silvery, darker dorsally, with a northern and southern ends of the lake bright greenish cast. Prominent black and these authors report the species as oblique stripe. According to Eccles & common in shallow (10m deep) sandy Trewavas, breeding males are sky blue, areas. with yellow spots on the flank scales. COMMERCIAL IMPORTANCE: Currently of Fins grey, with yellow spots on the soft- very minor importance in the south, rayed portions. The anal fin has a although according to Jackson (1961) broad yellow margin, with several often taken in seines. Occasionally ex- large white spots. Konings (1990a, ported for the ornamental fish trade. 1995b) illustrates ripe males which NOTES: According to Eccles & Trewa- more or less correspond to this descrip- vas, Haplochromis gigas Ahl is conspe- tion, but are somewhat greenish-blue. cific with B. lepturus. MAXIMUM SIZE: 42cm TL (Eccles & Trewavas, 1989). DIET: Morphology suggests a pisci- vorous diet. The stomach of two speci- mens (10-11cm SL) were examined: one was filled by 2cm long fish, probably of the genus Aulonocara, another con-

132 Chapter 14 Mylochromis and other oblique- striped species

The oblique striped pattern is appar- long snout and ventrally-positioned ently not found in any cichlid species mouth. In the south of the lake, it is except the endemic haplochromines of the only large, long-snouted species Lake Malawi (Eccles & Trewavas, with Lethrinops-type dentition and a 1989). This pattern is associated with a prominent black oblique stripe. It is number of very different morph- further distinguished from T. furcicauda ologies, which are placed by Eccles & by its longer snout, and from both T. Trewavas into a variety of genera. Most furcicauda and T. praeorbitalis by its of the morphologically unspecialised heavy-headed robust appearance. Dis- or molluscivorous deep-bodied forms tinguished from M. lateristriga and were placed in Maravichromis. Derijst Lichnochromis by dentition and greater & Snoeks (1992) discovered that this is body width, and from T. brevis by the a junior synonym of Mylochromis. longer snout, smaller eye, different Other oblique-striped species formerly form of anteriormost lower gillraker included in the genus Lethrinops, on the and of the lower pharyngeal bone. basis of the shape of the dental arcade, COLOUR: Females and immature have been placed by Eccles & Trewa- males dark grey dorsally, flanks yel- vas into the genera Taeniolethrinops and lowish, belly white. A continuous black Tramitichromis. Tramitichromis also in- oblique bar runs from just anterior to cludes species with very different the dorsal fin to the caudal peduncle. melanic patterns, but similar jaw and There are often 8-9 faint vertical bars pharyngeal bone morphology. under the dorsal fin. None of the oblique striped species MAXIMUM SIZE: 25cm TL. are found in very deep water (>100m). DIET: The stomachs of 2 specimens Some of the Mylochromis species have (12.5-14cm SL) contained mostly chiro- never been positively recorded since nomid larvae. Large quantities of sand, their original description, and it is detritus, and algal remains also in- probable that most of these are from gested (Grant & Turner unpubl.). very shallow water. REPRODUCTION: Robinson (1992) ob- served a female guarding fry off Thumbi East Island during October. Taeniolethrinops laticeps (Trewavas) DISTRIBUTION AND ABUNDANCE: In the colour photo on page 165 1992 survey, it was taken in trawl catches from 18-35m depth and was DISTINGUISHING FEATURES: One of a often quite numerous. It appeared to group of species characterised by a be confined to the eastern coast north

133 of the Chapola Shoal and to the area from most related species in the very from Nkhudzi to the Nankumba pe- long snout. More laterally compressed ninsula. A single large specimen was than T. laticeps, which also has differ- also collected from Domira Bay. I have ent dentition. Lichnochromis acuticeps, observed this species while Scuba div- which appears to be confined to rocky ing just outside Monkey Bay. shores, is less deep bodied, less wide, COMMERCIAL IMPORTANCE: Caught in and has a longer, less ventrally-angled pair trawl, gill net and seine catches. snout and more conspicuous teeth. NOTES: Previous authors doubted COLOUR: Females and immature whether this species was distinct from males silvery, darker dorsally with a T. praeorbitalis (e.g. Jackson, 1961; Ec- conspicuous black oblique bar. Ripe cles & Trewavas, 1989; Konings, 1989, males blue, turquoise or yellowish 1990a). In the southern part of the lake, ventrally with orange margin to flank the two species can be clearly distin- scales. Dorsal and caudal fins with or- guished, and although they prefer dif- ange-red spots. Dorsal margin white ferent habitats, they may be found with red-tipped lappets. Anal dark sympatrically and indeed may even with broad orange distal margin and forage together. I have not collected numerous with spots and streaks these species to the north of Domira (Konings 1989, 1990a). Bay, but I have observed some indi- MAXIMUM SIZE: 22cm TL. viduals while diving at Nkhata Bay. DIET: Arthropods (Eccles & Trewavas, These appeared to be more laterally 1989). compressed than typical T. laticeps, but REPRODUCTION: Konings (1990a) re- exhibited a prominent black bar not ports that males defend a territory seen in southern specimens of T. prae- among rocks, although males in breed- orbitalis. These specimens (at Nkhata ing colour may be observed foraging Bay) are similar to the type of Lethri- over sand. nops macrorhynchus Regan, which I DISTRIBUTION AND ABUNDANCE: A shal- have examined, and other small speci- low water species, sometimes ob- mens collected from Benga (northern served while snorkelling, but rarely Domira Bay). L. macrorhynchus has encountered in trawl catches. I col- been considered as a junior synonym lected this species in a few shallow of T. praeorbitalis since Trewavas's re- water trawl hauls, but not in the 1992 vision of 1931. The type is in poor con- surveys. It was often found in experi- dition — it has been cut in two. I have mental gill net catches in the Upper not yet examined the types of Lethri- Shire River. The type is a skin from an nops fasciatus Ahl, another species unknown location. Most of the speci- synonymised with T. praeorbitalis. mens examined by Eccles & Trewavas (1989) came from the southern end of the lake, but a single male was col- Mylochromis lateristriga (Günther) lected from Vua in the north. Eccles & Trewavas did not record it from their DISTINGUISHING FEATURES: A deep-bod- trawl surveys and Jackson (1961) re- ied oblique striped species. Differs ports it only from seine catches in the

134 Figure 38. Mylochromis lateristriga. Aquarium photo.

SE Arm. Konings (1990a) has recorded tition. It has fewer gill rakers than all the species only from the southern part Buccochromis and all Mylochromis ex- of the lake, mainly Senga Bay and the cept M. anaphyrmus, which has much Maleri Islands, although he also re- larger molariform teeth. It has a shorter ports a similar form from around snout and different shaped lower pha- Makanjila. ryngeal bone than species of the genus COMMERCIAL IMPORTANCE: May be lo- Taeniolethrinops. cally important in gill net and seine Colour (from Konings 1990a): Fe- catches, but not in trawls. males and immature males silvery grey NOTES: Konings (1990a) considers with conspicuous black oblique bar. that the more slender specimens found Ripe males blue, with numerous or- around Makanjila may represent a dif- ange spots and stripes in dorsal and ferent species from the deeper-bodied caudal fins. Anal fin dark with several form found at Senga Bay and the large yellow-orange egg-spots and Maleris. white margin. Pelvics dark with white leading edge. Dorsal with white mar- gin and orange lappets. Branchiostegal Tramitichromis brevis (Boulenger) membranes and chest orange. MAXIMUM SIZE: 16cm TL (Konings DISTINGUISHING FEATURES: A medium- 1990a). sized squat species with a large eye. DIET: Chironomid larvae (Fryer, 1959, Distinguished from other short- Eccles & Trewavas, 1989). snouted species of the Lethrinops group REPRODUCTION: Konings (1990a) re- by the prominent oblique black bar. ports that males build small sand cas- Differs from species of the genera tle nest near rocks. They form small Buccochromis and Mylochromis in the aggregations. shape of the first lower gillraker (broad DISTRIBUTION AND ABUNDANCE: Appar- and flat) and the Lethrinops-type den- ently found throughout the lake

135 Figure 39. Tramitichromis brevis, male, from Chisumulu Island (aquarium photo).

(Konings, 1990a). Eccles & Trewavas odon. Lower pharyngeal bone with report it from Nkhata Bay and the large heavy molariform teeth. M. 'deep' south, including a single trawl sample has a more blotchy oblique stripe and from 9m depth. Jackson (1961) re- a larger eye. ported it as abundant. It was not found COLOUR: Silvery on flanks, darker in the 1992 survey, but I did record it dorsally often with metallic green cast. in shallow water hauls around Makan- Females and immature males with con- jila. spicuous black oblique bar. Ripe males COMMERCIAL IMPORTANCE: Minor, in the pale turquoise on head and back. Dor- south at least. sal fin with bright yellow lappets. Anal fin with yellow lower margin and nu- merous egg-spots. Mylochromis anaphyrmus MAXIMUM SIZE: 23cm TL (Konings (Burgess & Axelrod) 1990a). colour photo on page 165 DIET: Morphology suggests that this species is mainly a pharyngeal crusher DISTINGUISHING FEATURES: A deep-bod- of molluscs. Five specimens (8-13cm ied, oblique-striped species. Differs SL) examined (Grant & Turner, un- from most related species in its small publ.) all contained large quantities of rounded head and small mouth. Rela- molluscan remains, mainly of gastro- tively smaller eye than T. brevis. pods. Considerable quantities of mol- Shorter, more rounded snout and lusc shell fragments were ingested. fewer gillrakers (8-9) than M. sphaer- Some individuals also contained a lot

136 of sand, detritus, and algal remains. catch by weight. Copepods, chironomids, and other ar- thropod material were also found. Mylochromis sphaerodon (Regan) REPRODUCTION: Robinson (1995) ob- served a small breeding arena at 5m DISTINGUISHING FEATURES: A deep-bod- depth. Males constructed small, asym- ied, oblique striped species. Differs metrical bowers. These fish were not from most related species in the small collected, and it is not certain that they rounded head and small mouth. Rela- were M. anaphyrmus. tively smaller eye than T. brevis and M. DISTRIBUTION AND ABUNDANCE: Eccles & 'deep'. Longer, more pointed snout Trewavas state that the species is com- than M. anaphyrmus. Lower pharyn- mon at 15-35m depth in the south of geal bone with large heavy molariform the lake and also report it from teeth. Nkhotakota. Konings (1990a) states COLOUR: Females and immature that it is mainly found at depths greater males with conspicuous black oblique than 10m in silty areas. In the 1992 sur- bar. Silvery on flanks, darker dorsally. vey, it was found in 42 out of 57 trawl Pelvic and anal fins bright yellow. Ripe samples at depths of 18-72m. It was males bluish, with orange spots on often one of the most abundant spe- flank scales, caudal and dorsal fins, cies. I also found this species at Domira and a white dorsal fin margin (Eccles Bay. Konings (1995b) suggests it is con- & Trewavas). Konings (1995b) illus- fined to the southern and western parts trates a ripe male, apparently of this of the lake. species, which is bright blue, with a COMMERCIAL IMPORTANCE: Occurs in yellow chest and belly. most samples of pair trawls and semi- MAXIMUM SIZE: 20cm TL (Konings pelagic trawls in the SE Arm, but does 1990a). not comprise a large proportion of the DIET: Morphology suggests that this

Figure 40. Mylochromis spaerodon, female, South-East Arm.

137 species is mainly a pharyngeal crusher COLOUR: Females and immature of molluscs, and this was reflected in males with conspicuous black oblique the stomach contents of a single 7.5 cm stripe. Silvery on flanks, darker dor- SL specimen which contained large sally. Mature male (perhaps not fully amounts gastropod material, along ripe) with dark blue head, vertical bars with copepods, cladocera, and algae under the dorsal fin, and dark un- and with some chironomid remains paired fins. Konings (1989) reports that (Grant & Turner, unpublished). male M. balteatus has a blue head, DISTRIBUTION AND ABUNDANCE: Types golden nape and grey-blue body. from unknown location. Eccles & MAXIMUM SIZE: At least 16cm TL. Trewavas examined one specimen DISTRIBUTION AND ABUNDANCE: M. cf. from Monkey Bay and state that it is a balteatus has been positively identified common shallow water species, but from a trawl catch at 25-30m in the SE did not record it in their trawl survey. Arm, south-west of Boadzulu Island. Konings (1990a) states that it has a In the 1992 survey it was tentatively lake-wide distribution but is less com- recorded from 3 trawl samples be- mon in the SE Arm. He illustrates tween 20 and 34m in the SE Arm and specimens from Senga Bay and the also from seine catches in the SE Arm 'southern part of the lake'. In his 1989 and Lake Malombe. The types of Mylo- book, he states that it is only found at chromis balteatus (Trewavas) are from Cape Maclear, the SE Arm and the the vicinity of Karonga, but three small coast near Makanjila. In his 1995 book, specimens from the SW Arm were as- he states it is confined to the southern signed to this species by Eccles & part of the lake. Jackson (1961), who Trewavas. Konings (1995b) has ob- worked mostly in the north, gives no served M. balteatus at Karonga in the information on its occurrence. In the north and Londo on the Mozambique 1992 survey, I recorded it from a single coast, but not elsewhere and suggests station at 26m off Chekopa in the that it may be confined to the north of north-eastern part of the SE Arm. It the lake, but has collected similar speci- seems to be a fairly uncommon patch- mens from Makanjila and Ntekete. ily-distributed species of shallow COMMERCIAL IMPORTANCE: Negligible sandy areas. in the south. Konings (1990a, 1995b) COMMERCIAL IMPORTANCE: Minor, but reports that it is common in beach seine may be locally important in seines. catches in the north. NOTES: Further work is required to determine whether northern and Mylochromis cf balteatus southern populations are conspecific. colour photo on page 16

DISTINGUISHING FEATURES: A deep-bod- Mylochromis ericotaenia (Regan) ied, oblique-striped species. Differs colour photo on page 165 from most related species in the larger head and jaws, large eye and concave DISTINGUISHING FEATURES: A deep-bod- profile above the eye. ied species, with a pointed snout and

138 Figure 41. Mylochromis ericotaenia, male, South-East Arm.

small mouth. The oblique band is gen- NOTES: The type specimens are very erally represented as a series of spots. small juveniles, and it is possible that COLOUR: Live coloration of females the larger specimens examined by Ec- and immature males silvery, darker cles & Trewavas (1989), on which my brown dorsally with a conspicuous identification is based, may represent oblique row of dark spots and 5-6 faint a different species. Some of the speci- vertical bars under the dorsal fin. mens illustrated by Konings (1990a) MAXIMUM SIZE: At least 21cm TL. have bright orange pelvic and anal fins. DIET: Insect larvae and molluscs (Ec- Others have clear fins. A single male cles & Trewavas, 1989). illustrated was blue with orange spots DISTRIBUTION AND ABUNDANCE: Rare in on the unpaired fins and a yellow 1992 trawl surveys: 4 specimens were chest. The anal fin is orange with a few taken in a single haul at 25m depth off egg-spots. These may represent a fur- the north-eastern shore of the SE Arm ther species. Konings (1990a) report and another one just north of Boadzulu that the guts of two specimens con- Island. According to Eccles & Trewa- tained phytoplankton, refers to this vas, who examined no less than 70 yellow-finned form (Konings pers. specimens from collections made in the comm.). 1920s and 30s, it is distributed through- out the lake, but was not recorded from their trawl surveys. Konings (1990a) Mylochromis melanonotus (Regan) reports seeing a few specimens at colour photo on page 166 Chiwindi in Mozambique and in Senga Bay and just south of Nkhata Bay at DISTINGUISHING FEATURES: A deep bod- Nkhomo Reef and Kande Is. Jackson ied species with the mouth less ven- (1961) was not familiar with the spe- trally placed than in most other simi- cies. lar oblique-striped species. In life, con- COMMERCIAL IMPORTANCE: Minor. spicuously bright yellow. Some

139 specimens with a characteristically flat- morphology, Eccles & Trewavas (1989) tened lower jaw. placed Haplochromis melanonotus in the COLOUR: Females and immature monotypic genus Platygnathochromis. males bright yellowish, more intense Similar specimens with more typical ventrally with prominent black oblique morphology had been described as stripe. Ripe males bright blue with Haplochromis semipalatus Trewavas. Ec- ochre spots on flanks and unpaired cles & Trewavas subsequently allo- fins. Anal fin dark, with broad orange cated H. semipalatus to the genus Mara- margin and prominent white egg-spots vichromis (now known to be a junior (Eccles & Trewavas, 1989; Baasch, synonym of Mylochromis Derijst & 1993). Snoeks 1992). Konings (1993b) reports MAXIMUM SIZE: 25cm TL (Konings, that the jaw shape is polymorphic, 1990a). even within a single breeding arena. DIET: Said to be small fish and mol- Consequently he considers M. semi- luscs (Eccles & Trewavas, 1989). Ko- palatus to be synonymous with P. mela- nings reports instances of raiding cat- nonotus, and that the latter properly fish nests (1993b) and also of cleaning belongs in Mylochromis. I concur with behaviour (1995b). The stomach of a this view, as the polymorphism in the single small individual (75mm TL) jaw shape clearly indicates this feature contained mostly cladoceran and is of no phylogenetic significance. copepod remains, along with algae, macrophytes, and sand (Grant & Turner unpubl). Mylochromis melanotaenia (Regan) REPRODUCTION: Konings (1990a) has colour photo on page 166 observed breeding colonies at depths of around 20m. Small sand castle nests DISTINGUISHING FEATURES: A deep-bod- or simple clear areas were defended. ied, oblique-striped species distin- In the aquarium, Baasch (1993) ob- guished by its relatively compressed served that males excavated a shallow shape, large jaws and eye, and thick depression in the sand. Females laid lips. around 60 eggs. COLOUR: Females and immatures sil- DISTRIBUTION AND ABUNDANCE: Material very with prominent black oblique examined by Eccles & Trewavas was stripe. Mature male (probably not fully collected from the SE and SW Arms ripe) has silvery flanks with orange- and the far north of the lake. Konings brown cast, dark blue head, fins dark (1990a) claims it has a lakewide distri- with orange spots. White dorsal fin bution, mainly over sand, but occasion- margin and yellow anal fin margin. ally over rocky substrates. During the MAXIMUM SIZE: 18cm TL. 1992 survey, it was often taken in small DISTRIBUTION AND ABUNDANCE: Rare. numbers at depths of 20-30m in the SE Known only from the south of the lake. Arm, north of Boadzulu Island. I also Only 7 specimens have been reported. found it in a trawl catch at 46-50m Two were taken by myself from a trawl depth, off Chirombo to Nkhudzi. near the southern tip of the SE Arm at NOTES: Because of its unusual jaw 15-18m depth. The three types, col-

140 lected by Wood, are of unknown prov- stripe is brownish and is broken into enance. Eccles & Trewavas identified 7-8 spots on the body and another at a single specimen from the Christy col- the posterior end of the caudal pedun- lection, from Malembo in the SW Arm. cle. Each spot may be continued as a Jackson (1961) records a single speci- faint vertical bar. Mature male (prob- men from a seine haul in Monkey Bay. ably not fully ripe) generally yellow- Konings (1989) describes the feeding ish, with a blue head and dorsum. behaviour of the fish and notes its oc- Dorsal and caudal fins dark with or- currence on the southern and eastern ange spots. Dorsal margin white with shores, but in his 1990 book claims not orange tips. Anal fin dark with whit- to have seen the species alive. The 1989 ish egg-spots. record appears to be a case of mistaken MAXIMUM SIZE: 18cm TL. identity (Konings pers. comm.). DISTRIBUTION AND ABUNDANCE: Re- COMMERCIAL IMPORTANCE: Negligible. corded only by the author and only from the Maleri Islands and Domira Bay areas at depths of 25-45m. Mylochromis 'deep' COMMERCIAL IMPORTANCE: Unknown. colour photo on page 166

DISTINGUISHING FEATURES: A deep-bod- Mylochromis plagiotaenia (Regan) ied, oblique-striped species distin- colour photo on page 166 guished by its relatively compressed shape. It has fewer lower gillrakers (9) DISTINGUISHING FEATURES: A small ob- than most other species (M. melanono- lique-striped species, with a delicate, tus, M. guentheri, M. obtusus, M. mollis, small-headed appearance. It has fewer M. epichoralis, M. mola). It has a smaller gillrakers (8-11) and a more ventrally- eye than M. cf. balteatus and lacks the positioned mouth than M. 'chekopae'. thick lips of M. melanotaenia. M. COLOUR: Females and immature labidodon and M. plagiotaenia have a males generally dark brownish dorsal- smaller head and more slender build, ly, with the oblique stripe often in the and M. incola (a rocky shore species) form of blotches which run together. has a longer snout. The eye is larger Konings (1990a) illustrates a ripe male (29-33% HL in fish of 12-17cm SL) than which has a bluish green colour, shad- that of similar-sized M. anaphyrmus ing to yellow ventrally. Dorsal and anal (26-27% HL). M. sphaerodon has a more fins are spotted with orange. slender shape and pointed snout and MAXIMUM SIZE: 14cm TL (Konings, orange pelvic and anal fins. Like M. 1990a). ericotaenia, the oblique stripe is gener- REPRODUCTION: Konings (1990a, ally expressed as a series of spots, but 1995b) reports that males clear a small that species has a less ventrally-placed area among vegetation in shallow wa- mouth, smaller head, and fewer verti- ter. cal bars. DISTRIBUTION AND ABUNDANCE: The COLOUR: Females and immatures sil- types and other material examined by very with a brownish cast. The oblique Eccles & Trewavas were collected from

141 unknown locations. Jackson (1961) de- Mylochromis 'chekopae' scribes it as a fish of sandy shores and calm waters, more common in the DISTINGUISHING FEATURES: A relatively south. Konings (1990a) reports this slender small-headed species, which species from the Nankumba Peninsula. has a body shape reminiscent of some I did not identify it in the 1991 or 1992 Copadichromis spp.. Distinguished from trawl surveys, but collected it from a most Mylochromis spp. by the higher single haul of the photographic survey number of gillrakers (13) and lateral in October 1991, in shallow waters (20- line scales (36-37) and fewer tooth rows 28m) south of Boadzulu Island. It may in lower jaw (2). Distinguished from be more numerous in shallower water. M. 'double spot' by the continuous COMMERCIAL IMPORTANCE: Minor. oblique stripe, higher number of gill- rakers, smaller eye and less ventrally placed mouth. COLOUR: Live colours not known. Pre-

Figure 42. Mylochromis 'chekopae', male, South-East Arm.

Figure 43. Mylochromis 'chekopae', female, South-East Arm.

142 served females and immature males Otopharynx 'productus' in that the sandy, paler ventrally, with prominent stripe is continuous and not broken up thin black oblique bar, occasionally into spots. The head profile is more broken into a series of spots. Ripe curved than that of M. formosus. males darker, but with bar still visible. COLOUR: Sandy, paler ventrally. Black Snout and head dark. Fins dark, dor- oblique stripe generally reaches about sal and anal with pale margin. half-way along the caudal peduncle. MAXIMUM SIZE: 15cm TL. Pelvic fins yellowish. Ripe males: head DIET: Stomachs of two specimens ex- dark blue; flanks dark metallic gold; amined indicate that they had been fins dark; anal with brilliant red lower feeding in the water column. Clado- margin; dorsal margin pinkish white. cera, copepods, and algae were found, MAXIMUM SIZE: 26cm TL. but no sand or detritus (Grant & DISTRIBUTION AND ABUNDANCE: Rare. Turner, unpubl.). Not recorded in 1992 survey, but taken DISTRIBUTION AND ABUNDANCE: Known in a haul from depths of 15-23m, near only from shallow waters (18-45m) off Boadzulu Island in the SE Arm. the north-eastern shore of the SE Arm, COMMERCIAL IMPORTANCE: Negligible. at the trawl stations Chekopa, Chi- NOTES: Konings (1989) illustrates a linda, and Chiponda. In the shallower species with a similar body shape un- part of its depth range, it is often the der the name Sciaenochromis species most abundant species. 'silver torpedo', but it is not mentioned COMMERCIAL IMPORTANCE: Minor, at in the text. In his 1990 book, it is la- present, but potentially locally impor- belled Maravichromis 'silver torpedo' tant. and said to be from Senga Bay. No melanin pattern is evident in the pho- tographs and it is possible that Ko- Mylochromis 'torpedo' nings' species may be O. 'productus'. colour photo on page 166

DISTINGUISHING FEATURES: An elongate, Champsochromis caeruleus oblique-striped species. Distinguished (Boulenger) from M. gracilis by its more pointed colour photo on page 167 snout and more laterally compressed body. Mouth more ventrally-placed DISTINGUISHING FEATURES: A slender, than Champsochromis species, which elongate, oblique-striped species, it has also have larger, more widely-spaced a rather cylindrical streamlined shape teeth. More elongate and laterally com- reminiscent of those of the horizon- pressed than most Buccochromis spe- tally-striped Rhamphochromis esox and cies. Compared to B. spectabilis the R. leptosoma. It has larger jaws and interorbital width is narrower (25-26% more widely-spaced teeth and larger head length in B. spectabilis; 20-21% in jaws than Mylochromis formosus and a M. 'torpedo') and the lower jaw shorter slimmer body and less prominent pre- (40-41.6% head length in B. spectabilis; maxillary pedicel than Champsochromis 36% in M. 'torpedo'). Differs from spilorhynchus. The pelvic fins and soft

143 dorsal and anal of ripe males are con- known as the 'trout cichlid'. siderably elongated. NOTES: Eccles & Trewavas (1989) con- COLOUR: Females and immature sider Haplochromis bellicosus Ahl to be males silvery, darker dorsally with a synonymous with this species. slender continuous black oblique band and a conspicuous black stripe on the preorbital bone. Ripe male (illustrated Champsochromis spilorhynchus by Konings, 1990a) similarly coloured (Regan) to that of C. spilorhynchus. colour photo on page 167 MAXIMUM SIZE: 32cm TL (Eccles & Trewavas, 1989). DISTINGUISHING FEATURES: A slender DIET: Feeds on small pelagic fish, es- elongate oblique-striped species, dis- pecially Engraulicypris sardella tinguished from most similar species (Konings 1990a). by the possession of a black stripe on REPRODUCTION: Reitz (1993) gives an the preorbital bone. It has larger jaws account of captive propagation. Eggs and more widely-spaced teeth than were laid on an open flat sandy sur- Mylochromis formosus and a deeper face — no bower was built. Females body and more prominent premaxil- were found to lay 50-200 eggs. lary pedicel than Champsochromis DISTRIBUTION AND ABUNDANCE: Eccles & caeruleus. Trewavas recorded this species as com- COLOUR: Females and immature mon in the SW Arm at depths of 15- males silvery, darker dorsally; a slen- 55m and less often in shallow water in der continuous black oblique band and the SE Arm. The preserved material a conspicuous black stripe on the they examined was from the extreme preorbital bone. A ripe male illustrated north and south of the lake. Jackson by Konings (1990a) is pale blue with a (1961) recorded this species from white dorsal fin margin, anal with a gillnets and occasionally seines and broad orange margin and numerous suggested it might be a midwater spe- pale spots. Another illustrated in his cies. Konings (1990a) also states that it later book (Konings, 1995b) is a darker is found throughout Lake Malawi and blue with faint vertical bars and an or- is an open water species. It was not ange dorsal fin margin. recorded in the ODA/SADC project MAXIMUM SIZE: 35cm TL (Konings, survey of the pelagic zone (Menz, 1995) 1995b). and is therefore not likely to be a truly DIET: Feeds on small fish (Eccles & offshore species. In the 1992 trawl sur- Trewavas, 1989). Konings (1990a) re- vey, it was found in a single sample at ports that it mainly feeds on small Co- 28m depth and photographed from padichromis. another trawl in shallow water. DISTRIBUTION AND ABUNDANCE: Eccles & COMMERCIAL IMPORTANCE: Minor. Per- Trewavas recorded this species from haps caught by chirimila, beach seine trawls between 15 and 55m in the SE and hook and line. Despite its large Arm and also in Upper Shire River. size, this species is a sought-after and Konings (1990a) also states that it is expensive aquarium fish, popularly found throughout Lake Malawi and

144 Malombe. Not recorded in the 1992 of the lake. Jackson regarded it as a rare trawl survey and not seen by the au- fish occasionally found off rocks. Ec- thor during an extensive survey of the cles & Trewavas did not record the spe- Upper Shire and Lake Malombe, al- cies in their trawl survey, but reported though frequently seen underwater it from ad hoc surveys at 10-15m depth. over sandy areas at Cape Maclear and Konings (1990a) records it from shal- in seine catches in shallow water low sandy areas at Senga Bay and the within Monkey Bay. The material ex- Maleri Islands, but states that it is amined by Eccles & Trewavas was all found throughout the lake. It was re- from the southern part of the lake (SE corded from a few samples at 25-65m Arm, SW Arm and Upper Shire) or in the 1992 trawl survey in the SE Arm from unknown locations. Jackson as well as at Domira Bay and the Maleri (1961) reported this to be a well-known Islands. commercially important species par- COMMERCIAL IMPORTANCE: Minor. ticularly in the SE Arm and Upper NOTES: Eccles & Trewavas (1989) re- Shire. port a ripe male from Malembo (SW COMMERCIAL IMPORTANCE: Minor. Per- Arm) which was blue with a white haps caught by chirimila, beach seine ventral surface and several rows of and hook and line. golden scales on the flanks. Dorsal and NOTES: Eccles & Trewavas consider anal fins with dull yellow spots and that Haplochromis longipes Regan is syn- stripes. Anal fin translucent with a onymous with this species. broad black margin tipped with red and several pale egg-spots. This colour description is very similar to that of M. Mylochromis formosus (Trewavas) 'torpedo' described above — a speci- colour photo on page 167 men I have checked and which is not M. formosus. The ripe male illustrated DISTINGUISHING FEATURES: A slender by Konings (1990a, 1995b) as Maravi- elongate oblique-striped species, dis- chromis sp. 'Kande' looks very similar tinguished from most similar species to M. formosus. It does not resemble the by the possession of a black stripe on colour description given by Eccles & the preorbital bone. It has more closely- Trewavas, having a white dorsal mar- packed teeth on smaller jaws than gin and lacking the red anal margin. Champsochromis species. Other specimens (Konings 1990a: COLOUR: Females and immature p.141, #6; p.156, upper photo; Konings, males yellowish with a broad continu- 1995b) look more like M. 'torpedo' but ous black oblique band and a conspicu- the lower lobe of the tail fin is yellow, ous black stripe on the preorbital bone. as Konings (1995b) points out, a fea- MAXIMUM SIZE: 15cm TL (Konings, ture of many Buccochromis species. 1990a). Clearly this group requires further DIET: Feeds on invertebrates and work. small fish (Konings, 1990a). DISTRIBUTION AND ABUNDANCE: The types were collected at Vua in the north

145 Mylochromis gracilis (Trewavas) livingstonii (Ps. lanisticola?) from gastro- colour photo on page 167 pod shells. REPRODUCTION: Baasch (1991) has ob- DISTINGUISHING FEATURES: An elongate served breeding in captivity. Males dig species with a prominent oblique a crater about 50cm in diameter. A stripe. Along with M. spilostichus this 15cm female produces about 100 eggs. species is distinguished from other DISTRIBUTION AND ABUNDANCE: Type elongate oblique-striped species by the material from Monkey Bay area. Eccles small head and large eye. It has a & Trewavas (1989) record the species smaller eye (23-24% HL), and deeper as uncommon and found from 18-55m preorbital (19-22% HL) than M. spilo- depth in the SE Arm. I have found only stichus. The oblique stripe is generally two specimens in trawls, one from 19- more continuous than in M. spilo- 26m near Namiasi and another from stichus. Mylochromis formosus is less lat- Mpemba at 44m. Konings (1990a, erally compressed, has an oblique 1995b) records the species from sandy stripe extending to the base of caudal areas around Senga Bay, but thinks that fin and usually has a prominent black it is found further north. lachrymal stripe. COMMERCIAL IMPORTANCE: Probably COLOUR: Females and immature occasionally caught in small numbers males yellowish, paler ventrally. The by trawls and seines. prominent dark oblique bar is continu- NOTES: Eccles & Trewavas (1989) ous anteriorly and generally ends just placed this species in the genus Sciaeno- at the anterior of the caudal peduncle. chromis, but Konings (1993a) has re- Dorsal and caudal fins faintly spotted. vised that genus to exclude the species Ripe males (illustrated by Baasch, with a continuous or broken oblique 1991) have a blue head and dorsum stripe, which he has placed in Mylo- with numerous yellow spots on the chromis. Eccles & Trewavas considered flanks. The belly is yellow. There is a that M. gracilis and M. spilostichus could black stripe on the preorbital bone and be distinguished on the ratio of inter- the chin and chest are black. Caudal orbital width to head length, but I did blue with yellow spots and stripes. not find this ratio to be diagnostic, al- Dorsal fin white with yellow spots though the mean values differed. with white border, yellow tips to the lappets, and a black submarginal band Mylochromis spilostichus on the anterior half. Anal fin black with (Trewavas) broad white margin and numerous colour photo on page 167 small white egg-spots. Pelvics black with white leading edge. DISTINGUISHING FEATURES: An elongate MAXIMUM SIZE: 25cm TL (Baasch, 1991; species with a prominent oblique Konings, 1995b) stripe, which is clearly broken into a DIET: Konings (1990a) reports that it series of spots. Along with M. gracilis, is a predator of small cichlids and in- this species is distinguished from other vertebrates and (1995b) has observed elongate oblique-striped species by the individuals attempting to extract Ps. small rounded head and large eye. It

146 has a larger eye (25-30% HL) and shal- (1995b) has observed the species over lower preorbital (11-18% HL) than M. sand at Makokola Reef at 35m depth. gracilis. The premaxillary pedicel is COMMERCIAL IMPORTANCE: Caught in more prominent than in other oblique- small numbers by trawlers. striped species. NOTES: See under M. gracilis. For- COLOUR: Females and immature merly placed in the genus Sciaeno- males brownish, paler ventrally. The chromis, but assigned to Mylochromis by prominent dark oblique bar is com- Konings (1993a). Baasch (1995) consid- prised of a series of 7-8 spots and gen- ers that it may be more appropriate to erally crossed by dark vertical bars consider it a member of the genus Stig- which are conspicuous dorsally, but matochromis, because of its blotchy pat- inconspicuous or absent ventrally. tern. He also gives an account of a simi- There is a dark spot at the posterior end lar species under the rather awkward of the caudal peduncle. Dorsal and name of Stigmatochromis 'spilostichus caudal fins faintly spotted. Eccles & type'. This has a longer, straighter Trewavas (1989) describe ripe males as snout than M. spilostichus and has a having golden flanks with each scale strongly projecting lower jaw. The ob- edged with blue, brownish dorsally lique stripe is made up of several elon- with gold and blue iridescence, white gated blotches. Male breeding colour ventrally. Lips and upper part of head and reproductive behaviour are very iridescent blue, chest and branchio- similar to those of M. gracilis. Konings stegal membranes yellow. Dorsal and (1990a) states that the 'Spilostichus caudal fins with dark yellow spots. type' is known only from aquarium Anal dark yellow with broad dark bar, specimens, but presents photographs no egg spots. Dorsal fin margin white labelled as this species from Likoma with orange tips and a dark submar- and Chisumulu Islands. These appear ginal band. Pelvics orange with a dark to be more heavily built than Baasch's bar and white leading edge. Faint ver- specimens and may represent another tical bars and oblique stripe on upper species. In the Monkey Bay field col- part of flank. lection, I found four specimens col- MAXIMUM SIZE: 25cm TL (Konings, lected from Chisi near Nkhata Bay by 1995b). the JFRO in 1959. These have a row of DIET: Small fishes (Konings, 1995b). vertically elongated spots and are up REPRODUCTION: Males defend small to 21cm TL. They are labelled Haplo- spawning pits on sand near rocks chromis pholidophorus, but appear to be (Konings, 1995b). a further species of the M. spilostichus DISTRIBUTION AND ABUNDANCE: Type group. from Monkey Bay area. Eccles & Tre- wavas record the species as common throughout the SE Arm at depths of 18- Aristochromis christyi Trewavas 70m. In the 1992 survey, it was re- colour photo on page 168 corded over the same depth range. Also collected by the author from 40- DISTINGUISHING FEATURES: Distin- 70m depth off Domira Bay. Konings guished from all other oblique-striped

147 species by its beaked premaxillae. from other oblique-striped species by COLOUR: Females and immature their powerful deep jaws and large males brownish, with a prominent thin broad teeth. Docimodus johnstonii has dark oblique band and another very broad flattened teeth and a thin diago- slender dark band just below the base nal band. The similar D. evelynae, a of the dorsal fin. Ripe males (illustrated species of rocky coasts, has more slen- by Konings, 1990a) bright blue. Anal der teeth and a broad blotchy diago- fin orange with numerous white egg- nal band with prominent partial verti- spots. cal bars. MAXIMUM SIZE: 30cm TL (Konings, COLOUR: Females and immature 1990a). males silver-grey with a thin black ob- DIET: Konings (1990a) reports that it lique band. Male breeding colours are takes small rocky shore cichlids with a unknown. sideways strike. I have observed this MAXIMUM SIZE: 30cm TL (Konings, species hunting in the company of N. 1990a). polystigma at Cape Maclear and with DIET: Eccles & Lewis (1976) report T. macrostoma at Nkhata Bay. that it feeds on the fins of clariid cat- REPRODUCTION: Konings (1990a) re- fishes. ports that males hold territories among REPRODUCTION: Eccles & Lewis (1976) rocks and females brood their young report that ripe males were taken from in caves. a trawl catch at 18m in the SE Arm. DISTRIBUTION AND ABUNDANCE: Appar- DISTRIBUTION AND ABUNDANCE: Eccles & ently found throughout the lake (Ec- Trewavas (1989) report that the species cles & Trewavas, 1989; Konings, 1990a), is uncommon, but found throughout although nowhere common. Konings Lakes Malawi and Malombe and the reports that it is found over rocks and Upper Shire River. Konings (1990a) sand and is mostly found at depths of reports that it is found throughout the 2-10m. Not recorded in the 1992 sur- lake and illustrates specimens from vey, but the author has recorded the Senga Bay. He believes it to be confined species in experimental trawls at about to sandy and muddy areas. It was rare 20m depth in the SE Arm, as well as in the 1992 survey, being recorded in from Lake Malombe and the Upper trawls at 42 and 46m in the SE Arm. Shire River. The author did not record this fish from COMMERCIAL IMPORTANCE: Negligible. Lake Malombe or the Upper Shire Exported in small numbers as an River despite examining many catches aquarium fish and regularly bred in in 1990-92. captivity. COMMERCIAL IMPORTANCE: Negligible. NOTES: Small specimens of the simi- lar Eccles & Lewis Docimodus johnstonii Boulenger are commonly seen in shallow rocky colour photo on page 168 areas (see Konings, 1990a).

DISTINGUISHING FEATURES: The heavy- headed Docimodus are distinguished

148 Corematodus taeniatus Trewavas the SE Arm of Lake Malawi. It forms a colour photo on page 168 small proportion of the catch weight in all cases. DISTINGUISHING FEATURES: Distin- guished from all other oblique-striped species by the large flat areas of closely Caprichromis liemi packed jaw teeth. C. shiranus is mor- (McKaye & MacKenzie) phologically almost identical, but is easily distinguished by its vertically- DISTINGUISHING FEATURES: A deep-bod- barred pattern. ied laterally-compressed species with COLOUR: Females and immature a thin black oblique stripe. It is distin- males yellowish with a thin black ob- guished from most other similarly lique band. Ripe males (illustrated by marked species by its long steeply-an- Spreinat, 1992b) are bright blue with gled jaws and thick lips. C. orthognathus numerous orange spots on the flanks, has shorter, more steeply-angled jaws. and orange spots and stripes on the COLOUR: Females and immature dorsal and caudal fins. The pelvic fins males silvery, darker dorsally with a are black, with a white leading edge. slender continuous black oblique Anal fin black with a broad orange band. Ripe male bright blue with or- margin and several irregular white ange spots in dorsal and caudal fins, egg-spots. and irregular pale streaks in the anal MAXIMUM SIZE: 20cm TL (Konings fin (Konings, 1991d, 1995b). 1990a). MAXIMUM SIZE: 23cm TL (Konings, DIET: It feeds on the scales of sand- 1990a). dwelling haplochromine cichlids (Ko- DIET: A paedophage which rams nings, 1990a). mouthbrooding female cichlids REPRODUCTION: Konings (1990a, (McKaye & MacKenzie, 1982). Konings 1995b) reports that ripe males defend (1991d, 1995b) suspects that it may also territories on the surface of rocks. feed on external parasites, such as DISTRIBUTION AND ABUNDANCE: Eccles & Argulus. Trewavas do not give a type locality or REPRODUCTION: Konings (1995b) states distribution, but report the colour of a that males form leks, build sand-cas- specimen taken at Malembo. Konings tle nest over one-metre in diameter and (1990a) states that the species has a brooding females remain in schools. lake-wide distribution, and is most DISTRIBUTION AND ABUNDANCE: Type common where small patches of rock material is from the Nankumba Penin- are near to sandy coasts. It was caught sula in the south of the lake. I have in several samples at 22-60m in the observed large individuals which re- 1992 trawl survey and also in shal- mained motionless on the substrate in lower water in Lakes Malawi and the middle of a lek of Copadichromis Malombe and the Upper Shire River. thinos near Monkey Bay. In the 1992 COMMERCIAL IMPORTANCE: Recorded trawl survey it was recorded from a occasionally from all types of seines single sample at Mpemba (south of and trawls from Lake Malombe and Boadzulu Island) at 44m depth.

149 Figure 44. Caprichromis liemi, male, South-East Arm.

1990a). Konings (1990a) reports that the spe- DIET: A paedophage which rams cies is also found at Nkhata Bay. mouthbrooding female cichlids COMMERCIAL IMPORTANCE: Negligible. (McKaye & MacKenzie, 1982). Konings NOTES: Konings (1990a) reassigned (1995b) states that it frequently attacks this species to Maravichromis (a junior female Fossorochromis rostratus. synonym of Mylochromis), but has sub- REPRODUCTION: Males build sand-cas- sequently adopted Eccles & Trewavas' tle bowers, usually alongside large classification (1995b). rocks (Konings, 1995b). DISTRIBUTION AND ABUNDANCE: Type material is from SW Arm, perhaps near Caprichromis orthognathus the Nankumba Peninsula, in the south (Trewavas) of the lake. Eccles & Trewavas (1989) colour photo on page 168 claim that it is found throughout the lake and Konings (1990a) states that it DISTINGUISHING FEATURES: A deep-bod- is also found in Lake Malombe. I have ied laterally-compressed species with recorded it from trawl catches and a thin black oblique stripe. It is distin- seines in southern Lake Malawi, but guished from most other similarly not in Lake Malombe. Eccles & Trewa- marked species by the extremely vas reported the species from trawl steeply-angled jaws and thick lips. catches between 35 and 55m in the COLOUR: Females and immature south of the lake. All the material ex- males silvery, darker dorsally with a amined by Eccles & Trewavas was slender continuous black oblique from the southern part of the lake, but band. Ripe males are blue with whit- Konings (1995b) illustrates specimens ish spots and streaks on the anal fin from Mdoka in the far north of the (Konings, 1995b). Malawian shore. MAXIMUM SIZE: 25cm TL (Konings, COMMERCIAL IMPORTANCE: Negligible.

150 NOTES: Konings (1990a) reassigned this species to Maravichromis (a junior synonym of Mylochromis), but has sub- sequently (1995b) adopted Eccles & Trewavas's classification with reserva- tions. Konings (1990a) illustrates and discusses a fish he calls Protomelas sp. 'paedophage' which appears to be identical to C. orthognathus except that it has two horizontal stripes instead of an oblique stripe. He has observed this species in the north and south of the lake and illustrates specimens from Mdoka and Chisumulu, including a blue ripe male. The diet and maximum size are identical to those of C. ortho- gnathus, but the horizontally-striped form attacks mouthbrooding females from above while C. orthognathus at- tacks from below. McKaye & Mac- Kenzie note that C. orthognathus can change colour, adopting an oblique stripe when attacking oblique-striped species and otherwise losing the stripe entirely. It is not inconceivable that it might be able to adopt a horizontally- striped pattern and thus Protomelas sp. 'paedophage' may be conspecific with C. orthognathus. Konings (pers. comm.) has preserved specimens of both and is certain that they are different spe- cies.

151 Chapter 15 Otopharynx and other spotted species

There are a number of species of Tramitichromis intermedius widely differing body form which (Trewavas) share a common pattern of three dark colour photo on page 168 spots. In some cases the spots are ar- ranged as if they were the remains of a DISTINGUISHING FEATURES: Distin- horizontal band, in others they are ap- guished from other short-snouted spe- pear to be the rudiments of an oblique cies of the Lethrinops group by presence stripe. Those species with small, highly of dark suprapectoral and supra-anal protrusible mouths have already been spots. Differs from most other 3-spot- discussed under Copadichromis. All spe- ted species in the shape of the first cies belonging to the genera Oto- lower gillraker (broad and flat) and the pharynx, Trematocranus, Ctenopharynx, Lethrinops-type dentition. It has a fewer Hemitilapia, Naevochromis, Stigmato- gillrakers (8-10) than all Ctenopharynx, chromis, and Exochochromis have this Trematocranus labifer, and all Otopharynx pattern. The latter three genera are except O. tetrastigma, O. selenurus and morphologically very distinctive. O. argyrosoma. O. argyrosoma has a less moorii also frequently exhib- deep body, O. selenurus is generally not its a three-spotted pattern, although spotted, O. tetrastigma has a more the mature fish of both sexes are nor- pointed snout and longer jaws. Trema- mally a uniform blue. A number of tocranus microstoma has a longer snout Lethrinops species frequently show and smaller mouth than T. intermedius. faint spots in addition to more promi- T. placodon is very similar morphologi- nent vertical bars these are mostly con- cally, but has much more larger, molari- sidered under 'Lethrinops and allied form pharyngeal teeth. genera'. The jaw dentition, pharyngeal COLOUR: Live coloration dark grey bone morphology, and male breeding dorsally, paler ventrally, with two dark colours of Tramitichromis intermedius blotches on flanks. Ripe males gener- are clearly most similar to other mem- ally dark blue with orange spots on bers of this genus, but the body pat- dorsal and anal fins. Anal fin orange- tern is very similar to that of Tremato- brown with numerous large yellow- cranus, so it is most likely to be con- white spots and stripes. Pelvics orange fused in the field with species of this brown with white anterior edge. Dor- genus, and is thus considered here. sal fin margin white with red tips to lappets. Branchiostegal membrane and chest red. Dark flank spot sometimes visible.

152 MAXIMUM SIZE: 15cm TL. larged sensory pits. Both of these fea- DIET: Konings (1995b) reports that it tures seem to be prone to convergent feeds on insect larvae and other soft- evolution. Subsequently Konings bodied invertebrates. (1995b) treated it under Tramitichromis, REPRODUCTION: Konings (1995b) re- with reservations. ports that males defend small pits built in the sand. DISTRIBUTION AND ABUNDANCE: The Trematocranus placodon (Regan) types are from the southern part of the lake. Jackson (1961) was not familiar DISTINGUISHING FEATURES: Differs from with the species. Konings (1995b) re- species of the genera Stigmatochromis ports a lake-wide distribution and and Exochochromis in the deeper body (1990a) illustrates specimens from and smaller jaws. Differs from Cteno- Senga Bay and from Ngara in the north pharynx, other Trematocranus, and of the lake, but they may not be con- Otopharynx species, except for O. specific. A shallow water species, in- tetrastigma in the lower gillraker count frequently caught at 20-30m depth (7-9). Deeper body, smaller jaws, and south of Monkey Bay. In the 1992 trawl less rounded head profile than O. survey they were taken in large num- tetrastigma and Hemitilapia. Lacks the bers at a single trawl station on the east wide flat first gillraker of T. intermedius. side of the central part of the SE Arm, Differs from most similar species in its at 20m. They were also taken in other large, heavy molariform lower pharyn- shallow water trawls in the same area geal teeth. and in seines near the mouth of Mid- COLOUR: Live coloration dark grey dle Shire dorsally, paler ventrally, with two or COMMERCIAL IMPORTANCE: Recorded in three dark blotches on the flanks. Ripe seine catches in the SE Arm. males (Konings, 1990a) are turquoise NOTES: Konings (1990a) illustrates blue, with red spots on flank scales. specimens from Ngara and Senga Bay. Pelvic and anal fins black. Numerous The ripe males have a turquoise sheen small egg-spots. and a white chin, rather than the dark MAXIMUM SIZE: 23cm TL (Konings, blue sheen and red chin of the south- 1990a). ern specimens. They also have a rela- DIET: Snails, bivalves and algal ma- tively large head. It is not clear if these terial were found in the stomachs of represent a different species or if they two individuals of 9-10cm SL (Grant are just small fishes at an earlier stage & Turner, unpublished). It feeds on of sexual maturity. Trewavas (1931) thin-shelled snails which are crushed assigned this species to Lethrinops on by the pharyngeal apparatus. the basis of its dentition. Eccles & REPRODUCTION: Konings (1990a) re- Trewavas (1989) removed it to Tramiti- ports that from July to September, chromis on account of its pharyngeal males defend large crater nests (70cm bone structure. Konings (1990a) re-as- diameter), but suggests that they might signed this species to Trematocranus, on use pits abandoned by other species. the basis of its colour pattern and en- DISTRIBUTION AND ABUNDANCE: Konings

153 Figure 45. Trematocranus placodon, female, South-East Arm.

(1990a) reports a lake-wide distribu- in controlling snails which act as vec- tion and an average depth preference tors for human schistosomiasis. The of 5m. Specimens examined by Eccles superficially similar Trematocranus & Trewavas came from northern and labifer (Trewavas) was not positively southern ends of the lake. They re- identified. It has 11-13 lower gillrakers ported its occurrence in Lake Malombe and lacks the molariform pharyngeal and the Shire River as far as the Majete dentition of T. placodon. Rapids, but only from a single trawl at 18m off Malembo. Jackson (1961), Ec- cles & Trewavas and Konings (1990a) Ctenopharynx nitidus (Trewavas) report that the species is associated colour photo on page 169 with Vallisneria beds. In the 1992 trawl survey it was found in most samples DISTINGUISHING FEATURES: Differs from from depths shallower than 20m and most other spotted species in the rela- was often abundant. I also found it in tively deep body with flat lower pro- artisanal catches from the Upper and file, and large ventrally placed mouth. Middle Shire and Lake Malombe and Suprapectoral and supra-anal spots frequently observed it underwater on large and elongated. Lower number of Chembe Beach on the Nankumba Pe- gillrakers than C. intermedius. ninsula. COLOUR: Live coloration dark grey COMMERCIAL IMPORTANCE: Small dorsally, paler ventrally with bronze catches in seines, gillnets and trawls. cast, with two dark blotches on flanks Frequently exported for the aquarium and a smaller one on caudal peduncle. trade. Throat and chest yellowish. Ripe males NOTES: Not reliably distinguished dark blue, yellow ventrally. Unpaired from T. labifer in the quantitative sur- fins dark with faint spots. Dorsal fin vey. Chiotha et al. (1991a,b) have lappets white. Anal fin dark with 6 shown that this species may be useful small whitish egg-spots. A male illus-

154 trated by Konings (1995b) has 7 large and with more lower gillrakers (32-41) yellow egg-spots. than C. nitidus. MAXIMUM SIZE: 13cm TL. COLOUR: Live coloration dark grey DIET: According to Eccles & Trewa- dorsally, silvery grey on flanks, white vas, it feeds on benthic invertebrates. ventrally, with two dark blotches on The stomach of a single 9cm SL speci- flanks and a smaller one on caudal men contained mostly sand and detri- peduncle. Ripe males generally golden tus, with some copepods and yellow, chin and chest orange. Head macrophyte material (Grant & Turner, and nape blue. Unpaired fins dark with unpubl.). Konings (1995b) has ob- faint orange spots. Dorsal fin lappets served the species filtering sediment. white or yellow. Anal fin dark with 6 REPRODUCTION: Konings (1995b) re- or more whitish egg-spots. ports that males are territorial but do MAXIMUM SIZE: 22cm TL (Konings not build bowers. 1995b). DISTRIBUTION AND ABUNDANCE: Some of DIET: The large number of long gill- the type material was collected at rakers suggests a diet of small crusta- Chilumba and Vua at the north end of ceans or algae, either obtained from the the lake (Eccles & Trewavas, 1989). Ec- water column or more probably (like cles & Trewavas report it from trawl the similar Ct. pictus) from the sedi- hauls between 8 and 35m, and it was ment. Two specimens examined indi- most abundant in the shallower water. cated dietary flexibility. A 7cm fish Konings (1995b) reports a lake-wide from Namiasi contained mostly distribution, and a preference for copepods and cladocera, with some sandy areas at a depth of 15m or more. algae material. An 11cm individual In the 1992 survey taken at depths of from Chilinda contained mostly large 18-65m, but most common in shal- insect nymphs (dragonfly?) and sand lower water on the east coast of the SE (Grant & Turner, unpubl.). Konings Arm. Also taken at 31-37m in Domira (1990a, 1995b) states that the species Bay. feeds by scooping small invertebrates COMMERCIAL IMPORTANCE: Occasion- from the sand and mud, but has ob- ally taken by midwater trawl. Probably served schools feeding on plankton 1- contributes to the pair trawl catch in 2m above the bottom (Konings, 1990a). areas north of Boadzulu Island. REPRODUCTION: Konings (1990a, 1995b) reports that males construct nests on the top of rocks or on nearby Ctenopharynx intermedius (Günther) sand. Robinson (1995) has observed a colour photo on page 169 breeding arena at 10m depth off Thumbi East Island. Territorial males DISTINGUISHING FEATURES: Differs from defended the tops of large (1-3m) most other spotted species in the rela- rocks. tively deep body with flat lower pro- DISTRIBUTION AND ABUNDANCE: Type is file, and large ventrally-placed mouth. badly preserved skin from an un- Suprapectoral and supra-anal spots known location. Other material exam- large and elongated. Deeper-bodied ined by Eccles & Trewavas from the SE

155 and SW Arms, and they also report it ventrally. Three dark blotches on flanks as far north as Nkhotakota. They re- and caudal peduncle. Ripe males ported it from trawl samples at 18m. bright metallic turquoise blue. Orange In 1992, it was frequently taken in trawl spots on flank scales and dorsal and samples throughout the SE Arm at caudal fin. Dorsal fin with white lap- depths of 18-60m. Konings (1990a) re- pets and red tips. Anal fin with whit- ports the species from Chilumba, ish spots and streaks. Faint vertical Usisya and Ngara in the north, as well bars often visible and there is usually in the south. a black stripe on the preorbital bone. COMMERCIAL IMPORTANCE: Comprised MAXIMUM SIZE: 20cm TL (Konings, around 1% of midwater trawl and deep 1990a). water bottom trawl catch. Also a small DIET: Principally scrapes algae from component of seine and pair trawl the surface of leaves of aquatic plants, catches. generally Vallisneria and Potamogeton. NOTES: There is a possibility that Also opportunistically takes plankton, there are two similar species, which do invertebrates, and small fish (McKaye not differ in colour, but have slightly & Marsh, 1983). different body proportions. The males REPRODUCTION: Konings (1990a) re- of the shallow water form have a larger ports that males clear a circular area of head and flatter ventral profile. This around 15cm diameter among weeds. form is illustrated by Konings (1990a, Males do not form arenas. Females 1995b), was collected by Robinson have not been reported to guard free- (1995) and more closely resembles the swimming fry. illustration in Eccles & Trewavas DISTRIBUTION AND ABUNDANCE: Jackson (1989). (1961) reports this species to be an in- habitant of the zone intermediate be- tween rocks and sand. Eccles & Trewa- Hemitilapia oxyrhynchus vas (1989) and Konings (1990a) record (Boulenger) the species as common all round the colour photo on page 169 lake in shallow vegetated areas. I have observed this species in large numbers DISTINGUISHING FEATURES: Differs from in shallow vegetated areas all around most other spotted species in the rela- the southern part of the lake and have tively deep body, terminal mouth, collected it from artisanal seine fisher- pointed snout and long jaws. The teeth ies in Lake Malombe and the SE Arm are characteristic, being unicuspid, of Lake Malawi, as well as experimen- long, and inclined forwards. The ends tal fyke nets and gillnets in the Upper of the closely-packed teeth form a con- Shire River. It was uncommon in Lake tinuous flat sharp blade which is used Malombe. It was taken from a single to scrape algae from the leaves of sample at 20m off Namiasi in the 1992 plants. trawl survey, and also seen in other COLOUR: Females and immatures shallow water hauls in the same area. generally with a yellowish-brown cast. COMMERCIAL IMPORTANCE: Contributes Dark grey-brown dorsally, silvery significantly to seine net catches in

156 some areas, and occasionally taken by MAXIMUM SIZE: 14cm TL (Konings, shallow water pair trawls. Occasion- 1990a). ally seen in the ornamental fish trade. REPRODUCTION: Seven ripe females NOTES: Feeds from the surface of veg- taken in Lake Malombe, ranging in size etation by turning onto its side, mak- from 75-97mm TL, contained 61-108 ing it highly conspicuous from the sur- eggs (Turner & Mwanyama, unpubl.). face. Probably for this reason, it moves Konings (1990a, 1995b) reports that into the shallowest part of its range males clear a circle of 10cm diameter (depths of less than 2m) in late after- among vegetation. noon, when it may be less visible to DIET: 26 specimens examined in Lake bird predators (Turner & Robinson, Malombe (Turner & Mwanyama, un- unpubl. data collected at Chembe publ.). Diet dominated by snails (18), Beach, Cape Maclear). In aquaria, the chironomids (4), or the copepod Meso- species forms dominance hierarchies cyclops (3). Other crustaceans, dragon- and dominant fish lose the typical fly, and chaoborid larvae, algae, nema- blotched pattern and develop vertical todes, and fish scales also taken. Ko- bars. This behaviour does not appear nings (1990a) reports a diet of insect to occur in the wild (Robinson & larvae and crustacea. Turner, 1990). DISTRIBUTION AND ABUNDANCE: An abundant species in Lake Malombe, the Upper Shire River, and shallow Otopharynx tetrastigma (Günther) swampy areas of Lake Malawi, such colour photo on page 169 as Monkey Bay. Material examined by Eccles & Trewavas came from the ex- DISTINGUISHING FEATURES: Differs from treme north as well as the far south of most other spotted species in the rela- the lake. Jackson (1961) reported that tively deep body, and pointed snout. this species was most common in the It has a more slender body than Hemi- south of the lake, probably on sandy tilapia and O. tetraspilus, and is distin- shores. Tweddle et al. (1979) found it guished from juvenile O. auromargina- in the Middle Shire River. Eccles & tus by the smaller number of gillrakers Trewavas reported that it was found (9-11). throughout Lakes Malawi and Ma- COLOUR: Live coloration sandy dor- lombe, which is repeated by Konings sally, paler ventrally, with three dark (1990a), who illustrates specimens blotches on flanks and caudal pedun- from Thumbi East Island. Konings be- cle. Dorsal and caudal fins faintly spot- lieves that it is confined to shallow veg- ted. Ripe males dark blue, with red etated areas from 1 to 7m deep. It was spots. Lappets white, with red or yel- not recorded in trawls by Eccles & low tips. Anal fin dull orange, with 6- Trewavas, nor by the present author. 10 white egg-spots, and red-yellow COMMERCIAL IMPORTANCE: Contributes margin. Konings (1990a, 1995b) also significantly to the seine net catches in describes male breeding colour as dark Lake Malombe, where an estimated blue, but illustrates a bright turquoise 375 tonnes were landed in 1990. The fish. average weight of 315 fish I examined

157 from the artisanal seine catch was 7g males not known for certain, although per fish, so the annual landing must Konings (1995b) illustrates a male la- be approximately 50 million fish. It is belled as 'Otopharynx cf. tetraspilus' also caught by seines in the SE Arm which has orange flanks and a blue and probably elsewhere in suitable head and nape. The dorsal fin margin habitats. in blue-white and the anal fin margin NOTES: Specimens collected at Mon- yellow. key Bay were more slender with shal- MAXIMUM SIZE: 16cm TL (Eccles & lower cheeks and less steep head pro- Trewavas). files than those from Lake Malombe. DIET: Algae, macrophytes, small It is not certain they are conspecific. crustacea (Eccles & Trewavas). They were initially identified by the DISTRIBUTION AND ABUNDANCE: Uncom- author as O. heterodon, but have fewer mon in trawls, it is probably confined dorsal fin spines than that species. The to shallow water. The type is from the identity of O. heterodon has not been eastern shore of the SE Arm. Eccles & clarified by field workers. Jackson Trewavas examined 97 specimens in- (1961) reports that this species builds cluding 28 types, all from the SE and large nests (50cm diameter) on the SW Arms. There is a record in the Shire sand, while Konings (1990a) is of the River from Tweddle et al. (1979). Al- opinion that it is a rocky shore species though Eccles & Trewavas state it is which does not build a nest. Konings widespread in the south in vegetated reports that a related species found at areas, it is otherwise unknown except Likoma builds large nests. O. 'heter- for two specimens I have collected, one odon Nankumba' from Cape Maclear from a trawl haul from between 5 and is a completely different species which 18m depth from Namiasi to the mouth superficially resembles Ct. pictus and of the Shire River, and a possible juve- is confined to rocky shores. nile collected from a seine haul in Mon- key Bay. COMMERCIAL IMPORTANCE: Minor. Otopharynx tetraspilus (Trewavas) NOTES: Konings (1995b) suggests that colour photo on page 169 the fish he previously named Oto- pharynx 'yellow fin mloto' (Konings, DISTINGUISHING FEATURES: Differs from 1990a) may be conspecific with O. most other spotted species in the rela- tetraspilus. tively deep body, steeply sloping head profile, small mouth and pointed snout. Lacks the long snout and jaws Trematocranus microstoma of Hemitilapia and is deeper bodied (Trewavas) than O. tetrastigma. Distinguished from colour photo on page 169 juvenile O. auromarginatus by the smaller number of gillrakers (9-11). DISTINGUISHING FEATURES: Differs from COLOUR: Females and non-breeding most other spotted species in the deep males silvery grey, darker dorsally, body, pointed snout, small mouth and with three small rounded spots. Ripe expanded sensory pits on underside of

158 head. The spots are often faint or in- gill net catches in suitable areas. visible in large females, as well as breeding males. COLOUR: Live coloration generally Lethrinops 'Domira blue' brownish. Juveniles with two large colour photo on page 170 blotches on flanks. Adult males dark blue with faint vertical bars. Dorsal fin DISTINGUISHING FEATURES: A small rela- with white lappets and red tips. Anal tively deep-bodied species, with faint fin with a large number of white egg- vertical bars and 1-2 dark blotches on spots, and red ventral margin. flanks and caudal peduncle. It has MAXIMUM SIZE: 25cm TL (Eccles & fewer dorsal fin spines (15-16) than O. Trewavas). heterodon and differs from that species DIET: The stomach of a 15cm SL indi- and other Otopharynx in its 'Lethri- vidual contained small whole gastro- nops-type' dentition and flattened pods as well as pieces of shell and flesh lower jaw. It differs from Lethrinops from larger snails (Grant & Turner, 'yellow-chin' in male breeding colours, unpubl.). Konings (1990a) reports that and lower number of gillrakers (10-12 they feed on small invertebrates which ceratobranchials). It is distinguished are detected by the lateral line system. from T. brevirostris and T. 'brevirostris Konings claims that they hover, al- deep' by its dentition, breeding col- though not close to the bottom, as do ours, and broader vertical bars. Aulonocara species. COLOUR: Females and immature REPRODUCTION: Konings (1995b) has males silvery, darker grey dorsally, recorded that males build craters on with 7-8 dark bars under the dorsal fin. muddy bottoms. Males in breeding Large dark irregular suprapectoral colour were found during November blotch, and a smaller dark spot on the and December. I have found a ripe caudal end of the caudal peduncle. male in July. Supra-anal spot apparently absent. DISTRIBUTION AND ABUNDANCE: Pre- Ripe males silvery blue, with dark blue served material examined by Eccles & snout and dorsal part of head. Trewavas (1989) mostly from the north MAXIMUM SIZE: 10.5 cm TL. of the lake, but one specimen was from DISTRIBUTION AND ABUNDANCE: Posi- the south. They did not record it in tively identified only from 55m depth trawl surveys and the species did not off Domira Bay. Not known from the seem to be known by Jackson (1961). SE Arm. Konings (1990a) reports a lakewide COMMERCIAL IMPORTANCE: Minor. distribution in shallow muddy bays NOTES: In body shape and colour pat- between 4 and 20m depth. He illus- tern, this species resembles Otopharynx trates specimens from Senga Bay. In the more than Lethrinops spp., but the den- 1992 trawl survey it was generally tition is characteristic 'Lethrinops'-type. trawled in shallow water (24-44m), and Few specimens have been collected was rare south of Boadzulu Island. and it is possible that there are two COMMERCIAL IMPORTANCE: Probably species. Males with egg-spots have a contributes a little to the seine net and straighter ventral profile and more

159 pointed snout than those with a dark Island its depth range was 18-70m. It anal fin margin. is less numerous south of Boadzulu Is., where it was recorded only at the most northerly station (Mpemba), where it Otopharynx 'argyrosoma deep' was quite common at 28-34m depth. colour photo on page 170 COMMERCIAL IMPORTANCE: Despite its abundance in experimental surveys of DISTINGUISHING FEATURES: 7-8 thin ver- the main trawling grounds, this spe- tical bars under the dorsal fin and cies did not seem to be a major com- prominent suprapectoral spot. 11-14 ponent of the commercial catch in the gillrakers, teeth in 4 rows arranged in area, being estimated at 0.3% of the 'Haplochromis' manner. More elongate haplochromine component of the than most spotted species, but deeper semipelagic trawl catch. bodied than O. decorus and O. 'pro- NOTES: It is possible that it may have ductus'. Preorbital bone shallower been mis-identified during the survey (depth <22% head length) than that of of the commercial catch. This species O. selenurus. Other O. argyrosoma spe- was identified as O. argyrosoma using cies are smaller, more slender and lack the key of Eccles & Trewavas. The type prominent spots and stripes. Lethrinops of O. argyrosoma (which I have exam- micrentodon has more gillrakers and L. ined and measured) is a small colour- 'yellow chin' is less laterally com- less juvenile from an unknown loca- pressed and has a 'Lethrinops'-type tion. I have been unable to determine tooth arcade. which population is conspecific with COLOUR: Females and immatures sil- the type, which is much more slender very with 7-8 thin vertical bars under than large specimens of this species, the dorsal fin and 1-2 on the caudal but has indistinguishable meristics. It peduncle. A dark spot lying on the is possible that the material examined upper lateral line unites the 3rd and by Eccles & Trewavas may represent 4th bars. Some specimens (especially several species. FAO (1976) reported the more slender ones) have small that Haplochromis argyrosoma was one supra-anal and caudal spots. Live male of the most common species in the colours silvery with a generally bluish trawl catches in the SE Arm. They gave cast. Vertical bars and spots present, a maximum length of 17cm TL, which but faint. almost certainly means that this spe- MAXIMUM SIZE: 16cm TL. cies (and perhaps several others) was DISTRIBUTION AND ABUNDANCE: Abun- that recorded in their survey. FAO gave dant in the SE Arm, particularly trawl a natural mortality rate of 1.64, and log area B (between the latitudes of Mon- (weight, g)= 2.579.log (length, mm) - key Bay and Boadzulu Island) where 3.025. it formed a larger proportion of the bio- mass than any other species during the 1992 survey. Overall, it was the second most abundant cichlid recorded, in terms of biomass. North of Boadzulu

160 Placidochromis hennydaviesae, male, off Monkey Bay. P. 'hennydaviesae III', male, off Maleri Island.

Placidochromis 'carnivore', male, South-East Arm. Placidochromis 'acuticeps', female, SE Arm.

Placidochromis 'macrognathus', male, South-East Arm.

Placidochromis johnstoni, immature, Monkey Bay. Sciaenochromis 'deep water', female, SE Arm.

161 Sc. psammophilus, male, SE Arm (Namiasi) Sc. psammophilus, female, SE Arm (Malindi)

Sciaenochromis benthicola, male, Domira Bay.

Placidochromis 'long', male, South-East Arm.

Placidochromis 'long', female, SE Arm. Sciaenochromis benthicola, female, South-East Arm.

162 Placidochromis subocularis, male, SE Arm (Namiasi)

Placidochromis subocularis, female, SE Arm (Malindi)

Placidochromis 'longimanus Chirombo', males, SE Arm (Chirombo Bay)

Corematodus shiranus, female, South-East Arm.

Placidochromis 'longimanus Namiasi', male, South-East Placidochromis 'longimanus Maleri', male, off Arm (Namiasi) Maleri Island.

163 Buccochromis nototaenia, male, South-East Arm (Chapola Shoal).

B. nototaenia, female, SE Arm (Namiasi). B. cf. oculatus, female, SE Arm (Chirombo Bay)

Buccochromis rhoadesi, juvenile, SE Arm. Buccochromis lepturus, juvenile, South-East Arm.

Buccochromis rhoadesi, male, Senga Bay (aquarium photo).

164 Taeniolethrinops laticeps, juvenile, South-East Arm. Mylochromis anaphyrmus, male, off Maleri Is- land.

Mylochromis anaphyrmus, immature male, SE Arm. Mylochromis ericotaenia, female, SE Arm.

Mylochromis cf. balteatus, male, South-East Arm.

165 Mylochromis melanonotus, female, South-East Arm.

Mylochromis 'deep', female, off Maleri Mylochromis melanotaenia, male (top) and female, SE Arm. Is.

Mylochromis 'deep', male, off Maleri Island.

Mylochromis plagiotaenia, female, SE Arm. Mylochromis 'torpedo', female, South-East Arm.

166 Champsochromis caeruleus, female, SE Arm. Ch. spilorhynchus, immature, Monkey Bay.

Mylochromis formosus, immature, off Maleri Island. Mylochromis gracilis, immature, South-East Arm.

Mylochromis spilostichus, immature, Domira Bay.

Mylochromis 'torpedo', male, South-East Arm.

167 Aristochromis christyi, female, South-East Arm. Caprichromis orthognathus, female, South-East Arm.

Corematodus taeniatus, male, South-East Arm.

Corematodus taeniatus, female, SE Arm. Docimodus johnstonii, female, South-East Arm.

Tramitichromis intermedius, male, SE Arm. Tramitichromis intermedius, female, South-East Arm.

168 Ctenopharynx nitidus, unripe male, Domira Bay.

Otopharynx tetrastigma, male (top) and female, Ctenopharynx intermedius, male, South-East Arm. Lake Malombe

Otopharynx tetraspilus, unripe male, SE Arm.

Hemitilapia oxyrhynchus, male, SE Arm. Trematocranus microstoma, male, South-East Arm.

169 Otopharynx 'argyrosoma deep', unripe male, SE Arm.

Lethrinops 'Domira blue', male (top) and female, South-East Arm. Otopharynx 'auromarginatus stripe', juvenile, SE Arm.

Otopharynx auromarginatus, male, SE Arm.

Otopharynx auromarginatus, female, SE Arm. Otopharynx decorus, female, off Maleri Island.

170 Otopharynx 'productus', female, SE Arm.

O. 'argyrosoma red', male, Lake Malombe

Otopharynx 'argyrosoma red', male (top) and female, Stigmatochromis pholidophorus, female, South-East Arm. South-East Arm.

Stigmatochromis woodi, female, South-East Arm. Stigmatochromis woodi, male, South-East Arm.

171 Otopharynx speciosus, immature, Domira Bay. Otopharynx brooksi, male (top) and female, South-East Arm.

Otopharynx speciosus, male, South-East Arm.

Stigmatochromis 'guttatus', female, SE Arm. Exochochromis anagenys, female, South-East Arm.

172 Trematocranus brevirostris, male, South-East Arm.

T. brevirostris, female, South-East Arm.

Trematocranus 'brevirostris deep', male (top) and female, Dimidiochromis kiwinge, immature, South-East Arm. South-East Arm.

Taeniochromis holotaenia, male, South-East Arm. Taeniochromis holotaenia, female, SE Arm.

173 Hemitaeniochromis urotaenia, male, SE Arm. Hemitaeniochromis spilopterus, male, South-East Arm.

Hemitaeniochromis urotaenia, female, South-East Arm.

Hemitaeniochromis 'insignis', female, South-East Arm.

174 Protomelas similis, female, South-East Arm. Protomelas kirkii, female, South-East Arm.

Protomelas triaenodon, female, South-East Arm.

Protomelas 'red dorsal', female, South-East Arm. Protomelas labridens, unripe male, SE Arm.

Chilotilapia rhoadesii, male, South-East Arm (aquarium photo).

175 Lethrinops lethrinus, male, South-East Arm. Nimbochromis livingstonii, female, South-East Arm.

Nimbochromis linni, male, Chisumulu Island.

Nimbochromis polystigma, female, SE Arm. , female, South-East Arm.

176 Otopharynx auromarginatus These authors state that it occurs all (Boulenger) round the lake, as well as the Shire colour photo on page 170 River, although not south of Liwonde. They did not record it in their trawl DISTINGUISHING FEATURES: A rounded survey, but report it from ad hoc shal- deep-bodied species with the typical low water trawls. In the 1992 trawl 3-spotted pattern of Otopharynx spp. It survey it was often taken at 20-35m has more lower gillrakers (14-18) than mainly on the east coast of the SE Arm, similar species, except Ctenopharynx south of Monkey Bay. North of Mon- nitidus, which has a larger mouth, and key Bay, it was replaced by O. 'auro- larger spots. Ct. intermedius has more marginatus stripe' although both spe- gillrakers. cies occurred sympatrically on occa- COLOUR: Live coloration generally sion. It was also occasionally seen to a grey or golden brown with three small depth of 40m and I have seen large rounded blotches on flanks and cau- shoals while diving around Mazinzi dal peduncle. Adult (ripening?) males Reef and also Chisale Beach near golden with bluish sheen, and bright Nkhata Bay. blue head. Branchiostegal membrane COMMERCIAL IMPORTANCE: Contributes bright yellow. Fins dark, spotted. Dor- to the seine net and gill net catches in sal fin with orange lappets. Anal fin suitable areas. Minor contribution to dark, with several orange egg-spots, pair- and midwater trawl catches. with orange ventral margin. Konings NOTES: The lack of records of this spe- (1995b) illustrates a territorial male, cies in Eccles & Trewavas's trawl sur- labelled as this species, which has blue vey may be due to misidentification. flanks and dark vertical bars. Specimens of O. auromarginatus and O. MAXIMUM SIZE: 25cm TL (Eccles & 'auromarginatus stripe' collected dur- Trewavas). ing the time of Eccles' trawl surveys DIET: This species can be observed in were deposited in the Monkey Bay large schools, feeding from the sedi- museum under the name 'Haplochro- ment (pers. obs.; Konings, 1995b). mis leuciscus'. Konings' (1990a) illus- Jackson (1961) says this species feeds trations of ripe males from Likoma Is- of bottom epiphytes. land are of a different species (referred REPRODUCTION: Konings (1995b) re- to as O. 'auromarginatus margrette' in ports that males construct bowers one his 1995b book) and thus his ecologi- metre in diameter on sand at depth of cal notes probably refer to that species. 10-25m. They are widely spaced and The illustrations in Konings (1995b) are do not seem to form leks. very likely of the true O. auromargina- DISTRIBUTION AND ABUNDANCE: Jackson tus. Two similar species from Chi- states that this species has been taken sumulu and Fort Maguire are also re- in shallow water between Mangochi ported. and Nkhata Bay and also at Likoma. Material examined by Eccles & Trewa- vas was from Chilumba, Karonga and Vua in the north as well as the SW Arm.

177 Otopharynx 'auromarginatus stripe' vey, it comprised about 10% of the bio- colour photo on page 170 mass of a sample at Nkope, but was otherwise rare. DISTINGUISHING FEATURES: A medium- COMMERCIAL IMPORTANCE: Minor. sized species with an elongated supra- Likely to contribute to the pair trawl pectoral spot, and an irregular stripe and seine net catches in suitable areas. on the posterior of the flank and cau- NOTES: In trawl records of the Malawi dal peduncle. Overall shape interme- Fisheries Dept., this species is confused diate between O. auromarginatus and O. with Nyassachromis leuciscus. The type decorus. More slender than O. auromar- of N. leuciscus is a small colourless ju- ginatus, which has smaller distinct venile, which has similar meristics, but spots. Fewer gillrakers (11-13) than O. is more elongate. According to the de- auromarginatus (14-18). Tail more scription by Eccles & Trewavas, N. deeply forked than in either O. auro- leuciscus may have a horizontal stripe marginatus or O. 'argyrosoma red'. Has on the posterior part of the flank, but a relatively shorter head (26-28% SL) no mention is made of a suprapectoral than O. selenurus (29-31% SL) or O. spot. 'argyrosoma red' (28-30%). Deeper bodied (34-40% SL) than O. decorus (29- 33%), which has higher meristic counts Otopharynx decorus (Trewavas) (LL Scales: 36-38 O. decorus; 34-35 O. colour photo on page 170 'auromarginatus stripe', Soft Dorsal Rays: 13-14 O. decorus; 11-12 O. 'auro- DISTINGUISHING FEATURES: Elongate. marginatus stripe'). Teeth in three Distinguished from most other spotted rows, 'haplochromis-pattern', outer species by its long body, small rounded ones unequally bicuspid. Lower pha- head, and high meristic values: 36-38 ryngeal bone with small crowded lateral line scales and 13-14 soft dorsal teeth, the three posterior teeth of inner- rays. O. 'productus' is more slender most row slightly enlarged. and has a more pointed snout. COLOUR: Sandy dorsally, silver later- COLOUR: Sandy, paler ventrally. Three ally and ventrally. Suprapectoral spot dark blotches on flanks and caudal large, elongate and rectangular. Supra- peduncle. Pelvic and anal fins bright anal spot generally drawn out into an yellow. Ripe males bright blue, silvery irregular horizontal stripe, often join- laterally. A whitish egg-spot on anal ing with the caudal spot, which is gen- fin. A specimen from Makokola Reef, erally faint. Pelvic fins bright yellow. illustrated by Konings (1990a), is simi- MAXIMUM SIZE: 18cm TL. lar, but has an orange chest. DISTRIBUTION AND ABUNDANCE: Re- MAXIMUM SIZE: 18cm TL (Konings, corded only from the trawl stations 1995b). Nkope, Mazinzi, Fowo, Chiponda, and DIET: Benthic insect larvae (Konings, Chekopa and from ad hoc trawls at 1990a). Chapola Shoal. All of these stations are DISTRIBUTION AND ABUNDANCE: Uncom- shallow (5-30m) sandy areas in the cen- mon. The types are from Vua in the far tral part of the SE Arm. In the 1992 sur- north, but Eccles & Trewavas also ex-

178 amined material from the south of the COLOUR: Sandy, paler ventrally. Three lake. They recorded it in several trawls large faint rectangular dark blotches on at 18m, and in single hauls at 35 and flanks and caudal peduncle. Pelvic and 55m. In the 1992 survey, it was fre- anal fins bright yellowish. quently taken in small numbers at 18- MAXIMUM SIZE: 17.5cm TL. 26m and also from single hauls at 34 DISTRIBUTION AND ABUNDANCE: Uncom- and 50m depth. Most records were in mon. In the 1992 survey, taken at Area B, south of Monkey Bay, and Nkope and Chilinda trawl stations at north of Boadzulu Island. Konings 18-20m. Also collected from trawls at (1990a) recorded this species from Ulande, Chekopa and near Boadzulu around the lake and states that it is Island around the Chapola Shoal from common at around 6m depth off depths of 5-50m. Usually taken in Namalenje Island. small numbers. COMMERCIAL IMPORTANCE: Occurs in COMMERCIAL IMPORTANCE: Minor, per- the semipelagic trawl catch, and it haps locally significant in seine fisher- seems likely to be locally caught in ies. seines and pair trawls. NOTES: Mylochromis 'torpedo blue' NOTES: Konings (1991e, 1995b) illus- (Konings, 1995b) may be conspecific trates a male from Senga Bay which has with O. 'productus'. It has been re- numerous small yellow egg-spots and corded from Ntekete to Narungu on a silver-grey chest. This is a deep-bod- the eastern shore of the Malawian part ied fish and may not be conspecific. He of the lake, in shallow sandy areas. A states that it feeds selectively among ripe male illustrated by Konings is blue the sediment on benthic invertebrates. with yellow spots on the flanks, dor- Konings (1991e) reports a maximum sal and caudal fins and two large yel- size of 18cm TL. Konings (pers. comm.) low egg-spots. Three dark spots are considers that this may be the true O. visible on the flanks. It has also been decorus. seen at Mdoka and a single specimen exported from Tanzania (Konings pers. comm.). Otopharynx 'productus' colour photo on page 171 Otopharynx 'argyrosoma red' DISTINGUISHING FEATURES: Elongate. colour photo on page 171 Distinguished from most other spotted species by long body and small angu- DISTINGUISHING FEATURES: A small elon- lar head. Has higher meristic values gate laterally compressed species. Dif- than most other spotted species (13 fers from other elongate species by its dorsal rays, 36 lateral line scales). Teeth deeper, more rounded body and faint in 5 rows, outer bicuspid. 11 gillrakers. midlateral blotch. Distinguished from Is more slender and has a more pointed O. 'argyrosoma deep' by its less deep snout than O. decorus. Differs from body, and the absence of vertical bars. Mylochromis 'torpedo' in lack of oblique Differs from N. 'argyrosoma blue' in stripe. coloration and less pointed snout.

179 COLOUR: Generally silvery, pale smaller sizes. sandy dorsally, usually with faint elon- DISTRIBUTION AND ABUNDANCE: A spe- gate blotch on upper part of flank. Ripe cies of shallow water in sheltered males dark with iridescent blue-green muddy areas. It is abundant in Lake sheen. An irregular patch of pinkish- Malombe. It was uncommon in 1992 red behind operculum. Dorsal fin with trawl survey and confined to waters white lappets and red tips. Anal fin of 20-28m south of Monkey Bay. I did dark with red margin. not record this species north of Mon- MAXIMUM SIZE: 12.5cm TL. key Bay and have only one uncon- DIET: Small gastropods were the firmed identification made from pre- dominant item in 12 out of 20 individu- served material from north of Boad- als collected from Lake Malombe zulu Island at the Chiponda trawl sta- (Mwanyama & Turner, unpublished). tion. Konings (1990a) illustrates speci- Copepods, chironomids, and clado- mens from Makokola Reef and Croco- cerans were also important. Quantities dile Rock which appear to be this spe- of algal remains, sand, and detritus cies, but later (1995b) reports that it is were also ingested. A specimen from also found in Senga Bay. the SE Arm of Lake Malawi contained COMMERCIAL IMPORTANCE: Abundant in gastropods, copepods, and diatoms nkatcha and kambuzi seine catches in (Grant & Turner, unpubl.). Konings Lake Malombe, where the annual catch (1990a) states that it feeds on small in- was estimated as 1,650 tonnes in 1990. vertebrates and plankton. At an average weight of 4g landed, this REPRODUCTION: Ovaries of ripe fe- translates to around 400,000,000 fish! males (n=5, SL 66-84mm) contained 18- In 1991-92, in small-meshed beach 78 eggs (Mwanyama & Turner, un- seine catches in the southern part of the publ.). Konings (1990a, 1995b) states SE Arm, it comprised a larger propor- that males construct sand-castle bow- tion of the catch by weight than any ers on open sandy areas. In Lake other haplochromine cichlid. Also Malombe (November, 1991), ripe fe- taken by pair trawls to small extent. males were found at lengths of 7-8.5cm NOTES: O. argyrosoma was described TL and ripe males from 7-9.5cm TL. It from a 60mm SL juvenile from an un- is possible that they mature at even known location in Lake Malawi. No melanic pattern is visible. The meristics of the type agree with those of the other Females Males specimens examined by Eccles & TL (cm) Immature Ripe Immature Ripe Trewavas (including the much deeper- 6.6-7.0 14 14 12 6 bodied figured specimen), as well as 7.1-7.5 17 4 13 2 O. 'argyrosoma red', N. 'argyrosoma 7.6-8.0 2 2 6 7 blue', O. 'argyrosoma deep' and prob- 8.1-8.5 2 1 4 3 ably several others. Previously-pub- 8.6-9.0 — — 1 3 9.1-9.5 — — — 2 lished field data pertaining to O. argyrosoma should be regarded as re- ferring to one or more indeterminable Table 4. Size at maturity; Otopharynx 'argyro- soma red', Lake Malombe, October 1991. species.

180 Figure 46. Mylochromis 'double spot', male, South-East Arm.

Mylochromis 'double spot' Fort Maguire. In the 1992 trawl survey, I recorded it from a single sample at DISTINGUISHING FEATURES: An elongate 28m off Chiponda, and collected other species with a small head and pointed specimens from 38m at Chiponda and snout. Less elongated and snout less 64m off Chilinda. Only occasional pointed than that of O. 'productus'. specimens found in trawl catches. Both the suprapectoral and supra-anal COMMERCIAL IMPORTANCE: Not record- spots are elongated and sometimes ed in commercial catches, perhaps due appear as double spots. The spots are to misidentification. occasionally joined into a blotchy ob- NOTES: Underwater observations in- lique band. Distinguished from M. dicate that this is a solitary species 'chekopae' by the lower number of (Konings, 1990a). I did not initially dis- gillrakers (10-11, against 13). tinguish this species from O. decorus COLOUR: Sandy, paler ventrally. Two and O. 'auromarginatus stripe'. large dark double blotches on flanks and a smaller spot on the caudal pe- duncle. Specimens from the east coast Stigmatochromis pholidophorus have bright yellow pelvic and anal fins (Trewavas) (Konings, 1991f). colour photo on page 171 MAXIMUM SIZE: 14cm TL (Konings, 1991f, 1995b). DISTINGUISHING FEATURES: A relatively DIET: Benthic invertebrates (Konings, elongate species with three prominent 1991f). It may also clean parasites from black spots and faint vertical bars. other fishes (Konings, 1995b). Head and jaws smaller than those of DISTRIBUTION AND ABUNDANCE: Konings S. woodi and S. 'guttatus'. Lacks the (1991f) reports this species from depths beaked premaxillae of Exochochromis of 5-30m near Makanjila in the south- anagenys. east and Mdoka and Chinteche in the COLOUR: Females and immature north. It is said to be abundant near males silvery, with three prominent

181 dark spots. The anteriormost spot is water (ca 90m) are probably not con- often extended from the anterior up- specific and one preserved specimen per corner. Seven faint vertical bars is now considered to be a slender beneath the dorsal fin occasionally vis- Otopharynx brooksi. Eccles & Trewavas ible. Ripe male colour not known, but (1989) describe a further species Stig- unripe males have 5-6 bright yellow matochromis pleurospilus (Trewavas) egg-spots on the anal fin, and faint red- which is known only from a single dish-orange spots on dorsal and cau- 40mm SL juvenile from the 'Lupembe dal fins and posterior part of the flanks. Sand Bank', the location of which is not Similar specimens illustrated by Ko- known with certainty, but may be in nings (1990a) have 4-5 dark spots on the extreme north of the lake, perhaps the body along the base of the dorsal in Tanzania. fin and unripe males have a white dor- sal fin lappets with red tips, generally dark unpaired and pelvic fins and a Stigmatochromis woodi (Regan) faint greenish iridescence on flanks. colour photo on page 171 Konings reports that ripe males are blue. DISTINGUISHING FEATURES: The deep MAXIMUM SIZE: 18cm TL (Konings, body and large mouth distinguish this 1990a). species from most other three-spotted DIET: Konings reports that it hovers species. The snout profile is much more about 1m from the sand, and strikes at acute than that of O. speciosus. It lacks small fish which are close to the bot- the beaked premaxillae of Exocho- tom. chromis anagenys. REPRODUCTION: Konings reports that COLOUR: Females and immature males are generally seen at around 20m males are silvery with a yellow-brown depth. cast, three prominent dark spots on the DISTRIBUTION AND ABUNDANCE: The flanks and smaller dark spots at the type is a small specimen (11cm TL) base of the dorsal fin. Ripe males gen- from Vua, at the north of the lake. Ec- erally very dark with black vertical cles & Trewavas were not aware of any bars, dark fins, numerous orange egg- other records of the species. Konings spots on the anal fin, and white dorsal (1990a) records populations from fin lappets with red tips. Konings Senga Bay and Fort Maguire and states (1995b) illustrates a pale bluish male that it is most commonly seen at 7m which has no vertical bars, but de- depth on sandy bottoms near rocks. It scribes the breeding colour as dark was not recorded in the 1992 trawl sur- blue to black. vey, but was collected by the author MAXIMUM SIZE: 25cm TL (Eccles & from trawl hauls from 15-26m in the Trewavas, 1989, Konings, 1995b) or SE Arm, but was generally rare. 30cm TL (Konings, 1990a- from an COMMERCIAL IMPORTANCE: Negligible. aquarium specimen) NOTES: This species is tentatively DIET: Recorded as piscivorous identified. A few similar specimens (Konings, 1990a), but one specimen taken from the 1992 survey from deep examined appeared to contain gastro-

182 pod remains (Grant & Turner, unpubl). Arm and on the Tanzanian coast. REPRODUCTION: I have observed a Konings (1990a, 1995b) illustrates a small lek of males and several small blue male of this species. Eccles & groups of brooding females at Mazinzi Trewavas (1989) state that the male of Reef in the SE Arm. Konings (1990a, S. woodi is blue — perhaps a reference 1995b) notes that territorial males can to S. 'tolae'. The deeper-water records be seen all year round, generally on of S. woodi reported by Eccles & Trewa- building a 1-m diameter crater nest on vas may be a result of confusion with sand near rocks at around 25-40m S. 'guttatus', although it could also be depth. The largest specimens appear that the more restricted depth range to be females (Stauffer, pers. comm.), found in the 1992 survey might be due which is very unusual for any mater- to a reduction in the abundance of the nal mouthbrooding species. species. DISTRIBUTION AND ABUNDANCE: Konings (1990a) records the species from throughout the lake. The material ex- Stigmatochromis 'guttatus' amined by Eccles & Trewavas (1989) colour photo on page 172 was from the extreme south (SE and SW Arms, Upper Shire River), or the DISTINGUISHING FEATURES: Very similar extreme north (Vua, Chilumba, Ka- to S. woodi, but much more slender. ronga, Lupembe Sand Bank). Ribbink COLOUR: Females and immature et al. (1983) recorded the species only males silvery, with three prominent from the SE Arm and the Nankumba dark spots on the flanks and smaller Peninsula. Jackson (1961) described the dark spots at the base of the dorsal fin. species as well-known and widely dis- Ripe males with prominent dark ver- tributed. The trawling records of Ec- tical bars. cles & Trewavas report the species from MAXIMUM SIZE: 16cm TL. 18-70m, but in the 1992 survey, it was DISTRIBUTION AND ABUNDANCE: In the rare, being occasionally trawled from 1992 survey, common from depths of 20-44m in the SE Arm. 50 to 100m in the SE Arm. Also re- COMMERCIAL IMPORTANCE: Taken in corded from hauls at 24 and 34m. small numbers by shallow water trawl- COMMERCIAL IMPORTANCE: Frequently ing and beach seines. Probably also seen in samples from deep-water trawl caught by hooks and gillnets. catches, but not comprising a large pro- NOTES: Eccles & Trewavas (1989) re- portion of the catch weight. port that specimens from the west side NOTES: Although numerous in the SE of the SE Arm and also those from the Arm, this species has not previously north of the lake differ in morphology been reported, unless perhaps some of and fin ray counts from the type mate- the more slender specimens of S. woodi rial. The status of these different popu- mentioned by Eccles & Trewavas are lations is unknown. Konings also re- this species. ports that a similar species S. 'tolae' occurs sympatrically with S. woodi on rocky shores at Mbenji Island, the SE

183 Otopharynx brooksi (Oliver) tern faintly visible. The dorsal fin has colour photo on page 172 a broad white margin with a yellow tip. The anal has numerous large whitish DISTINGUISHING FEATURES: A large- spots and stripes and a yellow margin. mouthed, rather heavily-built species This is a different species and in his with three large brownish spots, the later works is labelled as O. 'modestus anterior two generally elongated. Body makokola'. colour generally brownish. The mouth is more ventrally placed in O. speciosus. COLOUR: Females and immature Otopharynx speciosus (Trewavas) males brownish, whitish ventrally. 7-8 colour photo on page 172 faint vertical bars under the dorsal fin. Supra-anal and suprapectoral spots are DISTINGUISHING FEATURES: A large- both elongated and sometimes joined mouthed species which has a large to form an oblique stripe. Another dark spot on the dorsum at the ante- large spot on the caudal peduncle. rior end of the dorsal fin, in addition Maturing (?) male dark blue-grey with to large black suprapectoral, supra- dark fins. anal and caudal spots. MAXIMUM SIZE: 12.5cm SL (Eccles & COLOUR: Females and immature Trewavas), ca. 15cm TL. males brownish, whitish ventrally. DISTRIBUTION AND ABUNDANCE: The Large black caudal, supra-anal and types were taken from over rocks or suprapectoral spots. A prominent sand in the SE and SW Arms. The fourth spot at the anterior of the dor- depth distribution is given as 15-93m, sal fin. The suprapectoral spot is very but it is not clear whether specimens large and the anterior upper corner is were actually taken at 15m, or if they generally prolonged, sometimes to join were obtained from fishing gear which the fourth spot. There are sometimes were operated over the whole depth several thin, faint vertical bars occa- range. Eccles & Trewavas report that sionally intensified in the region be- the species was regularly trawled from tween the spots. Ripe males are bril- 40-50 fathoms (approx. 80-100m) in the liantly coloured. The flanks are whit- SE and SW Arms. Occasionally taken ish yellow, with a bright orange spot by the 1992 trawl survey at 60-102m, on each scale. Dorsal and caudal fins sometimes in substantial numbers, in with large bright orange spots. Dorsal the northern part of the SE Arm. There with white margin, tipped with yellow. are no records of the species outside of Anal fin dark with several large whit- the southern arms of the lake. ish egg-spots. Head bright blue COMMERCIAL IMPORTANCE: Negligible. dorsally, chin and chest bright yellow. Very occasionally in deep water trawls Lips orange-yellow. and probably also in hook and line fish- MAXIMUM SIZE: 25cm TL. eries. DIET: A predator (Eccles & Trewavas). NOTES: Konings (1990a) illustrates a DISTRIBUTION AND ABUNDANCE: Types ripe male which is bright blue with the from Vua and Monkey Bay. Eccles & spots and stripes of the melanin pat- Trewavas report it as widespread and

184 common between 35 and 55m, and The male does not construct a nest. The occasionally taken as shallow as 18m 16cm female incubated 98 fry for 3 or as deep as 70m. Also frequently re- weeks and continued to guard them corded in the 1992 survey of the SE after first release. Arm, where it was taken as deep as DISTRIBUTION AND ABUNDANCE: Konings 100m, and common between 40-70m. (1990a) records the species as common Collected by the author at 37-60m on rocky shores from 5-50m through- depth off Domira Bay. out the lake. It is rarely observed over COMMERCIAL IMPORTANCE: A minor sand (Konings, 1995b). It is rare in component of deep-water trawl trawl catches, but it was recorded from catches. Eccles & Trewavas (1989) re- a sample near Domwe Island in the port that the species is frequently taken 1992 trawl survey, and another trawl by hook and line. Apparently not ex- from near Boadzulu Island. The type ported as an aquarium fish, but the specimens are from rocky shores of bright male coloration suggests that it Thumbi West Island. would be a desirable specimen fish for COMMERCIAL IMPORTANCE: Negligible. the larger tank if it could be obtained NOTES: Konings (1995b) illustrates in shallow water. specimens with a yellow background colour. They have been found at Chi- sumulu Island and on the Tanzanian Exochochromis anagenys (Oliver) coast. Eccles & Trewavas (1989) report colour photo on page 172 that the species was found at all depths from 10-125m and over sandy areas as DISTINGUISHING FEATURES: A relatively well as near rocky shores. They report elongate, laterally compressed species that the species was known in the trawl with three prominent black spots. The surveys of the 1960s-70s as Haplochro- beaked premaxillae are unique among mis 'guttatus'. I have examined the spotted species. voucher specimens of this survey at the COLOUR: Females and immature Monkey Bay Fisheries Station, and males silvery-grey, with three promi- while one specimen was certainly E. nent dark spots, and smaller fainter anagenys, the great majority were the spots at the base of the dorsal fin. Male species I have discussed as Stigmato- breeding colour generally blue or sil- chromis 'guttatus'. As the latter species very blue. was frequently taken in deep water MAXIMUM SIZE: 30cm TL (Konings, and over sand in the 1992 survey, it 1990a). seems likely that Eccles & Trewavas DIET: Konings (1990a) states that the have confused the two. These authors species feeds on small rocky shore also speculate that the species may be cichlids (mbuna), which it catches with partially pelagic and that the deep a sideways strike. It lurks about 1m water records may be specimens taken from the bottom and attacks benthic as the trawl was being brought to the prey from above. surface. This is founded on the obser- REPRODUCTION: Baasch (1994) de- vation that it is sometimes caught by scribes reproduction in the aquarium. chirimila nets. Given Konings' obser-

185 Figure 47. Naevochromis chrysogaster. Katale Island, Chilumba. vations of its feeding behaviour, it merous large bright yellow egg-spots. seems most unlikely to be a truly MAXIMUM SIZE: 23cm TL (Konings, pelagic species. I have never seen it in 1990a). semipelagic trawl catches in the SE DIET: Konings (1990a) suggests that Arm, and it was not recorded from the it may be paedophage. pelagic zone by the ODA/SADC REPRODUCTION: Konings (1995b) has Pelagic Fish Project. Many demersal observed a male building a sand-cas- species found on rocky shores are oc- tle on top of pile of rubble, under an casionally taken by the chirimila. overhanging rock. DISTRIBUTION AND ABUNDANCE: Appar- ently found throughout the lake (Ec- Naevochromis chrysogaster cles & Trewavas, 1989; Konings, 1990a), (Trewavas) although nowhere common. Konings states that it is mostly found on areas DISTINGUISHING FEATURES: Distin- of rocks or mixed rock and sand. guished from all other spotted species Found in a single sample of the 1992 by the deep, steeply-angled lower jaw. survey at 92m off Monkey Bay. I have COLOUR: Females and immature also observed the species in a seine net males silvery-grey, with three promi- haul from shallow water in same area. nent dark spots, and faint vertical bars. Type specimens from SW Arm and Konings (1995b) reports that in some Karonga. populations the midlateral blotch is COMMERCIAL IMPORTANCE: Negligible. absent, while in others the spots are Occasionally exported as an aquarium joined to form a stripe. Ripe males (il- fish, most specimens being caught at lustrated by Konings, 1995b) blue with Chisumulu Island (Konings, 1990a). orange spots on the flanks and the dor- sal and caudal fins. Anal fin with nu-

186 Figure 48. Cyrtocara moorii. Aquarium photo.

Cyrtocara moorii Boulenger the south of the lake, I have observed numerous shoals of both Taeniolethri- DISTINGUISHING FEATURES: Spotted nops and F. rostratus, sometimes feed- when young, but uniformly dark blue ing together, and all C. moorii fed ex- when adult. Large specimens have a clusively with Taeniolethrinops. All pho- prominently humped head. tographs or films I have seen show C. COLOUR: Mature males and females moorii in the company of Taeniolethri- uniform dark blue, sometimes with nops spp. faint dark vertical bars. Immatures REPRODUCTION: Males do not defend blue grey with large dark blotches on territories or build nests and reproduc- the flanks. tion appears to take place opportu- MAXIMUM SIZE: 23cm TL (Konings, nistically when males and ripe females 1990a). meet. DIET: It feeds on small benthic inver- DISTRIBUTION AND ABUNDANCE: Konings tebrates stirred up by long-snouted (1990a) reports that this species is benthic feeding species such as Taenio- found throughout Lakes Malawi and lethrinops praeorbitalis, T. laticeps and Malombe. I did not record it in Lake (according to Fryer & Iles, 1972, and Malombe in 1990-92. It appears to be Eccles & Trewavas, 1989) also, along confined to shallow water and has not the coast from Nkhata Bay to Chi- been recorded from trawl catches in the lumba, Fossorochromis rostratus (Ko- 1992 survey, even those containing nings, 1995b). Eccles & Trewavas also large numbers of individuals of spe- report that Mylochromis lateristriga may cies normally used as feeding hosts. I also be used as a host in the south of have frequently observed this species the lake. At Nkhata Bay, as well as in while diving in shallow water at

187 Nkhata Bay and around the Nan- generalised benthic feeder: khumba peninsula. chironomids, gastropods, algae, and COMMERCIAL IMPORTANCE: Minor im- detritus were ingested in large quanti- portance as a food fish, but a popular ties. Annelids and ostracods also taken ornamental fish which is easily bred in (Grant & Turner, unpubl.). captivity. DISTRIBUTION AND ABUNDANCE: The types were collected at 'Bar House', at the southern tip of the SE Arm of the Trematocranus brevirostris Trewavas lake. I have collected specimens from colour photo on page 173 19-40m depth, south of Boadzulu Is- land, where it was one of the most DISTINGUISHING FEATURES: A small rela- abundant species, comprising 5.3% of tively deep-bodied species, with thin the total sample catch in the 1992 sur- vertical bars and 1-3 dark blotches on vey. Specimens from deeper water are flanks and caudal peduncle. Distin- probably not conspecific. guished from O. heterodon, O. tetra- COMMERCIAL IMPORTANCE: It comprises stigma, and O. tetraspilus by its more at least 1% of the pair trawl catch in rounded belly, larger eye, shorter the south of the SE Arm. snout, and less compressed body. Fe- NOTES: During my sampling of the males and immatures differ from those commercial fisheries, I did not distin- of Lethrinops 'yellow chin' in the pos- guish T. brevirostris from the following session of three clearly marked flank species, and relative importances are spots. Mature males can be distin- estimated subsequently from knowl- guished by their larger eyes, smaller edge of depth preferences of both the size and fewer gillrakers (11-13 cerato- fishes and the fishermen. I now con- branchials). It differs from T. 'brevi- sider these species to be distinct be- rostris deep' in its smaller eyes (32-38% cause of male colours, depth prefer- HL) and male breeding colours. ence and the (overlapping) difference COLOUR: Females and immature in the size of the eye, which is 32-37% males silvery, darker sandy-coloured head length in the specimens I have dorsally, with a large elongate dark collected from 19-30m depth south of suprapectoral blotch, a smaller fainter Boadzulu Island, 34% in both types supra-anal blotch and a third small and 37-42% in specimens from depth spot on the caudal end of the caudal of 90m, north of Monkey Bay. The peduncle. Ripe males silvery, with a specimens from 46-50m are doubtfully pinkish cast. 7-8 thin dark bars under assigned to this species. the dorsal fin, with a dark supra- Eccles (in Eccles & Trewavas, 1989) pectoral blotch. Supra-anal blotch of- placed T. brevirostris in the genus ten obscured by bars. Dorsal fin lap- Aulonocara, but at the same time de- pets white. Lips blue. Chin, cheek, and fined that genus as containing species operculum bright yellow. with vertical bars. As Eccles points out, MAXIMUM SIZE: 10cm TL. no other included species has a spot- DIET: Stomachs of three specimens ted pattern. Trematocranus microstoma, analysed indicate that the species is a the type species of Trematocranus, has

188 slightly inflated sensory pits on the MAXIMUM SIZE: 10cm TL. head, and a pattern of three spots, with DISTRIBUTION AND ABUNDANCE: Known occasional faint vertical bars. All of only from deep water, 90-102m depth, these features are shared with T. brevi- north of Monkey Bay. rostris, yet Eccles chose to reject Meyer COMMERCIAL IMPORTANCE: Members of et al.'s (1987) proposal to include T. the T. brevirostris complex comprise microstoma in Aulonocara, but included 2.5% of deep water bottom trawl catch. T. brevirostris. I feel it is better to be con- NOTES: Not reliably distinguished sistent and therefore cannot see any from T. brevirostris during sampling of justification for removing T. brevirostris commercial fisheries. from its original genus.

Trematocranus 'brevirostris deep' colour photo on page 173

DISTINGUISHING FEATURES: A small rela- tively deep-bodied species, with thin vertical bars and 1-3 dark blotches on flanks and caudal peduncle. 10-13 ceratobranchial gillrakers. Distin- guished from O. heterodon, O. tetra- stigma, and O. tetraspilus by its more rounded belly, larger eye, shorter snout, and less compressed body. Dif- fers from T. brevirostris in male breed- ing colours and in larger eye. COLOUR: Females and immature males silvery, darker grey dorsally, with a large dark suprapectoral blotch, a smaller fainter supra-anal blotch and a third small spot on the caudal end of the caudal peduncle. Ripe males sil- very grey. 6-7 dark bars under the dor- sal fin, with a dark suprapectoral blotch joining bars 2 and 3. Supra-anal blotch often obscured by bars. Dorsal fin dusky with yellow or yellow-or- ange spots and black lappets. Caudal fin dusky with yellow or orange spots, and black dorsal and ventral margin. Anal fin dusky with yellow egg-spots. Pelvic fins yellow-orange with black leading edge. Snout and lips dark grey with metallic green sheen.

189 Chapter 16 Protomelas and other horizontally-striped species

Most of the horizontally-striped spe- (1990a) prefers to place C. euchilus in cies are placed in the genera Protomelas Chilotilapia. Some specimens of some and Nyassachromis. There is no clear- of the Rhamphochromis species may ex- cut distinction between the genera. hibit a horizontal stripe. These are dealt Nyassachromis species are generally with under that genus. Horizontally- more elongated and most show a sin- striped species are almost all found in gle continuous horizontal stripe, while shallow water and few species are of most Protomelas are deeper-bodied, much significance in trawl fisheries, usually with two or three horizontal although some Protomelas species may stripes and some species often show be locally important in seines. elements of vertical barring. Some of the horizontally-striped species placed in the genus Copadichromis by Eccles & Taeniochromis holotaenia (Regan) Trewavas (1989) are considered by colour photo on page 173 Konings (1990a) to be Nyassachromis. Piscivorous species with horizontal DISTINGUISHING FEATURES: An elon- stripes are placed in Dimidiochromis if gated species with a continuous black they have a continuous stripe which stripe which extends through the eye does not extend across the snout, in and across the snout. This pattern ap- Taeniochromis if the stripe does extend pears to be unique among Malawian across the snout and in Hemitaenio- cichlids. chromis if the stripe is broken into a COLOUR: Females and immature series of spots anteriorly. Large-headed males pinkish brown, silvery ventrally, predators found exclusively on rocky with prominent black horizontal stripe shores are placed in . extending through the eyes and across Two species with specialised jaw mor- the snout. Ripe males bright blue, phology and sharing a pattern of two lighter ventrally. Dorsal and caudal broad horizontal bands are placed in fins with orange spots and stripes. monotypic genera — Chilotilapia Dorsal fin margin white. Anal fin with rhoadesii (which has short thick jaws for a broad black band and bright yellow crushing molluscs) and margin. Pelvic fins and chin black. euchilus (which has huge fleshy lips Horizontal and vertical bars some- which it uses to seal crevices between times evident. A black bar on the rocks when it sucks out small organ- preorbital bone is sometimes visible isms from within). The latter is exclu- (e.g. illustration in Baasch, 1992b). sively found on rocky shores. Konings MAXIMUM SIZE: 20cm TL (Konings,

190 1990a). Dimidiochromis kiwinge (Ahl) DIET: Pursues small fish and perhaps colour photo on page 173 invertebrates (Konings, 1990a). Two specimens I collected from the SE Arm DISTINGUISHING FEATURES: A stream- contained fish remains (Grant & lined, fairly elongated species with a Turner, unpubl.). continuous black horizontal stripe. REPRODUCTION: In the aquarium, COLOUR: Females and immature males excavate a small depression, of- males silvery ventrally, darker dorsally ten near or under a rock (Baasch, 1992b; with a prominent black horizontal Konings, 1995b). Females of 20cm may stripe. Small juveniles with a generally produce up to 100 fry. yellow-brown cast, but larger fish with DISTRIBUTION AND ABUNDANCE: Pre- a bright greenish iridescence, particu- served material examined by Eccles & larly on the dorsal surface. Ripe males Trewavas (1989) was from the far north brilliant blue with a bright orange spot of the lake (Vua and Chilumba) or un- on each flank scale (illustrated by known locations (including the holo- Konings, 1990a). type), but they report that the species MAXIMUM SIZE: 30cm TL (Konings, is found throughout the lake and 1990a). record it from trawl catches at 15-35m. DIET: Pursues small surface-living Jackson (1961) reports the species from fish (Konings, 1990a). Juveniles feed on areas of sand or weed near rocks, and zooplankton or aerial insects (Fryer, Konings (1990a) describes it as having 1959). a lakewide distribution, but specifi- REPRODUCTION: This is the most spec- cally reports and illustrates it only from tacular lek-breeder I have seen in the the area between Chilumba and lake, not just because of the size and Nkhata Bay. In the 1992 trawl survey, conspicuousness of the males, but be- it was found in many samples between cause of the incredibly close packing 20 and 64m in the SE Arm. In one sam- of their territories and almost continu- ple at 26m off Mazinzi Bay, this spe- ous aggression shown by the territo- cies dominated the catch, comprising rial fish. Leks I have observed them more one-third of the haul by weight. around the Nankumba Peninsula and I rarely recorded it south of Boadzulu at Nkhata Bay are small, generally 10- Island and did not see it in Lake 20 males, and are most common on the Malombe. rocky habitat, but occasionally over COMMERCIAL IMPORTANCE: Minor con- pure sand on steeply-shelving shores. tribution to shallow water trawl Males may defend surfaces of large catches. rocks, dig craters in the sand, or even NOTES: Eccles & Trewavas consider display in midwater. I have never ob- that Haplochromis bodyi Ahl is synony- served a solitary territorial male. Fe- mous with this species. males defend large broods of free- swimming fry and are commonly seen in shallow rocky areas. DISTRIBUTION AND ABUNDANCE: Pre- served material examined by Eccles &

191 Trewavas (1989) was from the north Popular with sport fishermen. Occa- and south ends of the lake. They also sionally exported as an aquarium fish. report that the species is found in the It must require an immense tank. Upper and Middle Shire and Lake NOTES: Eccles & Trewavas consider Malombe, but I have never observed that Haplochromis fuelleborni Ahl is this species in those areas or indeed synonymous with this species. anywhere far from a rocky shore. It is possible that it has been confused with D. strigatus. Ribbink et al. (1983) re- Dimidiochromis strigatus (Regan) corded the species at all of the their study sites on rocky shores through- DISTINGUISHING FEATURES: A large- out the Malawian sector of the lake. mouthed laterally-compressed species Although it lives and feeds at the sur- with a continuous dark horizontal face, it was not recorded by the ODA/ stripe. The mouth is larger and more SADC project's survey (Menz, 1995) steeply-angled than that of similar spe- and appears to be a littoral species cies. rather than a truly pelagic one. Juve- COLOUR: Females and immature niles often accompany shoals of Copadi- males silvery with a brownish-yellow chromis species and feed on zoo- cast and a prominent dark horizontal plankton. stripe. The ripe male is one of the most COMMERCIAL IMPORTANCE: Frequently beautiful in the lake, an unmistakable caught by hook and line, gill net and brilliant turquoise with a large red chirimila on rocky shores and reefs and patch behind the operculum, a yellow sometimes also by beach seines (Smith, chest, a broad red margin to the anal 1993). Not recorded in trawl catches. fin, and a red and white dorsal fin

Figure 49. Dimidiochromis strigatus, female, Senga Bay.

192 margin (illustrated by Konings, 1990a, traps in Lake Malombe, SE Arm of 1995b). Lake Malawi and the Upper Shire MAXIMUM SIZE: 25cm TL (Konings, River. Jackson (1961) reports it as a spe- 1990a). cies of sand and Vallisneria patches. It DIET: Predator of small fishes and seems to be confined to areas of veg- invertebrates (Eccles & Trewavas, 1989; etation and is not taken by trawls. Konings, 1990a). Jackson (1961) also COMMERCIAL IMPORTANCE: Occasion- reports vegetable remains in the stom- ally taken by beach seines, and prob- ach. ably also gillnets and hook and line. REPRODUCTION: Territorial males are Not recorded in trawl catches. Occa- solitary and according to Konings sionally exported as an aquarium fish. (1990a) defend a small nest on sand or among weed. Up to 230 fry may be produced in one brood (Konings, Dimidiochromis compressiceps 1995b). (Boulenger) DISTRIBUTION AND ABUNDANCE: Pre- served material examined by Eccles & DISTINGUISHING FEATURES: A large- Trewavas (1989) was from the north mouthed laterally-compressed species and south ends of the lake and includes with three horizontal stripes. The deep no less than 106 specimens collected by jaws and laterally compressed head are Christy in the 1920s. They also report immediately recognisable. that the species is found in the Upper COLOUR: Females and immature and Middle Shire and Lake Malombe. males generally silvery with 3 dark I have seen this species underwater at horizontal stripes, which are often dark Monkey Bay and Thumbi West Island grey, but sometimes bright metallic and in catches made by seines and green. Konings (1990a) illustrates some

Figure 50. Dimidiochromis compressiceps, immature at Chisumulu Island.

193 golden yellow specimens from Chisu- Hemitaeniochromis urotaenia mulu Island. Ripe males brilliant blue (Regan) with a broad red margin to the anal fin, colour photo on page 174 which has white egg-spots. Red spot- ted dorsal and caudal fins and a red DISTINGUISHING FEATURES: A heavily- and white dorsal fin margin (illus- built, large-mouthed species with two trated by Konings, 1990a). horizontal stripes, the lower continu- MAXIMUM SIZE: 23cm TL (Jackson, ous posteriorly, but broken into spots 1961). anteriorly. Hemitaeniochromis 'insignis' DIET: An ambush predator of small is more laterally compressed and has fishes which it stalks a head-down po- a smaller head, H. spilopterus has a sition (Eccles & Trewavas, 1989; more steeply angled-mouth. Konings, 1990a). It is extraordinarily COLOUR: Females and immature well camouflaged among vegetation. males generally silver-grey sometimes REPRODUCTION: Territorial males are with a yellowish tint, with a black hori- solitary and according to Konings zontal stripe, which is continuous (1990a) defend a small nest on sand. I posteriorly and broken into spots have observed females defending large anteriorly. There is a thinner upper numbers of very small free-swimming stripe which is also broken into spots. fry among long weeds (Potamogeton). Ripe male with a blue head and DISTRIBUTION AND ABUNDANCE: Pre- dorsum, a patch of red spots behind served material examined by Eccles & the head, and yellow flanks and belly. Trewavas (1989) was from the north Dorsal and caudal fins dark with or- and south ends of the lake. They re- ange spots. Anal fin dark with large port that the species is found in Lake yellow spots and stripes. Malombe and in the Upper Shire River, MAXIMUM SIZE: 22cm TL, but speci- where I have also recorded it. Jackson mens from Likoma, which may not be (1961) states that it is mainly a species conspecific, may reach 30cm TL of mixed rock and sand areas. Ribbink (Konings, 1990a). et al. (1983) recorded it at most of their DIET: A piscivore (Konings, 1990a). A study areas. It seems to be confined to 12.5cm specimen I collected contained vegetated areas and is quite common. a 5cm cichlid fish (Grant & Turner, COMMERCIAL IMPORTANCE: Occasion- unpubl.). ally taken by beach seines, and prob- REPRODUCTION: Territorial males con- ably also gillnets and hook and line. struct a sand castle nest near rocks Not recorded in trawl catches. Com- (Konings, 1990a). In rocky habitats at monly exported as an aquarium fish depths of 6-11m, females have been and easily bred in captivity. observed to defend free-swimming fry NOTES: It was formerly thought to until they attain a large size (Robinson, feed on the eyes of fishes (Fryer & Iles, 1995). 1972), and although there is no evi- DISTRIBUTION AND ABUNDANCE: Pre- dence for this, it is still often known as served material examined by Eccles & the 'eye-biter' in the aquarium trade. Trewavas (1989) was from the north and south ends of the lake, and they

194 report that the species is found Konings, 1995b). The breeding colour throughout the lake and in trawl reported by Eccles & Trewavas (1989) catches at depth less than 18m. In the is that of a non-territorial or ripening 1992 trawl survey it occurred at 22- male green dorsally and on the head, 26m. Underwater observations indi- yellow ventrally, with prominent cate that it is often numerous in deep stripes. areas where rocky reefs lie on sand, MAXIMUM SIZE: 23cm TL (Konings, such as Mazinzi Reef in the SE Arm. 1990a). COMMERCIAL IMPORTANCE: Minor. Oc- DIET: Probably a paedophage (Ko- casionally taken by beach seines and nings, 1990a). shallow water trawls, and probably REPRODUCTION: Territorial males con- also gill net and hook and line. struct a sand castle nest near to or NOTES: Eccles & Trewavas (1989) con- against a rock. In rocky habitats, fe- sidered Hemitaeniochromis to be mono- males defend free-swimming fry (Ko- typic, but Konings (1990a) treated it as nings, 1990a). Large numbers of terri- a subgenus of Protomelas. Konings torial males and brooding females (1990a, 1995b) illustrates a ripe male, were observed by the author at Ma- from Mdoka in the far north of the lake, zinzi Reef in February 1995. A couple which is completely yellow and has of mature males were also observed rather small jaws and also illustrates among rocks near Monkey Bay. Ko- (Konings, 1990a) a specimen in which nings has observed territorial males at the lower stripe is broken into blotches Likoma and Maleri Island. along its entire length. There are prob- DISTRIBUTION AND ABUNDANCE: The ably several species. holotype is from the SE Arm, and other preserved material examined by Eccles & Trewavas (1989) came from the ex- Hemitaeniochromis spilopterus treme northern and southern ends of (Trewavas) the lake. These authors report that the colour photo on page 174 species is found throughout the lake (as does Konings, 1995b), but did not DISTINGUISHING FEATURES: A broad fish record it in trawls. Also rarely seen in with large, deep, steeply-angled jaws 1992 survey at 28-32m, but collected by and horizontal stripes. H. 'insignis' is the author from several shallow water more laterally compressed, H. urotaenia trawls in the SE Arm and Domira Bay. has a less steeply-angled mouth. Very abundant at Mazinzi Reef and COLOUR: Females and immature occasionally seen underwater over males generally golden yellow with shallow rocky areas elsewhere. two black horizontal stripes. Ripe male COMMERCIAL IMPORTANCE: Minor. Small blue head with orange spots on dorsal specimens recorded from beach seine and caudal fin, a dark anal fin with catches in the SE Arm. May occasion- numerous large bright yellow egg- ally occur in pair trawl catches. spots. Dorsal fin margin red and white. NOTES: Eccles & Trewavas (1989) Horizontal stripe not shown while ac- placed this species in the genus Proto- tually on territory (pers. obs., see also melas, but Konings (1995b) notes that

195 the melanic pattern and overall mor- authors follow McKaye (1983) in as- phology are more similar to those of signing the name Protomelas insignis to Hemitaeniochromis urotaenia than to the small striped fish often seen near other Protomelas species. Konings rocky shores and Copadichromis breed- (1995b) has also reported a number of ing arenas, where it snatches cichlid intermediate forms, which probably eggs before the spawning females have merit specific status. Under Eccles & been able to take them into their Trewavas' classification it would be mouths. The striped specimens are of- unclear to which genus they should be ten accompanied by 3-spotted fish gen- assigned, therefore I support Konings' erally regarded as Otopharynx ovatus. proposal. Their body shape, background colour and feeding behaviour are identical. In Hemitaeniochromis 'insignis' the confusion of an attack on a spawn- colour photo on page 174 ing pair of Copadichromis, one these fishes appeared to change between the DISTINGUISHING FEATURES: A deep-bod- spotted and striped pattern. The ied species with 2 horizontal stripes, striped pattern illustrated by Konings the upper broken into a series of spots, (1990a) clearly shows signs of under- the lower broken anteriorly and con- lying spots. Konings (1990a) mentions tinuous posteriorly. More laterally that P. insignis and O. ovatus breed to- compressed and with a smaller head gether at Chinyankwazi and Chin- than H. urotaenia or H. spilopterus. yamwezi Islands. During fights for ter- COLOUR: Females and immature ritories, may change colour and be- males silvery grey with two horizon- come practically indistinguishable. tal stripes, the upper broken into a se- The ripe males illustrated by Konings ries of spots, the lower broken appear to differ only in the extent that anteriorly and continuous posteriorly. the blue colour overlies the underly- MAXIMUM SIZE: 20cm TL (Konings, ing yellow, which is often the kind of 1990a). difference shown between conspecifics DISTRIBUTION AND ABUNDANCE: The at different stages of reproductive state. types of P. insignis are from Monkey I think the spotted and oblique-striped Bay. Eccles & Trewavas (1989) did not paedophages may be conspecific and record it in trawls. In the 1992 survey, will presently regard them as O. ovatus. H. 'insignis' was recorded only from a Konings (pers. comm.), who has ob- single sample, taken at 128m off served these fishes more that I have, Domwe Island, but it has also been re- thinks there may be as many as three corded by the author in semipelagic species in this group. I have not exam- and pair trawl catches in the SE Arm, ined the types of P. insignis, but the and off Domira Bay at 74m, Monkey melanin pattern, deep body and slight Bay 90m and 24-28m Palm Beach- nuchal hump of the drawing of the Maldeco. type in Eccles & Trewavas (1989) seem COMMERCIAL IMPORTANCE: Minor com- to fit the species I have collected rather ponent of trawl catches. better than they do the shallow water NOTES: Konings (1990a) and other paedophage. The paedophage species

196 lacks a mental process, which is fairly vas (1989) came from the extreme prominent in the specimens I have as- northern and southern ends of the lake. signed to H. 'insignis', which also have They recorded this species from a few a more steeply-angled gape. The speci- trawls made between 15 and 18m men illustrated by Konings (1990a: 147 depth. Jackson (1961) reports that it is #5) as Protomelas cf insignis appears to abundant, and found in river mouths be the same species, and the ripe male as well as the main lake. Konings illustrated as Protomelas sp. 'insignis (1990a) states that it is found through- mumbo' may also be conspecific. Fur- out Lake Malawi and Lake Malombe. ther study of the types is needed be- In 1990-92, it was one of the few larger fore it would be possible to determine haplochromine cichlids (i.e. mature at if H. 'insignis' is actually the species a length of greater than 8cm TL) still classed as Protomelas insignis by Eccles present in reasonable numbers in Lake & Trewavas. Malombe. In the 1992 trawl survey it was recorded from a single station at Protomelas similis (Trewavas) 18m depth on the eastern shore of the colour photo on page 175 SE Arm, north of Boadzulu Island, but was also recorded from shallow water DISTINGUISHING FEATURES: A deep-bod- trawls made further south. It is princi- ied species with two horizontal stripes, pally a species of shallow weedy areas and generally golden-brown back- and can often be observed by snorkel- ground colour. P. kirkii has a longer, ling. more acute snout and longer jaws, P. COMMERCIAL IMPORTANCE: Minor com- labridens has a more steeply-angled ponent of trawl catches. Frequently mouth. taken by beach seines indeed the COLOUR: Females and immature small-meshed beach seine most com- males silvery, brownish dorsally, with monly used is known as the 'Kambuzi a generally golden cast and yellowish Seine' after the local name for P. similis pelvic and anal fins. There are two thin and its relatives. At present beach seine black horizontal stripes, the upper one catches in southern Lake Malawi and is often indistinct. Males are bright tur- Lake Malombe are dominated by spe- quoise blue with fins conspicuously cies which mature at far smaller sizes, patterned with red, white, and orange but the name of the gear suggests that (see Konings, 1990a). it was formerly of greater importance MAXIMUM SIZE: 17cm TL (Eccles & than it is at present, or that it may be Trewavas, 1989). more important in the northern parts DIET: Macrophytes and algae (Fryer of the lake which are less heavily & Iles, 1972). fished. Occasionally exported as an REPRODUCTION: Males clear a small aquarium fish. circle among short weeds, such as Vallisneria, in very shallow water (less Protomelas kirkii (Günther) than 2m depth). colour photo on page 175 DISTRIBUTION AND ABUNDANCE: The material examined by Eccles & Trewa- DISTINGUISHING FEATURES: A deep-bod-

197 ied species with two horizontal stripes, and generally golden-brown back- and generally golden-brown back- ground colour. It is distinguished from ground colour. P. similis has a shorter, most similar species by its more less acute snout and shorter jaws, P. steeply-angled mouth and shorter labridens has a more steeply-angled jaws, and from H. spilopterus by its thin- mouth. ner jaws and less steeply-angled COLOUR: Females and immature mouth. males silvery, brownish dorsally, with COLOUR: Females and immature a generally golden cast and yellowish males silvery, brownish dorsally, with pelvic and anal fins. There are two thin a generally golden cast and yellowish black horizontal stripes, the upper one pelvic and anal fins. There are two thin is often indistinct. Males are bright tur- black horizontal stripes, the upper one quoise blue with fins conspicuously is often indistinct. Males (not fully patterned with red and white (see ripe?) turquoise blue, yellowish ven- Konings, 1990a). trally with fins conspicuously pat- MAXIMUM SIZE: 11cm TL (Eccles & terned with red and white (see Ko- Trewavas, 1989). nings, 1995b). DIET: Invertebrates, especially those MAXIMUM SIZE: 16cm TL (Eccles & found on macrophytes and among Trewavas, 1989). sand (Eccles & Trewavas 1989). Crus- DIET: Eccles & Trewavas suggest it taceans and snails (Konings, 1995b). may feed on snails, on the basis of its REPRODUCTION: Males build small enlarged pharyngeal dentition. sand-castle nests in very shallow wa- DISTRIBUTION AND ABUNDANCE: The ter (less than 2m depth), often coloni- material examined by Eccles & Trewa- ally (Konings, 1995b). vas (1989) came from the extreme DISTRIBUTION AND ABUNDANCE: The northern and southern ends of the lake. material examined by Eccles & Trewa- Jackson states that it is less abundant vas (1989) came from the extreme than P. kirkii and is mainly found in the northern and southern ends of the lake. south of the lake. Principally a species Jackson records it as abundant in shal- of shallow weedy areas, it was re- low sandy shores and sheltered areas. corded in a single shallow water trawl Neither I, nor Eccles recorded it in catches by Eccles & Trewavas and once trawl catches and it seems to be prin- seen by the author from a trawl haul cipally a species of shallow weedy ar- made at 15-18m in the SE Arm. eas. COMMERCIAL IMPORTANCE: May be of COMMERCIAL IMPORTANCE: Frequently minor importance in beach seine taken by beach seines, but not of ma- catches. jor importance. Protomelas marginatus (Trewavas) Protomelas labridens (Trewavas) colour photo on page 175 DISTINGUISHING FEATURES: A horizon- tally striped species with a large mouth DISTINGUISHING FEATURES: A deep-bod- and eye. ied species with two horizontal stripes, COLOUR: Females and immature

198 Figure 51. Protomelas marginatus, female, Upper Shire River. males silvery grey with two horizon- sheltered sandy areas, being most tal stripes. Male breeding colour (from abundant at Nkhotakota and in the Upper Shire) dark blue, with a red spot south. Eccles & Trewavas did not on each flank scale and a faint dark record the species from the field. midlateral stripe. Dorsal fin dark blue, Konings (1995b) photographed a sin- lighter posteriorly with faint orange gle specimen at Mdoka, in the far spots on soft rayed area. White mar- north. I have found it in experimental gin and red lappets. Caudal rays dark fyke net catches in the Upper Shire blue, membranes white with orange River and have tentatively identified spots. Anal fin membranes dark proxi- individuals while Scuba diving around mally, white distally. Spines white with shallow rocky areas just south of Mon- pink tips. Pelvics white anteriorly, key Bay. black posteriorly. Rays white. Konings COMMERCIAL IMPORTANCE: Negligible. (1995b) illustrates a male from the NOTES: Eccles & Trewavas (1989) con- north of the lake, which is generally of sidered that northern and southern a similar colour, but paler blue. specimens represent distinct subspe- MAXIMUM SIZE: 17cm TL (Eccles & cies (P. m. marginatus and P. m. vuae). Trewavas, 1989). They differ in mean eye diameter and DIET: Eccles & Trewavas (1989) record in modal values of lower gillraker and stomach contents consisting of plant dorsal spine counts. All morphometric material, sponge and other inverte- and meristic values presented show brates (unspecified). considerable overlap. Konings (1989) DISTRIBUTION AND ABUNDANCE: The treated them as separate species, but material examined by Eccles & Trewa- later treats them as subspecies (Ko- vas (1989) came from the extreme nings 1990a, 1995b). northern and southern ends of the lake. Jackson reports that it is common in

199 Protomelas triaenodon (Trewavas) red margin to the dorsal fin. Eye larger colour photo on page 175 (32-33% HL) than that of P. pleurotaenia (24-29 % HL). Lower gillrakers (10-11) DISTINGUISHING FEATURES: A heavily- fewer than P. triaenodon or P. marginatus. built, deep-bodied species with a bright It lacks the enlarged pharyngeal teeth silvery body, two thin horizontal stripes, of P. macrodon. and a red margin to the dorsal fin. COLOUR: Females and immature males COLOUR: Females and immature males silvery with two thin horizontal stripes silvery with yellowish fins, two thin (the upper generally broken into spots) horizontal stripes and a number of dark and a number of dark spots on the base spots on the base of the dorsal fin. Dor- of the dorsal fin. The dorsal fin margin sal fin margin bright red. is bright red. MAXIMUM SIZE: 15.5cm TL (Eccles & MAXIMUM SIZE: 8.5cm TL. Trewavas, 1989). DISTRIBUTION AND ABUNDANCE: A shal- DISTRIBUTION AND ABUNDANCE: The ma- low water species, known from beach terial examined by Eccles & Trewavas seine catches in Monkey Bay and a sin- (1989) came from the SE and SW Arms, gle trawl haul NW of Boadzulu Island but they also report it from trawls be- at 13-23m depth. tween 10 and 18m depth and from Lake COMMERCIAL IMPORTANCE: Minor, prob- Malombe and the Upper and Middle ably taken in seines. Shire River. Not recorded by Jackson. NOTES: Confused with P. triaenodon Konings (1995b) has not observed the during the first part of my survey. This species underwater and suggests that it species may be conspecific with what does not occur north of Senga Bay. In Konings (1995b) calls P. pleurotaenia. the 1992 survey, it occurred in small numbers between 20 and 45m, but be- Chilotilapia rhoadesii Boulenger tween 1990 and 1992 was recorded at colour photo on page 175 the Nkope, Mazinzi, Fowo and Chekopa stations in the SE Arm. I did not record DISTINGUISHING FEATURES: A distinctive the species to the south of Boadzulu Is- heavy-headed species with a short land, in Lake Malombe nor in the Shire snout, short powerful jaws and two River. horizontal stripes. COMMERCIAL IMPORTANCE: Minor, occa- COLOUR: Females and immature males sionally recorded in trawls. golden brown with two broad dark hori- NOTES: The specimens illustrated by zontal stripes. Ripe males bright blue Konings (1990a) as Nyassachromis nigri- with numerous pale egg-spots on the taeniatus appear to be P. triaenodon. anal fin. MAXIMUM SIZE: 30cm TL (Konings, Protomelas 'red dorsal' 1990a). colour photo on page 175 DIET: It feeds on snails, Melanoides and Lanistes (Konings, 1990a), although DISTINGUISHING FEATURES: A small spe- small specimens (9-10cm SL) may also cies with a bright silvery body, two thin ingest copepods and diatoms (Grant & horizontal stripes, faint vertical bars and Turner, unpubl.).

200 REPRODUCTION: Konings (1990a) states pattern of two horizontal broken stripes that the species lives in schools in which and bright red dorsal fin lappets. only a single male maintains reproduc- COLOUR: Females and immature males tive coloration and spawning takes silvery with two broken horizontal place within the foraging area of the stripes, the upper usually manifested as school. He also notes that males may be a series of spots. Dorsal fin lappets red. colonial and construct nests. This behav- Ripe males turquoise-green with numer- iour has also been noted by Stauffer ous orange spots in dorsal and caudal (pers. comm.) from the vicinity of fins. Dorsal fin margin white with red Kanchedza Is. Konings further notes tipped-lappets. Anal fin with numerous that male bowers are apparently marked large whitish-yellow egg-spots. with heaps of empty Lanistes shells. MAXIMUM SIZE: 17cm TL (Eccles & Stauffer (unpublished data) marked Trewavas, 1989). hundreds of empty Lanistes shells and DIET: Konings (1990a) reports that it scattered them near a Chilotilapia arena. feeds on small benthic invertebrates. The He found that the males did not move stomachs of two specimens from the SE them onto their bowers and concluded Arm contained chironomid larvae, that the shells present in the bowers had copepods, diatoms, sand, and detritus merely been uncovered in the course of (Grant & Turner unpubl.). the males' digging activities. Ribbink et DISTRIBUTION AND ABUNDANCE: Konings al. (1983) report that the males may de- (1990a) reports that this species is found fend territories over rocks. throughout Lakes Malawi and DISTRIBUTION AND ABUNDANCE: Eccles & Malombe, and illustrates specimens Trewavas state that the species is found from Senga Bay and the SE Arm. Eccles throughout Lake Malawi on sandy sub- & Trewavas state that it is widespread strates and recorded it from trawls at in the southern part of the lake and in depths of 30-60m, and once (off Nkhota- inflowing rivers. Jackson mentions that kota) at 90m. Konings (1990a) reports a it is commoner in the south. In 1992, it lake-wide distribution. In the 1992 sur- was abundant south of Monkey Bay and vey it was frequently taken in small especially south of Boadzulu Island at numbers from 20-30m. It was common depths of 34m or less. I also found it to in fyke net catches in the Upper Shire, be common in Lake Malombe and the and in seines and trawls in the SE Arm. Upper and Middle Shire River. It seems It was not recorded in Lake Malombe. to prefer muddy areas. COMMERCIAL IMPORTANCE: A minor com- COMMERCIAL IMPORTANCE: An signifi- ponent of seine and trawl catches. Prob- cant component of seine and pair trawl ably also taken in gill nets. Occasionally catches. Probably also taken in gill nets. exported as an aquarium fish. NOTES: The type species of Lethrinops, but rather a distinctive form, really Lethrinops lethrinus (Günther) only similar to L. argenteus, which is to colour photo on page 176 date known only from the far north of the lake. It may be unrelated to the DISTINGUISHING FEATURES: A long- majority of the species presently placed snouted species, which has a distinctive in Lethrinops.

201 Chapter 18 Nimbochromis and other blotched species

Nimbochromis is a small genus of dis- whitish with large dark brown blotches tinctively marked species. All share a over the body and fins. There is a general colour pattern of irregular dark prominent dark bar on the preorbital brown blotches distributed over the bone. Mature males silvery blue with whole body. All have large mouths and a white and red dorsal fin margin and are piscivores. and Fos- a broad red anal fin margin. sorochromis are other genera character- MAXIMUM SIZE: 25 cm TL (Konings, ised by a pattern of numerous large 1990a). spots. DIET: An ambush predator of small In addition to the species discussed fish, which often lies, partly buried, on below, Nimbochromis fuscotaeniatus its side on sandy or muddy bottoms. (Regan) is occasionally observed in It is very cryptic. shallow water in the south of the lake. REPRODUCTION: Konings (1990a) re- It is the most elongate Nimbochromis, ports that the males defend an area of with the dark blotches tending to join sand adjoining a rock. together to form horizontal stripes. DISTRIBUTION AND ABUNDANCE: Material Konings (1990a) and Eccles & Trewa- examined by Eccles & Trewavas came vas (1989) illustrate the species and from both northern and southern ends discuss its biology. Although Konings of the lake. Ribbink et al. (1983) re- reported that it was found in Lake corded it from all of their study sites Malombe, it did not occur in the on rocky shores throughout the Ma- catches I sampled in 1990-91. It may lawian sector of the lake. Eccles & sometimes be taken by seines in south- Trewavas report the species from ern Lake Malawi. Malombe and the Upper Shire, and I also recorded it in these areas in 1990- 92. Jackson (1961) states that this spe- Nimbochromis livingstonii cies is found only among Vallisneria (Günther) beds, which is clearly not true of the colour photo on page 176 deep water individuals. In the 1992 survey, I recorded it from the shallow- DISTINGUISHING FEATURES: A blotched est trawls to as deep as 114m. Also fre- species with a pale whitish back- quently observed by snorkelling in ground colour. Pectoral fins spotted, shallow water. but no small spots elsewhere on the COMMERCIAL IMPORTANCE: A minor body. component of many fisheries and a COLOUR: Females and immatures major ornamental species, easily bred

202 in captivity. illustrated by Axelrod (1993) are gen- NOTES: Jackson (1961) and McKaye erally pale blue with numerous small (1981) interpret the ambush strategy of orange flank spots and many whitish this fish as functioning by attracting egg-spots on the anal fin. The anal fin small prey fish by mimicking a dead margin is orange-red. fish. I consider this unlikely, both from MAXIMUM SIZE: 23 cm TL (Konings, observing the behaviour of the poten- 1990a). tial prey and on theoretical grounds — DIET: Eccles & Trewavas describe it mimics are always rarer than the as a piscivore. It adopts a variety of 'model' that they mimic, and dead fish feeding strategies: sometimes acting as are far less often seen in the lake than a solitary ambush predator, but also hunting N. livingstonii. I have seen hunting in packs (pers. obs.; Konings, small fish attracted to the clouds of 1990a). When adopting the ambush sand stirred up when an individual strategy, it often lies on the substrate, buries itself, but not otherwise ap- but it has not been observed to lie on proaching a stationary fish. I think that its side, in the manner of N. livingstonii. the colour pattern probably serves as At Chembe Beach, Cape Maclear, I disruptive camouflage. This species is have often observed this species to unusual in its occupation of such a hunt in groups of 2-10 individuals, of- wide range of habitats and depths. Jay ten in the company of other species Stauffer (pers. comm.) is presently car- such as F. rostratus, P. johnstonii, A. rying out taxonomic work to deter- christyi, and C. spilorhynchus. Konings mine if deep water and shallow water (1990a) has also observed small Bucco- populations are conspecific. chromis heterotaenia in such groups. These groups move quickly over sandy and rocky substrates, attacking groups Nimbochromis polystigma (Regan) of small cichlids, often fry guarded by colour photo on page 176 females, or foraging in the sand or mud, presumably for benthic inverte- DISTINGUISHING FEATURES: A blotched brates. species with a pale yellowish back- REPRODUCTION: Konings (1990a) re- ground colour. Small brown spots ports that the males defend an area of cover the body and fins. The snout is sand adjoining a rock, and they may not decurved as in N. linni. spawn on the rock surface or the sand. COLOUR: Females and immatures yel- Females guard free-swimming fry. He lowish with large orange brown to reports it to be a non-seasonal breeder. dark brown blotches over the body and DISTRIBUTION AND ABUNDANCE: Material fins. Superimposed on this pattern are examined by Eccles & Trewavas mostly numerous small brown spots. The came from the SE and SW Arms, or smaller juveniles are generally an or- from unknown locations (including the ange hue. Konings (1990a) illustrates types), although a single specimen was a mature male which is blue with a from Karonga in the north. Jackson white and red dorsal fin margin and a records it as a rock-frequenting species. broad red anal fin margin, while those Konings (1990a) reports it as having a

203 lake-wide distribution and being Nimbochromis venustus (Boulenger) found in all habitats, although most colour photo on page 176 commonly in areas of mixed rocks and sand or in vegetated areas. Eccles & DISTINGUISHING FEATURES: A blotched Trewavas regard it as characteristic of species with a pale yellowish back- weeded areas, especially near rocks ground colour and bright yellow pel- and also report a lake-wide distribu- vic and anal fins. It lacks the small tion, although it occurred only in two spots on the body and fins. The pecto- of their trawls, at 18 and 55m in the SE ral fins are unspotted. It is deeper-bod- and SW Arms respectively. Ribbink et ied than other congeneric species and al. (1983) found it at all of their survey the large blotches tend to be vertically areas on rocky shores. In the 1992 sur- — rather than horizontally-expanded. vey, it was recorded at two stations, COLOUR: Females and immatures yel- both at 26m, and I also collected the lowish with large dark brown blotches species at 15-18m in the SE Arm and over the body and fins. The dorsal fin have frequently observed it by snorkel- has a white margin. Mature males gen- ling or diving in shallow water. erally exhibit the spotted pattern, but COMMERCIAL IMPORTANCE: A minor the background colour becomes bright component of seine net catches in the yellow. The dorsal fin is bright yellow SE Arm and probably elsewhere. A and this colour extends as a forehead popular ornamental species, easily blaze to the snout. The rest of the head, bred in captivity. as well as the chest, bright metallic NOTES: Two specimens assigned to N. blue. The anal fin is blue with a broad pardalis (Trewavas) by Eccles & Trewa- white margin. vas were collected from Chilumba and MAXIMUM SIZE: 22.5 cm TL (Konings, Nkhata Bay in the north of the lake. 1990a). These individuals are more slender DIET: Eccles & Trewavas describe it than the material assigned to N. as a piscivore. Konings (1990a, 1995b) polystigma or N. livingstonii. They also states that small individuals, at least, have a pronounced dark bar between are ambush predators which rest partly the eyes like N. livingstonii, which is not buried in the sand. Axelrod (1993) found in typical N. polystigma. A fur- states that they lie on their sides, but it ther poorly preserved specimen from is not clear if this is a misinterpreta- an unknown locality has been de- tion of the reports of Konings, for it scribed as N. maculimanus (Regan). does not appear to have been reported This differs from N. polystigma only in elsewhere. Konings (1995) reports that having a larger number of jaw teeth. I it also feeds on invertebrates. am inclined to agree with Konings REPRODUCTION: Konings (1990a) re- (1990a), who believes that both N. ports that the species breeds on sand, maculimanus and N. pardalis are syno- but has not observed a nest. nyms of N. polystigma. DISTRIBUTION AND ABUNDANCE: Material examined by Eccles & Trewavas came from the northern and southern ends of the lake. Jackson states that it is a

204 species of sandy and probably also (1990a) has a dull bluish cast. The anal weedy areas. Konings (1990a) consid- fin has a broad orange margin with ir- ers the species to be distributed regular white stripes and spots. throughout Lakes Malawi and Malom- MAXIMUM SIZE: 30 cm TL (Konings, be, but that it prefers depths of 15m of 1990a). more (1995b). Ribbink et al. (1983) re- DIET: An ambush predator which corded the species only in the SE Arm feeds on small mbuna hiding under and Nankumba Peninsula. Eccles & rocks. It often remains motionless, up- Trewavas reported it to be widespread right, resting on the top of a rock for a in shallow sandy areas, and often taken while before striking downwards un- in trawls at 10-18m, where it was some- der the rock. This behaviour has been times abundant. In the 1992 survey, it observed by Konings (1990a), Stauffer was recorded in small numbers at two (pers. comm.), and myself. Axelrod stations, at 20 and 42m, and I also col- (1993) claims that individuals 'play lected the species at 15-18m in the SE dead' by lying on the bottom. Arm. I did not record it in Lake REPRODUCTION: Konings (1990a) re- Malombe. ports that males defend territories in COMMERCIAL IMPORTANCE: A minor the rocky habitat, or on the sand be- component of seine net catches in the side a rock (Konings, 1995b), but do not SE Arm and probably elsewhere. A build nests. Females guard fry over popular ornamental species, easily rocks or nearby areas of sand. bred in captivity. DISTRIBUTION AND ABUNDANCE: A wide- NOTES: Eccles & Trewavas consider spread fish of rocky shores. Eccles & Haplochromis simulans (Regan) to be a Trewavas and Konings (1990a) con- synonym of N. venustus. sider the species to be distributed throughout the rocky areas of Lake Malawi. Ribbink et al. (1983) recorded Nimbochromis linni the species from all of their study ar- (Burgess & Axelrod) eas except Usisya and the vicinity of colour photo on page 176 Senga Bay. I did not record it in the 1992 survey, but collected a single specimen DISTINGUISHING FEATURES: A blotched from a trawl at around 50m depth close species with numerous small spots on to the rocky points of Chirombo and the body and fins, including the pec- Nkhudzi. torals. It is distinguished from N. COMMERCIAL IMPORTANCE: Trivial. Per- polystigma by its characteristically haps taken by gillnets or hooks over decurved snout. rocky areas. A popular ornamental COLOUR: Females and immatures species, bred in captivity. whitish brown with large dark brown blotches over the body and fins. The pattern of large blotches is almost ob- Eclectochromis ornatus (Regan) scured by the large number of small dark spots covering the entire surface. DISTINGUISHING FEATURES: Small spe- A mature male illustrated by Konings cies with a long snout, thick lips and a

205 Figure 52. Eclectochromis ornatus, male, Lake Malombe. pattern of irregular vertical bars and kota and the type of E. festivus is from spots. Nkhudzi. Konings (1990a) reports the COLOUR: Females and immature species from all round the lake. I have males silvery-grey with six vertical observed specimens which appeared bars under the dorsal fin and two more to be this species from muddy areas on the caudal peduncle. The bars are near Kanchedza Island, Chirombo Bay irregular and tend to be broken into and have seen specimens collected three rows of spots. Konings (1990a) from Lake Malombe and the Upper illustrates ripe males which are blue, Shire River. Konings considers that this with an orange chest and chin. There species is confined to very shallow are small orange spots on the flank water (1-10m) in 'intermediate' habitat scales, caudal and dorsal fins. The dor- — areas where a few rocks are scattered sal has a broad white border with a red amidst a sandy habitat. Occasionally tip. The anal fin has numerous whit- seen in groups of 5-20. Not abundant. ish egg-spots and a broad orange bor- COMMERCIAL IMPORTANCE: Probably der. A black stripe on the preorbital taken in seines and gillnets in suitable bone is visible in some fish. habitats, but unlikely to be of much MAXIMUM SIZE: 25cm TL (Konings, importance. 1990a). NOTES: Eccles & Trewavas (1989) rec- DIET: Invertebrates and small cichlids ognised three species: E. ornatus which are often sucked from horizon- (Regan), E. festivus (Trewavas) and E. tal cracks (Konings, 1990a). lobochilus (Trewavas). The genus was REPRODUCTION: Ripe males are seen all poorly known at that time — they ex- year round and defend a cave under amined only 4 specimens of all three rocks (Konings, 1990a). species combined. The lip folds of E. DISTRIBUTION AND ABUNDANCE: An un- ornatus and E. lobochilus were continu- common shallow water species of shel- ous across the lower jaw, while the left ter soft-bottomed areas. Eccles & Tre- and right lip folds of E. festivus were wavas report the species from Nkhota- separate. E. ornatus had 11-13 lower

206 gillrakers and molariform teeth on the he had previously (1989) confused E. lower pharyngeal bone. E. festivus (12 lobochilus and E. ornatus. The holotype gillrakers) and E. lobochilus (15-16) had of E. lobochilus was collected from enlarged central teeth on the lower Chilumba, and Eccles & Trewavas re- pharyngeal, but these were sharp not port a single tentatively identified rounded. I collected a single sexually specimen from Mozambique. Konings immature 10cm TL specimen from an reports it is common on rocky coasts experimental fyke net on the Upper throughout the lake. This may explain Shire River, which keys out as E. Jackson's (1961) records of E. ornatus festivus on the basis of the gillraker as a species of rocky shores. Konings count (13) and lack of molariform (1989, 1990a) also chose to include the teeth. Its lips are not enlarged at all and species of Eclectochromis within the ge- it was not possible to assess whether it nus Protomelas, a move I do not adopt, has paired or unpaired folds. An ap- in the interests of nomenclatural sta- parent male (20cm TL) from Lake bility, although there seems little to dis- Malombe, was found in a voucher col- tinguish between these genera. lection in the Monkey Bay Fisheries Station, and had been labelled by Ec- cles as E. ornatus. It has 12 lower Fossorochromis rostratus gillrakers and continuous lip folds. (Boulenger) Konings (1990a) considers E. festivus to be a junior synonym of E. ornatus DISTINGUISHING FEATURES: A large spe- (Regan) a possibility already raised by cies with a long snout and three rows Eccles & Trewavas. I am inclined to of bold spots. accept this — both specimens under COLOUR: Females and immature 10cm TL key out as E. festivus and all males yellowish with three rows of three greater than 14cm TL key out as dark spots. Ripe males dark blue, al- E. ornatus. Konings (1990a) admits that most black, with an irregular area of

Figure 53. Fossorochromis rostratus, female, middle Shire River.

207 pale metallic blue on the flanks and but other specimens overlap in these upper surface. respects. In neither case is the location MAXIMUM SIZE: 35cm TL (Konings, of the types known with more preci- 1995b). sion than 'Lake Nyasa'. This species is DIET: Invertebrates buried in the sand almost always seen in small groups (Konings, 1990a). In the south, I have Konings (1990a) states 20-40, but I observed medium-sized specimens think it could sometimes be far fewer. hunting small fish in the company of Konings states these consist of adults, N. polystigma and other piscivores. but I have observed that there is often REPRODUCTION: Konings (1990a) a striking variation in size. It is not stated that ripe males are seen all year known if these groups persist for long round and do not seem to build a nest periods, but if so, it could be a harem or defend a territory, but later (1995b) polygynist, as there is generally only a noted that males sometimes formed single male in each foraging group. aggregations and dug large crater nests in shallow water (1-2m deep). Repro- ductive activity is highest in the morn- ing (Konings, 1995b). At Nkhata Bay, near rocks, I have seen a female de- fending free-swimming fry. DISTRIBUTION AND ABUNDANCE: Eccles & Trewavas examined material from the northern and southern ends of the lake. Tweddle et al. (1979) recorded it from the Shire River, extending downstream from Liwonde. Konings (1990a) reports it from throughout Lake Malawi and Lake Malombe. It has not been re- corded from trawls. I have found it in seine catches in the SE Arm, and the middle Shire River, and gillnets from Cape Maclear and the Upper Shire. I did not find it in Lake Malombe. From underwater observations, I have found it to be abundant in shallow sandy ar- eas around Cape Maclear, the SE Arm, and Nkhata Bay. COMMERCIAL IMPORTANCE: Probably important in seines and gillnets in suit- able habitats. NOTES: Eccles & Trewavas consider Haplochromis macrorhynchus to be a jun- ior synonym. The types of the two dif- fer in gillraker counts and head shape,

208 Chapter 19 Discussion

Species inventory mis trilineatus, T. variabilis, Trematocra- nus labifer. Although some of species The present work cannot be consid- are likely to be synonymous with those ered as a comprehensive inventory of I have discussed above, many are defi- the species of the offshore areas of nitely not and are well-known from Lakes Malawi and Malombe. Virtually shallow water habitats (Konings, all of the material came from a very 1990a, 1995a). I suspect that many off- small portion of the lake surface. Many shore species have restricted distribu- species recorded by Eccles & Trewavas tions within the lake, and thus that were not encountered by me and are there are many more species awaiting thus not discussed, except sometimes discovery in parts of the lake I have yet in the course of distinguishing them to survey. No collections of offshore from taxa I collected. Leaving aside fishes have ever been made in Mozam- those species known to be confined to bique, and very little material has been rocky shores, the following may (or collected from the whole stretch of the may not) be found in offshore habitats: Malawian shoreline from Senga Bay to Aulonocara auditor, A. nyassae, A. Nkhata Bay. The last collection from trematocephalum, Buccochromis atri- the Tanzanian coast was described by taeniatus, B. oculatus, B. spectabilis, Co- Ahl in 1927. Clearly much more work padichromis mloto, C. pleurostigmoides, needs to be done. Dimidiochromis dimidiatus, Lethrinops albus, L. argenteus, L. auritus, L. leptodon, Habitat preferences L. lunaris, L. macrophthalmus, L. margin- atus, L. microstoma, L. oculatus, L. Previous studies (e.g. Fryer, 1959; parvidens, Mylochromis balteatus, M. Ribbink et al., 1983; Konings, 1989, epichoralis, M. guentheri, M. incola, M. 1990a, 1995a) have emphasised that labidodon, M. mola, M. mollis, M. obtusus, rocky shore cichlids often have very Nimbochromis maculimanus, N. pardalis, restricted habitat preferences. The Nyassachromis boadzulu, N. breviceps, N. present work demonstrates that many leuciscus, N. microcephalus, N. nigri- offshore species certainly have re- taeniatus, N. prostoma, N. purpurans, N. stricted depth distributions, although serenus, Otopharynx heterodon, O. ovatus, the depth range occupied is generally O. selenurus, Placidochromis electra, P. greater than those of many rocky shore stonemani, Protomelas annectens, P. species. There is some indication of insignis, P. macrodon, P. pleurotaenia, P. habitat preferences within a particular phenochilus, P. virgatus, Sciaenochromis depth range, according to substrate ahli, Stigmatochromis pleurospilus, type: species which are abundant in Taeniolethrinops cyrtonotus, Tramitichro- Lake Malombe tend to be found in

209 similar habitats (sheltered muddy crustacea, and insect larvae. Special- bays) in Lake Malawi. However, some ised fin biters, scale eaters, and paedo- species are almost ubiquitous, such as phages represent only a few uncom- Nimbochromis livingstonii and Copadi- mon species in each habitat. Overall, chromis virginalis, notably species there have been only a few quantita- which are also found over rocky tive diet studies (reviewed in Turner, shores. Overall, it appears that offshore 1994d). It is certainly possible that di- species are much less restricted to etary specialisations may be important patches of favourable habitats than are seasonally or during intermittent pe- rocky shore species, although large riods of crisis. A fundamental problem topographic features such as deep is that stomach contents are samples rocky coasts (e.g. the Nankumba Pe- over a short period and give only a ninsula) or major river mouths may act snapshot of the recent food intake, as barriers to movement of some spe- which may vary seasonally or accord- cies. This means that it is less easy to ing to age. Stable isotope studies, see how allopatric speciation could which provide an integrated sum of the have occurred in the non-rocky shore lifetime's diet of an individual, may be species. a promising approach (Bootsma et al., in press), but year-round behavioural Trophic specialisation observations of how and where food is gathered will also be essential. At The pioneering work of Geoffrey present, there is no compelling reason Fryer (e.g. Fryer, 1959; Fryer & Iles, to think that trophic specialisation has 1972) demonstrated the incredible played any role in the speciation of range of feeding structures and behav- Lake Malawi cichlids. iour within the Malawian cichlids. Many subsequent accounts have rep- Reproduction and life cycles resented these as feeding specialisa- tions which have evolved to limit com- All of the endemic cichlids of Lake petition for food, allowing the co-ex- Malawi which have been studied are istence of so many different species. In maternal mouthbrooders. What is fact, Fryer (op cit.) suggested the re- known of the less well-studied species verse, namely that many species ap- is consistent with the hypothesis that peared to occupy virtually identical this is the universal reproductive sys- ecological niches in violation of tem of the haplochromines and Oreo- Gausse's competitive exclusion princi- chromis species, in Lake Malawi or else- ple. Fryer's suggestion seems all the where. more plausible now that we believe Males of most shallow-water species that many of the forms he considered construct spawning platforms or bow- to be colour morphs may in fact be ers out of sand or mud. Most species separate species (Ribbink et al., 1983). aggregate in arenas or leks. Particular The present work indicates that most species complexes or genera often have offshore species feed on a few basic characteristic bower forms (Konings, resources: plankton, diatoms, benthic 1995b). Although there may be consid-

210 erable variation in size or shape of rental behaviour of many genera, in- structures produced by closely-related cluding such important ones as Lethri- species (Turner et al., 1991b, Stauffer et nops, Alticorpus, Mylochromis, Rhampho- al., 1993), these characteristics may also chromis, and Diplotaxodon. Many spe- be influenced by substrate type, water cies, irrespective of whether they guard depth and degree of water movement. fry, tend to release them in shallow It is likely that most bottom-living off- nursery areas. Species using inshore shore species also construct bowers nurseries include members of some and certainly large numbers of ripe genera with offshore representatives males are often trawled together, sug- such as Rhamphochromis and Copadi- gesting they form leks. Males of some chromis. Juveniles of other taxa, such species may defend rock surfaces while as Diplotaxodon, Alticorpus and the others are not known to be territorial Placidochromis hennydaviesae complex and may breed opportunistically when have never been found in shallow wa- ripe females encounter sexually active ter and it seems likely that many deep- males. Male Copadichromis 'chryso- water or pelagic species complete their notus blue' defend territories in mid- entire life cycle within the adult habi- water, although in the vicinity of rocks, tat. and females spawn in midwater, catch- ing the eggs in their mouths before they Sexual selection and fall to the bottom (Eccles & Lewis, species recognition 1981). It is likely that some deep-wa- ter pelagic species, particularly Diplo- Exclusively maternal care, lek forma- taxodon spp., may breed in midwater tion, and strong sexual dimorphism are far offshore, possibly without needing all features of strongly sexually se- to orientate their territories to under- lected species. Sexual selection is the lying substrate features. If so, they process by which traits evolve because would be the first known cichlid spe- they are beneficial in competition over cies to be permanently pelagic. mates, either through fighting or be- Parental care is exclusively by the cause they are attractive to the oppo- female in all known haplochromine site sex. Bright colours, long fins, and Oreochromis species. Among shal- bower construction, lek formation, and low water species no mbuna or Aulono- the larger size of males are all likely to cara has ever been reported to practise be sexually selected traits. While larger care of free-swimming fry in the natu- size may have evolved mainly through ral habitat, but many other haplochro- advantage in combat, the other fea- mine females have been observed tures must all have arisen mainly guarding fry, including members of the through female choice. To date how- genera Copadichromis, Protomelas, ever, only the work of Hert (1991) on Otopharynx, Dimidiochromis, Hemi- egg-spot numbers and McKaye et al. taeniochromis, Nimbochromis, Fossoro- (1990) on bower size have actually chromis, and Buccochromis. Oreochromis demonstrated that females select males spp. apparently all guard free-swim- on the basis of any of these features. ming fry. Nothing is known of the pa- Bright colours are likely to attract

211 predators, long fins to slow down es- It has been suggested that differences cape and both of these features as well in the territorial sites of breeding males as construction of bowers are probably represent a form of niche separation. I energetically costly to males. It has think this suggestion can probably be been suggested that such traits have dismissed because all of the endemic evolved mainly for the purpose of spe- cichlids are polygamous maternal cies recognition, but this seems un- mouthbrooders. Each male can thus likely as this purpose could be served mate with many females, and so popu- by much less costly means, such as lations cannot be regulated by short- fairly subtle colour differences unlikely age of male territories, except in the to attract the attentions of predators. extremely unlikely event that virtually Extravagant and costly signals can no males of a particular species are able only evolve where there is competition to defend a territory even for the short to 'persuade' the receiver of the signal time required for mating. The remain- — in this case the female. Where the ing possibility is that Fryer was right interests of the sender and receiver of and Malawi cichlids violate the com- a signal are identical, as with species petitive exclusion principle. Many recognition, selection will favour a re- ecologists are now sceptical about the duction in the costliness of the signal principle. Marine ecologists whether (Maynard Smith, 1991). However, working on rocky shores, coral reefs, characteristics which evolved because or commercial fish stocks now recog- of sexual selection can certainly serve nise that apparently random processes the purpose of species recognition. have a fundamental role in community structuring. In particular, the order in Co-existence which species colonise an area may profoundly affect the resulting com- How can so many species co-exist in munities — remove the fauna and dif- Lake Malawi? Many species are sim- ferent species may get a foothold first ply found in different areas of the lake, and prevent or even facilitate other but many other species appear to be species from becoming established. wide-ranging. There is certainly some Fisheries scientists have expended segregation through habitat prefer- great efforts trying to work out stock- ences, most clearly by depth, but also recruitment relationships, but have substrate type, although again many found that recruitment — the number species are almost ubiquitous, and of juveniles surviving their first few many more certainly overlap to some weeks of life — is extremely variable extent. Some species have rather dis- and largely independent of stock size tinct diets or different means of gath- — the number or biomass of adults in ering food, and although at present the population — at least until adult there have been few incisive studies, it numbers become very low indeed. may emerge that many species co-ex- Could similar processes be operating ist sympatrically while feeding on the with Lake Malawi cichlids? We have same resources. Would this be difficult no idea. Malawi cichlids have small to explain? broods of young which are provided

212 with long-term parental care, in con- tle phylogenetic value, evolving rap- trast to the huge numbers of tiny eggs idly and unpredictably under sexual or larvae produced by all the well- selection. Similar patterns often seem studied marine fish species. No-one to reappear in apparently very distant knows what difference this makes to lineages, e.g. the yellow-white 'blaze' stock-recruitment relationships, but if in some Aulonocara, Cynotilapia, Proto- Lake Malawi follows the pattern for melas, Diplotaxodon, and Copadichromis marine systems, most of the critical — all of which have closely-related processes in community structuring species without the blaze. Eccles & may occur during the juvenile phase. Trewavas provided a generic classifi- cation in which non-breeding colour Phylogenetics was given considerable weight as a phylogenetic trait. They clearly recog- Prior to the advent of modern mo- nised a number of exceptions, such as lecular techniques, phylogenetic recon- the scale-eater Corematodus in which struction was based on the study of mimicry of host species has almost cer- anatomical characters. The most use- tainly been the reason why one species ful phylogenetic characters are often is vertically barred and the other ob- those which are not of importance to liquely-striped. This is an extreme case, the ' chances of surviving and but it indicates the general truism that reproducing, because these features are colour is often under strong selection, likely to be strongly influenced by the particularly as means of camouflage, environment and the way of life of the which in turn will be influenced by the . For example, the shape of the diet and habitat preference of each spe- jaws may be important in determining cies. To make matters worse, colour how the animal gets its food and so pattern is clearly variable, both be- animals feeding the same way will tween geographically separate popu- tend to develop similar shaped jaws, lations and also within an individual, but the method of development of the according to mood. However, it re- particular bones that go to make up the mains a useful way to allocate new jaw structures may be irrelevant to species to genera, so long as it is not their function — as long as the end re- given too much weight. sult is a good strong jawbone — and Recent developments in molecular so it can provide critical phylogenetic biology have revolutionised phylo- information. genetics. The reason is not because it Among Malawi cichlids, very few is possible to identify genes for species phylogenetically useful anatomical recognition, an extremely arduous features have been found. Most varia- task, but because it provides the large tion is in colour patterns or anatomi- set of selectively neutral characters that cal features connected with feeding phylogeneticists have always dreamt adaptations. Trophic structures are of finding. In other words, most infor- particularly likely to be the product of mation comes from the 'junk DNA' convergent or parallel evolution. Male which persists in the genes simply be- breeding colours also seem to be of lit- cause it does no harm and there is no

213 easy way for evolution to get rid of it. cies definition accepted by relatively To date, few studies have been carried few practising biologists, but there is out, and most of these have looked at not a shred of evidence that it is appli- a small number of taxa or used rela- cable (or indeed inapplicable) to Ma- tively crude techniques. Broadly, the lawi cichlids. For example, many ani- main findings are that Malawian mal species occasionally hybridise suc- haplochromines are more closely re- cessfully with other sympatric species lated to each other than to species from or at geographic boundaries with allo- Lake Tanganyika or Victoria (Meyer et patric species. Paterson's concept sim- al., 1990, Kocher et al., 1993), that the ply does not accomodate this. Obvi- ancestor of all the Lake Malawi haplo- ously, it remains quite possible that chromines, with the possible exception Paterson's definition is applicable to of Rhamphochromis (Moran et al., 1994), Malawi cichlids, but there are many was Astatotilapia calliptera or its ances- other possible species concepts which tor (Meyer, 1993), that Aulonocara, also need to be tested. Many biologists Alticorpus and at least some Lethrinops have now gone back to the idea of species are more closely related to the Charles Darwin that a species is noth- mbuna than to other Malawian haplo- ing more than a well-characterised va- chromines (Moran et al., 1994) and that riety, and the exact point at which two the chambo group is more closely re- populations diverge from varieties into lated to other 'tasselled' species from species is impossible to determine. For outside Lake Malawi, such as Oreochro- practical purposes, a species can be mis (Nyasalapia) macrochir, than to the defined as a collection of populations sympatric O. shiranus, which is related which either (i) comprises individuals to Oreochromis mossambicus and O. which do not reproduce with those of placidus (Sodsuk et al., 1995). sympatric populations and probably would not do so with those of allopat- ric populations if they were to become Species concepts sympatric or (ii) in which some degree of crossing occurs but this does not Most recent literature on African cause a loss of the overall 'distinctness' cichlids assumes that we can safely use of the populations. This definition in- Paterson's (1980) Recognition Species volves some subjectivity of judgement Concept, whereby each species can be and applies only to species with sexual determined by identification of its reproduction existing in the present. unique 'Specific Mate Recognition Sys- tem'. These are important and useful ideas, which have helped to clarify Speciation — where? thinking on some of the older species concepts, and focus research on the Leaving aside the question of exactly importance of male colour in species what causes reproductive isolation identification (e.g. Ribbink et al., 1983). between species, we can consider 6 However, it may come as a surprise to possible modes of speciation: many readers that not only is this spe-

214 Allopatric (gene flow stopped before Lake Malawi consisted of a single ba- speciation) sin (Ribbink, 1994a), so this explana- tion is inapplicable. (i) invasion of a lake by multiple lin- eages, (iii). Isolation in peripheral lagoons. (ii) splitting of the lake into multiple The classic example of this process is basins at low water levels, Lake Nabugabo, which has been iso- (iii) isolation of populations in pe- lated from for 3,500 years ripheral lagoons, and now contains five endemic species (iv) isolation of populations to habi- (Greenwood, 1974). The fundamental tat patches within the lake, difference between this and the previ- ous explanation is that small lagoons Sympatric (gene flow stopped dur- are unlikely to contain species of deep ing speciation) water, pelagic, or rocky shore habitats, but will mainly be populated by spe- (v) speciation along a cline of con- cies frequenting sheltered swampy tinuous distribution, bays. Such lagoons have almost cer- (vi) speciation within a fully mixing tainly existed around Lake Malawi and population. some still do (e.g. Lake Chiwondo), but there is no direct evidence that they have ever contained endemic species. (i) Multiple invasions. Seven distinct cichlid lineages have invaded the Lake (iv). Isolation on habitat patches Malawi basin: Oreochromis (Nyasalapia) within the lake. Mbuna have no dis- spp., Oreochromis (Oreochromis) shir- persal phase in their life cycle and are anus, Tilapia rendalli, Tilapia sparmanni, rarely seen far from rocky habitats. Pseudocrenilabrus philander, Serrano- Many species are endemic to particu- chromis robustus, and Astatotilapia lar islands or stretches of rocky shore calliptera. Yet, apart from the three Oreo- within the lake. Thus, there has been chromis (Nyasalapia) spp., all (or almost ample opportunity for allopatric all) of the cichlid speciation in Lake speciation within the rocky shore spe- Malawi appears to have taken place cies (Fryer, 1959; Ribbink et al., 1983). within the lineage descended from Very large stretches of rocky coastline Astatotilapia. Multiple invasions seem may represent a geographic barrier to unlikely to be important in Lake Ma- the movement of some sandy shore lawi, at least. fishes. Some species may range widely, but will only breed in a particular kind (ii). Isolation in multiple basins. At of habitat which may be patchily dis- low water levels, Lake Tanganyika was tributed. So habitat patchiness is prob- certainly split into three basins. This ably the single most important cause has left a clear mark on the present day of speciation within Lake Malawi, but distributions of some Tanganyikan it does not provide a complete expla- cichlids (Sturmbauer & Meyer, 1993; nation for rapidity of cichlid Konings, 1994). At low water levels, speciation. It is far more difficult to

215 identify barriers to the movement of ticular, it is now known that under cer- pelagic or deep-water species or to tain conditions, it is theoretically fea- such as Nimbochromis livingstonii which sible for sexual selection to cause sym- occupy all habitats. Previously, it may patric speciation (Turner & Burrows, have been possible to claim that these 1995), although it is not yet known ex- groups simply comprised very few actly how general these conditions are, species, but the present work indicates nor whether they apply to Malawian that deep-water and pelagic commu- cichlids. Certainly there is absolutely nities are rich in endemic species. no justification for concluding at present that all speciation in Lake Ma- (v). Speciation along a cline. I know lawi has taken place allopatrically, al- of no detailed quantitative study of though elaborate untestable stories can variation in morphology, coloration or always be produced to explain every molecules along an uninterrupted dis- past speciation event in allopatric tribution of any Lake Malawi species. terms. For example a pelagic Diplo- Konings (1995b) illustrates geographic taxodon might somehow get into a pe- variation in male colours in a number ripheral lagoon and speciate in allo- of mbuna species, but it is impossible patry without losing any of its adapta- to know if these populations are all tions to deep water. spatially isolated or if there is some gene flow between them. Speciation how? (vi). Speciation within a location. There are numerous examples of The lake is large and provides many closely-related sympatrically-occur- potential ecological niches, but ecologi- ring species in Lake Malawi. Some cal (especially trophic) specialisations workers prefer to explain all such oc- are unlikely to be the driving force be- currences as the result of secondary hind speciation, because the funda- contact between previously allopatric mental process in speciation is the es- populations (e.g. Ribbink et al., 1983), tablishment of reproductive isolation, but this is a non-parsimonious expla- which is not directly caused by adap- nation and it could equally be said that tations in trophic structures. Some re- all presently allopatric populations are searchers argue that speciation is the result of sympatric speciation fol- mainly a result of acccidental changes lowed by local of one of the in mate recognition traits resulting species pair. Most biologists find it from adaptation to the environment in easier to accept allopatric than sympat- allopatry (e.g. Ribbink, 1994b). How- ric speciation, but recent reviews ever, this theory is unlikely to be the (Bush, 1994), theoretical (Turner & Bur- whole explanation, as changes in male rows, 1995), and empirical studies coloration appears to evolve before any (Schliewen et al., 1994) indicate a grow- significant morphological change ing acceptance that geographic isola- (Greenwood, 1974; Ribbink et al., 1983; tion may not be a necessary prerequi- Turner, 1994d) and there is no evidence site for speciation in animals. In par- that male colour differences between

216 closely related species are in any way tween them (Ribbink, 1994). Likewise related to differences in their environ- in Lake Victoria, there are only 17 en- ment. As Ribbink (1986) has pointed demic species out of 11 non-cichlid out, if colour mainly serves the pur- families (Greenwood, 1994). Neither pose of mate recognition, it is likely to lake has a great diversity of endemic be under strong stabilising selection. invertebrates, but both have probably Of course it is always possible that more than 500 endemic haplochro- change in colour could occasionally mines. This is in marked contrast to result from pleiotropic effects resulting Lake Tanganyika which has 61 species from selection on other traits, but this of endemic non-cichlid fishes, 389 en- is not really consistent with the obser- demic invertebrates, and even two en- vation that colour is the trait that ap- demic reptiles, compared to 'only' pears to vary most. The only way out around 180 cichlids (Coulter, 1994). I of this dilemma that I can see is to con- would suggest that the great age of sider that species recognition is prin- Lake Tanganyika and its regular split- cipally an effect of sexual selection, re- ting into 3 basins during low water lev- sulting from choice of mates within a els has meant that virtually any taxon species. getting into the lake has undergone Theoretical studies indicate that speciation. Thus, we are really looking sexual selection is very likely to lead for an explanation for the richness of to speciation and recently several good the haplochromine species flocks of explanations have been given for the Lakes Malawi and Victoria. variability of traits under sexual selec- Theories based on habitat fragmen- tion (see Turner, 1995b, 1996). Ma- tation and low dispersal capabilities of lawian cichlids do seem to be subject haplochromines have to contend with to strong sexual selection. Future ex- (i) the lack of speciation in many other perimental work should concentrate taxa in many other places which exist on this possibility. There is no evidence as geographically isolated populations that Malawian cichlids are unusually (ii) the apparent lack of habitat barri- prone to develop chromosomal or ers to non-rocky shore species. Expla- physiological reproductive incompat- nations based on morphological inno- ibility, indeed they seem to hybridise vation permitting invasion of empty readily in aquaria. niches have to accomodate (i) the lack of evidence that morphological inno- Speciation — why haplochromine vation leads to exploitation of differ- cichlids? ent niches (ii) the similarity of mor- phology of many sibling species (iii) Probably the most intriguing puzzle the likelihood that reproductive isola- about the speciation process in Lake tion is not caused by morphological Malawi is that the haplochromine differentiation. Theories emphasising cichlids alone have speciated so exten- the role of sexual selection have the sively. There are 10 other families of difficulties that (i) Oreochromis spp. are fishes in Lake Malawi, but these have also strongly sexually selected, but do produced only 25 endemic species be- not appear to speciate rapidly (ii)

217 haplochromines in riverine habitats introduced forms does not seem to be have not diversified to the same extent occurring, although it may be too early as those in lakes. Given the present to say for sure. Unfortunately, the best state of knowledge, it would be fool- data on population sizes (Ribbink et al., ish to strongly support any one of these 1983) comes from counts of territorial theories at the expense of the others. males and as we have already seen, this is likely to be largely irrelevant to a The aquarium trade population's reproductive capacity. It is possible that the main detrimental Malawi cichlids are popular in the effect of introduced species may be in aquarium trade. Many species are bred competition between juveniles for fry in small numbers in aquaria in Europe refuges (Trendall, 1988). and North America, mostly by ama- Overall, the aquarium trade brings teur or semi-professional breeders. large amounts of foreign currency to Large-scale breeding in ponds in SE Malawi (and increasingly Tanzania Asia and the southern USA supplies and Mozambique), both through fish large numbers of fishes, but often these sales and tourism and appears to do are artificially selected mutants, such little, if any, harm. as albinos, or hybrids. Neither are Where might the aquarium trade popular among discerning aquarists expand in the future? Export from Tan- specialising in African cichlids. Be- zania has been flourishing in the last cause of the dangers of hybridisation, decade, although there is the worry- but also because of the demand for ing possibility that competition be- novelty, wild-caught fishes are much tween exporters may result in in demand and command premium overexploitation or translocations. prices. While there has been some con- Stuart Grant has now started to collect cern that wild populations may be en- in Mozambique. Limited and tightly dangered by overexploitation (Ribbink controlled collection from the southern et al., 1983), there is no real evidence part of the lake, including the Lake that this has happened yet and at Malawi National Park could be possi- present the risks are small because of ble. Trade in aquarium fishes has been the small numbers taken and because concentrated on mbuna, which are of- of recent development of lakeshore ten colourful as juveniles as well as pond breeding at Stuart Grant's facil- adults, remain relatively small and are ity in Senga Bay, Malawi. However, abundant on rocky shore areas where there is no question that the aquarium they are easily seen and caught by trade has had unfortunate conse- divers. Over the last 10 years or so quences, through the introduction of Aulonocara and Copadichromis species species from the north of the lake have become increasingly sought after, (Nkhata Bay and Likoma) to the south, as have some of the piscivores such as around Cape Maclear. Exporters are Nimbochromis. Deeper water species now aware of these risks, and future and sandy shore species have not been translocations seem less likely, and the exploited to the same degree as com- feared exclusion of endemic species by parable species from Lake Tanganyika,

218 perhaps because they are more diffi- appears that until the 1980s, the num- cult to catch or to identify, but perhaps bers of fishermen and the development because of the large resource of rocky of their technology did not permit shore species. Some of the species dis- overexploitation, except perhaps of a cussed in the present work could be few vulnerable stocks such as the caught alive and may make desirable cyprinid Labeo mesops which was eas- aquarium species, some may be con- ily trapped in large numbers as it mi- fined to waters too deep for safe ex- grated upriver to spawn (FAO, 1976). ploitation. Thus, development aid was wisely in- vested in the exploration of un- exploited stocks, the development of Fisheries management fishing techniques and infrastructure (e.g. jetties, processing equipment) and For the majority of people living in the subsidy of fishermen, particularly Malawi and the lakeshore regions of trawl operators. However, it is now Tanzania and Mozambique, fish are clear that most major fisheries are fully important principally as a source of exploited, and in some cases seriously food and income. Under free economic overexploited (Turner, 1994e, 1995). At development, all fish stocks must in- present, the governments of the evitably be overexploited. What little lakeshore nations cannot afford ad- evidence there is suggests that catch- equate enforcement or education cam- per-unit effort (CPUE, e.g. catch per paigns and do not really have access day with a particular kind of fishing to adequate data to know if such meas- technique) of fisheries aimed at large ures as are undertaken have proved high-value species has been steadily effectual. Little can be done to reverse declining, leading to calls for mesh size the decline, unless external aid money restrictions etc.. However, decline in is invested in fisheries management, CPUE is an inevitable consequence of rather than fisheries expansion. Fish- increasing fishing effort, even while eries do not regulate themselves, and yields are well below optimal levels. It in the absence of alternative income- is impossible to assess the state of ex- generating activities, more and more ploitation of a fishery without either a fishermen are likely to join the indus- long series of catch and effort data or a try, further accelerating the decline. detailed knowledge of the population Most fisheries in Malawi involve structure of the species. Neither of small-meshed unselective gears, such these are available for most fisheries as trawls and seines, and all species are or species. The situation is further com- taken for food. It has already become plicated by the huge number of spe- clear that heavy exploitation by these cies exploited, the great variety of fish- gears has led to reductions in popula- ing techniques in use and the fact that tions or even the local elimination of no-one really knows which techniques many of the larger cichlid species. This catch which species, let alone how has been documented in Lake much of each is being caught. Malombe and the SE Arm of Lake Leaving aside these difficulties, it Malawi (Turner, 1994e; Turner et al.,

219 1995; Tweddle et al., 1995) and may fishing is limited to relatively innocu- have occurred in other areas, which ous techniques such as traps or an- have not yet been studied. gling, or by convincing the fishermen that they are genuinely realising some Conservation practical benefit from their restraint. Malawi already has reserves for the At present it has not been proven that preservation of rocky shore fishes, but any species of Lake Malawi cichlid has these are not at present endangered, gone extinct, but extinction of aquatic except perhaps by translocations of animals is notoriously difficult to species which are probably irrevers- prove. Many species have not yet been ible. Rocky shore species are not eas- described, and there is presently no ily caught in large numbers as seines information about the population sizes and trawls cannot operate over rocky of most species. The only attempts at bottoms. population estimation have been for In the long term, pollution or species the tilapiines in the SE Arm and Lake introductions may prove the greatest Malombe (FAO, 1993; Turner, 1995) danger to Lake Malawi's unique fishes, and the experimental trawl surveys of but at present no major industrial de- the SE Arm demersal haplochromines velopment has taken place on the (Turner, 1994e; Turner et al., 1995). Both lakeshore and no species have been of these studies have indicated sharp introduced from outside the lake. Fish- declines in populations of larger ing represents a real and continuing cichlids, and hint at local . danger, particularly to the offshore spe- In both cases, the only plausible expla- cies and sandy shore species. This is nation is that populations have been the principal problem which conserva- reduced by fishing. tionists, fisheries scientists, and policy- At present fisheries are managed ac- makers must address if irreversible cording to the principle of maxi- damage to the unique ecosystem of misation of sustainable yields, not spe- Lake Malawi is to be prevented. cies conservation. Indeed, it is quite possible that maximum fishery yields can best be obtained from practises that are likely to lead to the extinction of large slow-maturing species, especially predators. Although much well-mean- ing rhetoric is expended about the ben- efits of conservation to indigenous peo- ples, in truth there may be no economic reason for fishermen to conserve these species. If citizens of the developed world wish to ensure that these spe- cies survive, it will be necessary for them to pay it, either through the crea- tion of biodiversity reserves where

220 Glossary

Definitions of technical terms used trawlers, freezer plants, echo sounders. in this book are given, together with Artisanal fishermen in Malawi are com- other terms which may often be seen mercial fishermen — they generally sell in the technical and semi-technical lit- most of their catch, rather than use it for their own families' consumption. erature on African cichlids. I have also Aufwuchs. Epilithic algae (q.v.) and mi- included brief synopses about some of cro-invertebrates (q.v.). the researchers whom I have fre- Axelrod, Dr. Herbert R. Founder and head quently mentioned in the text. Inclu- of the Tropical Fish Hobbyist publishing sion reflects their overall published empire. Made several collecting trips to contribution to date to the study of the Malawi in the early 1970s, and collected fishes of the offshore region, rather offshore cichlids from a catch by the fish- than representing any personal assess- eries research trawler Ethelwynn Trewa- ment of individual scientific merit! vas. Bauplan. Overall body form. Active. State of fish gonads (reproductive Beach Seine. A net which is set in a semi- organs) indicating that eggs or sperm circle and pulled onto the shore. are developing. Benthic. Fish, invertebrates or plants liv- Adaptation. The process of evolutionary ing on the bottom. change of any feature of an organism Bicuspid. Used about teeth, meaning that which improves its overall chances of they have two points or 'cusps'. survival or reproductive success. Char- Biogeography. The study of the distribu- acters which evolve through sexual se- tion of animals and plants in relation to lection (q.v.) are sometimes wrongly their evolution. considered to be non-adaptive because Biomass. Total weight of living organisms they may interfere with survival, but any in an area or a sample. adaptive change may have both posi- Bivalves. Molluscs with two shells, such tive and negative effects on different as clams. Several species are found in functions. the mud of Lake Malawi. Allopatric. Found in different places. Allo- Bower. Structure built by an animal to help patric species are those which presently it attract mates, but not used for rearing have non-overlapping distributions. Allo- of young. Sometimes referred to as a patric speciation is a hypothesis that a spawning platform, or more loosely, species was divided into two non-over- nest. lapping populations which subsequently Bottom Trawl. A large net pulled along evolved into separate species. The ma- the bottom. In Malawi, generally used jority of biologists contend that this is to refer to a trawl (q.v.) operated from a the only mode of speciation possible. single large vessel. Annelids. Segmented worms. Branchiostegal Membrane. A membrane Artisanal Fishery. A fishery which does forming the lower part of the mouth cav- not employ expensive equipment such ity. It is often brightly coloured and can

221 be extended for display in some cichlids. ure of primary productivity (q.v.). Also known as the 'gular membrane'. Cichlid. Fishes of the family Cichlidae, Burgess, Dr. Warren E. Writer on aqua- one of the major families of freshwater rium fish for Tropical Fish Hobbyist Pub- fishes in Africa, and South and Central lications. Described several cichlid spe- America and Madagascar. A few spe- cies from specimens collected by Axel- cies are also native to southern Asia and rod (q.v.). North America. All species have a dor- Bycatch. Fish which are caught and usu- sal fin with both spiny and soft rays, pha- ally killed, but are not the main target of ryngeal jaws, and a closed swimbladder. the fishermen, usually because they are As far as is known all species practise of low economic value. Around Lake complex courtship behaviour and long- Malawi all bycatch is used for human term parental care. consumption. Cladocera. Small aquatic crustaceans, Caudal Peduncle. The fleshy part of the with paired shells, such as Daphnia. tail. Cladistics, Cladistic Taxonomy. Taxo- Cephalic Pores. Sensory pores of the lat- nomic study which aims to base classi- eral line system (q.v.), expanded in fication on phylogeny (q.v.), in which some genera. each taxon is intended to represent a Ceratobranchial Gillrakers. Gillrakers on monophyletic (q.v.) group. the lower arm of the gill arch. Lower Clariid. Catfishes of the family Clariidae, Gillrakers. native to Africa and Asia. Generally large Ceratophyllum. A common macrophyte fishes with strong pectoral fin spines and in shallow water of Lake Malawi. Noted an air-breathing organ on the roof of the for its tough skeleton of silica which gill chamber. Clariid catfishes form the makes it virtually inedible to most second largest species flock in Lake aquatic animals. Malawi. Chambo. Tilapiine cichlids of the genus Cleithrum. The bony support for the pec- Oreochromis, subgenus Nyasalapia, toral and pelvic fins. endemic to the Lake Malawi system. Competition. The situation where two or Chambo Seine. A seine net (q.v.) used to more organisms require access to some catch chambo (q.v.), usually with rela- resource, and where the survival or re- tive large meshes (75-100mm). These productive success of one may be low- nets may be up to 1.5km long in south- ered by the other's gaining access to ern Lake Malawi. that resource. Competition need not in- Chaoborus. A fly with aquatic predatory volve aggression, and may occur be- larvae, which are pelagic for most of tween organisms that never meet. Com- their life history. petition between conspecifics (q.v.) is Chironomids. Flies with aquatic larvae widespread and is one of the driving which live buried in the sediment. forces of evolutionary change, but the Chirimila. A small 'D'-shaped net operated role of competition between species is offshore and near the surface, from ca- more controversial. noes or small plank boats, mainly to ex- Conspecific. Belonging to the same spe- ploit usipa (q.v.) or utaka (q.v.). cies. Chlorophyll-a. Chlorophyll is a coloured Copepods. Small aquatic crustaceans, pigment used by plants to trap the en- such as Cyclops. ergy from sunlight. Concentration of one Countershading. Dark on top, and lighter class of these pigments- chlorophyll-a- coloured on the flanks and belly. This is often used as an approximate meas- pattern counteracts the natural tendency

222 for the lower part of the fish to seem con- other molecules used to construct their spicuously dark because it is shadowed. bodies and ultimately their behavioural It is the camouflage pattern typical of traits. Recent developments in analys- open water fishes. ing the structure of the molecule have CPUE, Catch per-unit-effort. Term used by vastly increased the number of traits fisheries scientists meaning the weight available for phylogenetic reconstruc- of fish caught per unit time with a par- tion. ticular unit of fishing effort- i.e. a par- Eccles, David H. Fisheries researcher for ticular gear and boat used in the same the ODA during the 1960s and early manner. Used to give an indication of 1970s, mostly based at Monkey Bay. the biomass (q.v.) of a species at a par- Carried out important limnological stud- ticular time. ies, fisheries surveys and together with Cyanophytes. So-called 'blue-green al- Lewis (q.v.) taxonomic work on Lethri- gae', they lack the nucleus and cell nops. After leaving Malawi in 1978, organelles of higher plants and animals worked on fish taxonomy at the JLB and are actually more properly regarded Smith Institute of and the as bacteria. The tough cell-walls of the British Museum of Natural History be- blue-greens means that the can only be fore moving to Australia. digested by fish with very acidic stom- Egg-spots. White or yellow spots on the ach secretions. anal fin of haplochromine cichlids (q.v.), Cyprinid. Fish of the family Cyprinidae, which probably evolved to increase fer- the dominant family of freshwater fishes tilisation success in mouthbrooding in Europe, North America and much of cichlids. In some species at least, this Asia. Abundant in Africa rivers, cyprinids function has been supplanted by a role are a much less significant part of the in sexual selection. fish communities in large lakes. Char- Endemic. Of an animal or plant species, acterised by the presence of pharyngeal found only in a particular place, where it jaws, lack of jaw teeth, soft dorsal and is assumed to have originated. In evo- anal fins. Malawian species do not prac- lutionary biology, a species which is na- tise parental care and most spawn in tive to a particular place, but also found the streams and rivers flowing into the elsewhere, is said to be 'indigenous'. main lake. Ephemeroptera. Mayflies. A group of in- Demersal. Fish living on or near the bot- sects with aquatic juveniles or nymphs. tom. Epilithic. Growing on the surface of rocks. Dental Arcade. The arrangement of teeth Epiphyte (adj. Epiphytic). Growing on the in the jaws. surface of plants. Detritus. Decaying organic matter lying FAO. The Food and Agriculture Organi- on the bottom, together with the micro- sation of the United Nations, which has organisms that feed on it. supported a number of projects on Lake Diatom. Algal cells with an external skel- Malawi, notably on demersal trawl fishes eton of silica. Numerous pores in the (completed 1976), pelagic ecosystem & skeleton mean that the internal contents fishes (1982) and tilapias (1992). are easy digested by fish. The most con- Fauna. The animal species of a particular spicuous component of the phytoplank- area. ton (q.v.) and the dominant large algae. Fryer, Dr. Geoffrey A. British freshwater DNA. Deoxyribonucleic acid, the molecule biologist who worked for the JFRO (q.v.) which is used by all animals, plants and on Lake Malawi in the 1950s. Notable bacteria to code for the proteins and for his monograph on the mbuna, par-

223 ticularly of the Nkhata Bay region and name is retained by the holotype, un- his classic book with T.D. Iles (q.v.). Sub- less it is considered to belong to a pre- sequently worked for the Freshwater viously described species. Biological Association at Windermere, ICEIDA. Icelandic International Develop- England, principally on freshwater ment Agency, which is presently jointly crustacea, but has continued to write funding fisheries development and re- reviews on evolution and conservation search on Lake Malawi, along with the of African fishes. World Bank. Fyke Net. A kind of trap fixed to the bot- Interorbital Width. The shortest distance tom with 'wings' or fences to herd fish between the eyes across the bony roof into the net mouth. of the skull. Gastropods. Molluscs with a single shell. Iles, Dr. T. Derrick. British fisheries biolo- Snails. gist working with the JFRO (q.v.) on GEF. Global Environment Facility of the Lake Malawi during the 1950s. Most no- World Bank, an organisation set up to table for studies on the taxonomy and promote environmental protection. Sup- population biology of the utaka and for porting research and conservation on his classic book with G. Fryer (q.v.). Lake Malawi from 1995. Subsequently a distinguished marine Gene Flow. The exchange of genetic ma- fisheries scientist based in Canada. terial (DNA) between populations of Jackson, Peter B.N. Director of the JFRO sexually reproducing organisms which (q.v.) during the 1950s. Notable for his results from their interbreeding. annotated checklist of the fishes of Lake Genus. A taxonomic group containing one Malawi, and for taxonomic work on the or more species. The principal taxo- clariid catfishes. nomic division above the level of the JFRO. Joint Fisheries Research Organi- species. The generic name forms the sation, a British Government-funded re- first part of the formal latin name. search organisation which for a few Gillnet. A stationary net used to capture years in the 1950s carried out research fish by entanglement. on the fishes of northern Lake Malawi. Gillrakers. Bony projections from the front It also worked elsewhere in Africa. of the gill arch, used to retain food items Junior Synonym. A species name ren- in the mouth cavity. dered invalid, when it is subsequently Handline. Hook and line, generally baited. discovered that a different name had Haplochromine. A subdivision of the fam- previously been applied to the same ily Cichlidae, in some works given the species in a valid description. formal status of a , it includes all Kambuzi Seine. Small-meshed beach the endemic cichlid genera of Lake Ma- seine (q.v.) mainly used to catch small lawi, plus many from Lakes Victoria and haplochromine cichlids (kambuzi), juve- Tanganyika, as well as riverine genera nile chambo (q.v.) and usipa (q.v.). such as Astatotilapia and Serrano- Konings, Dr. Ad. Self-employed author, chromis. All known species are mater- publisher, film-maker and diver special- nal mouthbrooders. ising in aquarist literature on cichlid Holotype. A specimen selected in the fishes. Notable for superbly illustrated original description of a species as the guides to Lake Malawi cichlid fishes and holotype or type. In a subsequent revi- taxonomic work on various inshore spe- sion of the species, if the original type cies. Formerly a biomedical researcher. series in found to include members of Lectotype. A holotype was often not des- more than one species, the species ignated in earlier descriptions. A

224 lectotype is a specimen selected by a ecology. later worker to serve the same role as a Macrophyte. Large plants, usually higher holotype. plants with complex multicellular struc- Lappets. The tip of the dorsal fin mem- ture, but also some larger algae. branes, which are generally separated Macro-invertebrate. Larger inverte- on the spinous part of the fin. brates, such as molluscs, crabs and in- Lateral Line. A sensory system used by sect larvae. fish to detect pressure waves generated Maternal care. Parental care by females by objects in its vicinity. The most promi- only. nent feature is a series of pored scales Mbuna. Small haplochromines (q.v.), prin- along the midline of the flanks, but also cipally found on rocky shores. including sense organs on the head. McKaye, Prof. Kenneth R. US behavioural Laterally Compressed. Flattened from ecologist. Carried out numerous stud- side-to-side. ies of Lake Malawi cichlids during the Lek. An aggregation of males. Each male 1970s and 80s, latterly in collaboration defends a territory in which it mates, if with J.R.Stauffer (q.v.) and others. Has given the opportunity. No parental care subsequently concentrated more on his or other resources of use to the female first study area, Nicaragua. Notable for are provided by such males. his ideas on sympatric speciation and Lewis, Dr. Digby S.C. British fish biolo- sexual selection. gist, employed by the ODA to work on Melanin Pattern. The basic underlying biology of commercial fishes in south- pattern of dark markings, generally more ern Lake Malawi during the 1970s, col- clearly seen in female and immature laborating with Eccles (q.v.) on tax- fishes than in sexually active males. Re- onomy of the Lethrinops species. Later garded by Eccles & Trewavas as of key the leader of a WWF-funded team which phylogenetic significance. prepared the first guidebook to the Lake Mental Process. A bony knob on the un- Malawi National Park. Subsequently derside of the chin. worked on fisheries of Lake Kariba, in Meristics. Counts of scales, fin rays, Zimbabwe. gillrakers etc.

Linf. Length at infinity (usually spoken as Micro-invertebrate. Smaller aquatic in- 'El-infinity'). Fisheries biology term for vertebrates, such as cladocera (q.v.), asymptote of the length when fitted as copepods (q.v.) and ostracods (q.v.). a curve against age. Can be thought of Dominate the zooplankton (q.v.) but also as the approximate average maximum numerous in the sediments. size. Midwater Trawl. A trawl (q.v.) net set to Longline. Hook and line fishing gear in catch fish off the lake bottom. On Lake which a number of hooks are attached Malawi, experimental trawl surveys to a float which is left baited and unat- have occasionally used fully pelagic tended. Generally used to catch catfish. trawls in which no part of the gear rests Lowe (later Lowe-McConnell), Rose- on the bottom, but the commercial mary H. British fish ecologist. In the midwater trawl fishery is actually a semi- 1940s, carried out a single-handed pelagic trawl where the otter boards rest study of the tilapias and other commer- on the bottom, but the net is set to float cial fish mainly in southern Lake Malawi. with the foot rope a few metres clear of Subsequently worked in Uganda, S. the substrate. The commercial gear America and elsewhere, and author of mainly catches ndunduma (q.v.), several classic textbooks of tropical fish chambo (q.v.), ncheni (q.v.) and utaka

225 (q.v.). Nomen Nudum. A species name used in Molariform. Of teeth, expanded and flat- scientific literature without it having been tened for crushing hard food items such previously applied to a type specimen as molluscs. A number of Lake Malawi in a formal description. A nomen nudum haplochromines have Molariform teeth remains available for use by later tax- on the pharyngeal bones, which are onomists who may wish to apply it to broader and have strong muscular at- the same or another species. tachment than usual. It is likely that a ODA. Overseas Development Administra- great deal of this variation is the result tion of the UK, which has supported nu- of phenotypic plasticity (q.v.). merous projects on Lake Malawi, nota- Molluscivore. A predator of molluscs. bly the ODA/SADC Pelagic Resources Monophyletic. A taxonomic group con- Project (completed 1994) and the cur- sisting of a number of lineages de- rent Pelagic Cichlid Biodiversity Project scended from a single common ances- (Ncheni Project), as well as stationing tor. In cladistic taxonomy, there is the several ODA-funded fisheries officers in further restriction that none of the line- Malawi (see Eccles, Lewis, Tweddle). ages descended from the common an- Oliver, Dr Michael K. US fish taxonomist cestor are excluded from the group. who carried out extensive systematic Mormyrid. Fish of the family Mormyridae. studies of Lake Malawi cichlids in the Mormyrids lack spiny rays on the fins, course of a PhD study completed in and all species generate electric pulses 1984. This work included several spe- which are used to detect objects, includ- cies descriptions which he did not pub- ing prey, as well as for communication. lish, but subsequently appeared in the The eyes are generally reduced and the work of Eccles & Trewavas, sometimes fishes are primarily nocturnal. Several in modified form. species occur in Lake Malawi. Operculum. The bony gill-cover plate. Morphometrics. Body measurements Ostracod. A small benthic crustacean used in taxonomic studies. which is enclosed in a pair of shells. Mouthbrooder. A fish which carried its Paralectotype. Specimen of a type se- eggs and/or fry in its mouth. ries where a holotype (q.v.) was not origi- Najas. A prickly macrophyte (q.v.) which nally designated, remaining after the is generally not rooted to the bottom. designation of a lectotype (q.v.) by a later Nape. The upper part of the head in front worker. of the dorsal fin. Paraphyletic. A group of taxa descended Natural Mortality. Term used in fisheries from a single common ancestor, but one biology. The rate of deaths per unit time in which some of the lineages de- in a particular age class of an un-ex- scended from that ancestor are ex- ploited population. In simple fisheries cluded. models (single-species models) it is as- Paratype. Specimens other than the holo- sumed to be unaffected by fishing. type (q.v.) included in an original spe- Ncheni. Pelagic predatory cichlids of the cies description. genus Rhamphochromis. Large indi- Parsimony. The assumption that the sim- viduals also known as 'batala' in some plest explanation is most likely to be true areas. in the absence of positive proof. A ba- Ndunduma. Pelagic or deep water sic assumption of all science, and for- cichlids of the genus Diplotaxodon. The malised in cladistic taxonomy, to the name of the Malawi Fisheries Depart- extend of its incorporation into compu- ment's latest research trawler. ter programmes used to help determine

226 phylogeny (q.v.). (q.v.), at least two species of which are Pair Trawl. A trawl (q.v.) pulled between found in Lake Malawi. It forms exten- a pair of boats. The most common com- sive beds of plants up to 2m tall in the mercial trawl gear on Lake Malawi, gen- southern part of the lake, where it is the erally used in shallow waters less than preferred haunt of most epiphyte feed- 40m deep. ers. Paedophage. A predator feeding on the Polymorphism. The occurrence of two or eggs and fry of other fish. Generally re- more forms within a species. More stricted to those preying on young of strictly, the morphs should occur sym- mouthbrooding cichlids after they have patrically, be clearly distinct (discontinu- been taken into the mouth, but before ous), be genetically-determined and the they have been released to feed inde- rarest type should not be maintained by pendently. mutation alone. Differentiation into two Pharyngeal Bones. The bones of the sexes is not normally considered to con- pharyngeal jaw apparatus, possessed stitute polymorphism. by a number of fish groups including Polyphyletic. A taxonomic group de- cichlids, catfish and cyprinids. In scended from two or more distinct an- cichlids, the upper pharyngeal bones are cestral taxa. separate, while the lower elements are Premaxillary Pedicel. The upper exten- fused to form a single bone. Earlier sion of the premaxilla. In cichlids this is cichlid taxonomists, e.g. Trewavas (q.v.) visible as a bulge on the upper part of considered the pharyngeal bones to be the snout which slides back and for- of particular taxonomic significance, but wards as the jaws are opened and more recent work indicating extensive closed. phenotypic plasticity (q.v.) casts doubt Preorbital Bone. A flattened bony plate on this. under each eye extending to the upper Phenotypic Plasticity. Variation in the lip. Also called the lachrymal bone. structure or behaviour of an organism Primary Productivity. The rate of produc- induced by differences in the environ- tion of plant biomass, which in turn lim- ment rather than genetic differences. its the potential productivity (q.v.) of ani- Phylogeny. The evolutionary history of a mals. group, particularly the order of diver- Productivity. The production of biomass gence of various lineages generally rep- (q.v.) by a species, or group of species resented by a 'tree'. the most important in a given area in a given period of time. traits for determining phylogenetic rela- It can be used to cover the potential rate tionships are generally those which ei- of production of resources, assuming ther have no adaptive value or are indi- they are not subjected to predation or cations of different ways of construct- harvesting by man. ing the same structure. Redescription. A description of a previ- Phytoplankton. Microscopic plants, gen- ously-described species which may or erally algae and cyanophytes (q.v.), may not involve additional specimens, which float freely in the water. intended to make it easier to identify the Piscivore. A predator of fish. Generally species or to provide information which used in connection with predators of may be helpful in elucidating phyloge- adult or subadult fish which kill their prey, netic relationships. The authority cred- i.e. excluding fin and scale eaters and ited with describing the species remains paedophages. with the original author. Potamogeton. A rooted macrophyte Ribbink, Dr. Anthony J. South African

227 ethologist and conservation biologist, Darwin in 1859 to explain extreme ex- worked as director of a research project amples of sexual dimorphism (q.v.), but on inshore aquarium fishes (mainly has recently become accepted by the mbuna) of Lake Malawi in the 1970s and majority of evolutionary biologists. 80s, then at the JLB Smith Institute of Speciation. The formation of a new spe- Ichthyology, Grahamstown, S. Africa cies. It is generally restricted to cases where he continued to be involved with where one species splits into two, rather Lake Malawi. Manager of the GEF than cases of change within a lineage. biodiversity project on Lake Malawi from Species. The fundamental unit of biologi- 1995. Renowned for classic monograph cal classification, the exact definition of on the mbuna published in 1983. An ad- which remains controversial. In practice vocate of Paterson's recognition species a species can be considered as a popu- concept. lation or group of populations which Ring Net. A net operated offshore, being rarely interbreed with other such groups first cast in a circle, before the bottom is or would rarely interbreed if they were closed by pulling on a rope. On Lake not spatially separated. Some biologists Malawi, these nets are operated by a consider that species, but no other taxo- pair of boats and are only used in the nomic division represent a biological re- SE Arm. ality, while others consider them to be Ripe. Of fish reproductive organs, at a arbitrary man-made conveniences. stage where a fish will shortly be capa- Spent. Used of fish reproductive organs, ble of spawning or fertilisation of eggs. when reproduction has been recently SADC. Southern African Development completed. Community, an organisation formed to Standard Length. Body length excluding co-ordinate states of the region. Partici- the caudal (tail) fin. This is the preferred pated in the co-ordination of the ODA- taxonomic length measurement, be- funded pelagic resources project and cause damage to the tail can cause in- the present GEF-funded biodiversity accuracies. project. Stauffer, Prof. Jay R. US ichthyologist Secchi Disc. A black and white disc of based at Pennsylvania State University standard size which is lowered into the who has worked on a number of stud- water. The point at which it can no longer ies of Lake Malawi cichlids since the be seen from the surface is a crude 1980s, generally in collaboration with measure of water transparency (q.v.). K.R. McKaye (q.v.). Although principally Sediment. Organic and inorganic mate- concerned with inshore fish, he has rial of small particle size resting on the made notable contributions to the tax- bottom. onomy of a number of deep-water Sexual Dimorphism. Differences be- groups and to studies of sexual selec- tween male and female of a species. tion. Generally confined to external aspects Stern Trawl. A trawl (q.v.) net pulled be- not directly concerned with egg-laying hind a boat, generally used to designate or fertilisation. the single-boat bottom set trawls used Sexual Selection. The process whereby in deep water on Lake Malawi. traits evolve because they are favour- Submarginal Band. A band of dark pig- able in competition over mates, either ment on the dorsal fin lying below the through fighting or mate choice. This lappets or fin margin. mechanism has been controversial Subspecies. A geographic variety of a since it was first proposed by Charles species thought sufficiently distinct to

228 warrant a formal name. Some taxono- held up by floats and the bottom pulled mists disapprove of the term. down by weights. In the centre is a long Sympatric. Found in the same place. 'bag'. This is the main gear used by Sympatric species presently have over- large-scale fishing operations on Lake lapping distributions. Sympatric Malawi. speciation is the hypothesis that during Trewavas, Ethelwynn. British fish taxono- the process of speciation the distribu- mist, who worked at the British Museum tions of the two speciating populations of Natural History from the 1920s to the overlapped to some extent. Many biolo- 1990s. Although working on a wide va- gists contend that this is impossible. riety of fish groups, she published many Syntype. One of a series of type speci- notable works on the tilapiines, culmi- mens of which no holotype (q.v.) has nating in her classic monograph in 1983. been designated. She published brief abstracts of descrip- Systematics. The classification of organ- tions of many Malawian haplochromines isms, emphasising the determination of in 1931 and 1935, eventually publish- phylogeny (q.v.). ing full descriptions, along with a new Taxon, (plural= taxa). A group of individu- generic scheme, in 1989 (with Eccles, als belonging to any level of taxonomic q.v.). In 1939, she participated in a field division. study on the fish and fisheries of Lake Taxonomy. The science of naming and Malawi. Fisheries research trawler was classifying organisms. named after her. Terminal Mouth. The condition where the Trophic. Relating to feeding or diet. mouth lies at the end of the snout. Trophic structures are those such as Tilapia/ Tilapiine. A subdivision of the jaws, teeth, pharyngeal bones (q.v.) and family Cichlidae, sometimes given the gillrakers (q.v.) involved in food collec- formal status of a 'tribe'. The principal tion and processing. genera are Sarotherodon, Oreochromis Turbidity. Cloudiness of water. Low trans- and Tilapia, the latter two with repre- parency (q.v.). sentatives in Lake Malawi. The Ma- Turner, Dr. George F. UK fish and fisher- lawian species are relatively large and ies biologist. Author of the present book. largely herbivorous. No relation to J.L. Turner (q.v.). Total Length. Body length including the Turner, Dr Jerry L. US fisheries scientist, caudal fin. Preferred length measure- manager of the FAO (q.v.) projects on ment for fisheries biology, because it is demersal fisheries (completed 1976) quicker to perform. and pelagic ecosystems (1982). Transparency. A measure of how far light Tweddle, Denis. UK fisheries scientist, penetrates the water. Waters with high employed on a variety of projects in planktonic productivity (q.v.) generally Malawi in 1970s-90s, including FAO have low transparency, but this may also demersal fish project (completed 1976) be caused by disturbance of sediments and various ODA-funded work. Notable by wind or by material carried into the for compilation of fisheries catch statis- lake from rivers during periods of high tics from 1976 to 1990 and for rainfall. biogeographic work on non-cichlid Trawl. A net pulled behind a boats (or be- fishes, presently being pursued at the tween two boats) with long 'wings' to JLB Smith Institute of Ichthyology, South herd the fish, usually held open by 'ot- Africa. ter boards'- heavy plates which normally Type. The holotype (q.v.) or lectotype rest on the bottom. The top of the net is (q.v.).

229 Type Locality. Location where the type (q.v.) was collected. In early works, of- ten simply given as 'Lake Nyasa'. Type Series. All of the specimens used in the original description, i.e. holotype plus paratypes, lectotype plus paralectotypes, syntypes. Type Species. Species to which a generic name is permanently attached and thus having the same status for a generic name as the holotype has for a species name. Unicuspid. Of teeth, having a single point, or cusp. Also called 'simple' or 'conical'. Usipa. A small pelagic cyprinid (q.v.) Engraulicypris sardella, endemic to Lake Malawi. Utaka. Zooplankton (q.v.)-feeding cichlids (q.v.) of the genera Copadichromis and Nyassachromis, especially those caught near rocky reefs using the chirimila (q.v.) net. Sometimes mistakenly thought to be pelagic, these fishes are generally found less than a couple of metres above the substrate. Vallisneria. A rooted macrophyte (q.v.) which produces strap-like leaves from a single growing point at the base. Rarely exceeding 20cm in height, it forms extensive beds in shallow water. Ventral Profile. The outline shape of the lower surface of the fish when viewed from a lateral aspect. Voucher Specimens. Specimens depos- ited at a museum to act as reference material for the verification of published identification of species reported in a field study. World Bank. An international financial or- ganisation providing loans on generally favourable terms for development pro- grammes. Presently supporting fisher- ies expansion work on Lake Malawi. Yield. Amount of fish caught in a particu- lar time, usually annually. Zooplankton. Small animals floating freely in the water.

230 References

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231 ECCLES, D.H. & LEWIS, D.S.C. (1981) Mid- KONINGS, A. (1989) Malawi Cichlids in Their water spawning in Haplochromis chryso- Natural Habitat. Verduijn Cichlids, Zeven- notus (Boulenger) (Teleostei, Cichlidae) huizen, Holland. in Lake Malawi. Env. Biol. Fish. 6, 201- KONINGS, A. (1990) Cichlids and all the other 202. fishes of Lake Malawi. TFH Publications, ECCLES, D. H. & TREWAVAS, E. (1989) Ma- Neptune City, New Jersey. lawian Cichlid Fishes: A Classification of KONINGS, A. (1990) Descriptions of six new Some Haplochromine Genera. Lake Fish Malawi cichlids. Trop. Fish Hobbyist, 38, Movies: Herten, Germany. 110-129. FAO (1993). Fisheries management in KONINGS, A. (1991a) Copadichromis sp. south-east Lake Malawi, the Upper Shire 'virginalis Gome' p.46 in Konings, A. (ed) River and Lake Malombe. CIFA Techni- The Cichlids Yearbook, Volume 1, Cichlid cal Paper 17 106pp. FAO, Rome. Press, St. Leon-Rot, Germany. FRYER, G. (1959) The trophic relationships KONINGS, A. (1991b) Copadichromis sp. 'vir- and ecology of some littoral communi- ginalis Chitande' p.47 in Konings, A. (ed) ties of Lake Nyasa with especial refer- The Cichlids Yearbook, Volume 1, Cichlid ence to the fish, and a discussion of the Press, St. Leon-Rot, Germany. evolution of a group of rock-frequenting KONINGS, A. (1991c) Taeniolethrinops sp. cichlidae. Proc. Zool. Soc. Lond. 132, 11- 'Furcicauda Ntekete'. p.50 in Konings, A. 281. (ed.) The Cichlids Yearbook, Volume 1, FRYER, G. & ILES, T.D. (1972) The Cichlid Cichlid Press, St. Leon-Rot, Germany. Fishes of the Great Lakes of Africa. Oliver KONINGS, A. (1991d) Maravichromis (Capri- & Boyd, Edinburgh. chromis) liemi (McKaye & McKenzie, GREENWOOD, P.H. (1974) The cichlid fishes 1982). p.40 in Konings, A. (ed) The of Lake Victoria, : the biology Cichlids Yearbook, Volume 1, Cichlid and evolution of a species flock. Bull. Brit. Press, St. Leon-Rot, Germany. Mus. Nat. Hist. Suppl. 6, 1-134. KONINGS, A. (1991e) Otopharynx decorus GREENWOOD, P.H. (1994) Lake Victoria. Ad- (Trewavas, 1935). p. 42 in Konings, A. vances in Limnology. (Arch. Hydrobiol. (ed) The Cichlids Yearbook, Volume 1, Beih. Ergebn. Limnol.) 44, 19-26. Cichlid Press, St. Leon-Rot, Germany. HERT, E. (1991) Female choice based on KONINGS, A. (1991f) Maravichromis sp. 'dou- egg-spots in Pseudotropheus aurora Bur- ble spot'. p.43 in Konings, A. (ed) The gess, 1976, a rock-dwelling cichlid of Cichlids Yearbook, Volume 1, Cichlid Lake Malawi. J. Fish Biol. 38, 951-953. Press, St. Leon-Rot, Germany. ILES, T.D. (1960) A group of zooplankton KONINGS, A. (1992) Tramitichromis lituris feeders of the genus Haplochromis (Trewavas, 1931). pp. 38-40 in Konings, (Cichlidae) in Lake Nyasa. Ann. Mag. A. (ed) The Cichlids Yearbook, Volume Nat. Hist. 13, 257-280. 2, Cichlid Press, St. Leon-Rot, Germany. ILES, T.D. (1971) Ecological aspects of KONINGS, A. (1993a) A revision of the ge- growth in African cichlid fishes. J. Conseil. nus Sciaenochromis Eccles & Trewavas, 33, 363-385. 1989 (Pisces, Cichlidae). pp 28-36 in JACKSON, P.B.N. (1961) Check List of the Konings, A. (ed.) The Cichlids Yearbook, Fishes of Nyasaland. Cambridge Univer- Volume 3, Cichlid Press, St. Leon-Rot, sity Press. Germany. KOCHER, T.D., CONROY, J.A., MCKAYE, K.R. KONINGS, A. (1993b). The yellow black line- & STAUFFER, J.R. (1993) Similar morph- the current generic placement of Haplo- ologies of cichlids fish in Lakes Tangan- chromis melanonotus Regan. pp. 43-44 yika and Malawi are due to convergence. in Konings, A. (ed) The Cichlids Year- Mol. Phylogen. Ecol. 2, 158-165. book, Volume 3, Cichlid Press, St. Leon-

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235 Index

Alticorpus 23, 100, 214 C. liemi 149 A. 'deep' 61, 102, 104, 107 C. orthognathus 149, 150, 168 A. 'dwarf mentale' 101 Carcharinus leucas 12 A. 'geoffreyi' 17, 61, 101, 102, 104 Cephalic pits 23 A. macrocleithrum 17, 61, 105 Champsochromis 132, 143 A. mentale 17, 61, 101, 102, 104 Ch. caeruleus 143, 144, 167 A. pectinatum 17, 61, 102, 103, 104 Ch. spilorhynchus 143, 144, 167, 203 A. peterdaviesi 102, 103, 104 Chaoborus 18, 68 A. profundicola 102, 104 Cheilochromis euchilus 190 Aristochromis christyi 147, 168, 203 Chilotilapia rhoadesii 175, 190, 200 Astatotilapia calliptera 12, 27, 214 Chisawasawa 39 Aulonocara 23, 100, 188, 213, 218 Christyella nyasana 36 A. aquilonium 100 Clarias gariepinus 12, 13 A. 'blue-orange' 16, 63, 108, 109, 110 Cleaning behaviour 33, 140 A. brevinidus 100 Copadichromis 38, 39, 66, 142, 152, A. 'copper' 63, 106, 107, 112 192, 213, 218 A. 'dark stripe' 64, 110 C. azureus 67 A. 'deep' 17, 63, 107 C. borleyi 67 A. 'deep yellow' 18, 64, 110 C. 'chrysonotus black' 13, 53, 72 A. gertrudae 100, 109 C. 'chrysonotus blue' 67, 211 A. 'gold' 62, 107 C. chrysonotus 72 A. 'green' 64, 110 C. inornatus 73 A. guentheri 62, 106 C. likomae 66 A. 'long' 112 C. jacksoni 67 A. cf. macrochir 16, 17, 62, 106 C. mbenjii 67 A. 'minutus' 64, 111 C. mloto 67, 68, 69, 73, 74 A. nyassae 107, 109 C. pleurostigma 54, 66, 70, 71 A. 'orange' 14, 63, 108, 109, 110 C. pleurostigmoides 70, 71, 72 A. 'pyramid' 18, 63, 108 C. quadrimaculatus 19, 53, 69, 71 A. rostratum 62, 105, 106 C. thinos 149 A. 'stonemani' 64, 111 C. trimaculatus 71 A. 'yellow' 14, 17, 64, 109, 110 C. verduyni 67 Barbus spp. 12 C. virginalis 13-17, 45, 53, 67, 68, Buccochromis 129, 135, 143 69, 72, 73, 74, 210 B. atritaeniatus 129 Corematodus shiranus 128, 149, 163 B. heterotaenia 129, 203 C. taeniatus 128, 149, 168 B. lepturus 129, 131, 132, 164 Ctenopharynx 152, 153 B. nototaenia 129, 131, 164 Ct. intermedius 124, 154, 155, 169, 177 B. cf. oculatus 129, 130, 164 Ct. nitidus 17, 124, 154, 169, 177 B. rhoadesii 129, 131, 164 Ct. pictus 155, 158 B. spectabilis 129, 132, 143 Cyathochromis obliquidens 36, 50 Caprichromis 149 Cynotilapia 33, 213

236 Cyrtocara moorii 152, 187 H. micrentodon 76 Dimidiochromis 190 H. mloto 68 D. compressiceps 193 H. pholidophorus 147 D. kiwinge 173, 191 H. semipalatus 140 D. strigatus 192 H. sexfasciatus 128 Diplotaxodon 8, 38, 65, 211, 213, 216 H. simulans 205 D. argenteus 39, 45, 46, 47, 51 H. stonemani 111 D. 'bigeye' 18 Hemitaeniochromis 190 'D. big head' 65 H. 'insignis' 174, 194, 195, 196 D. 'big-eye copper' 39 H. spilopterus 174, 194, 195, 196, 198 D. 'big-eye offshore' 39 H. urotaenia 174, 194, 195, 196 D. 'brevimaxillaris' 46, 47, 52 Hemitilapia 152, 153, 158 D. 'copper' 48, 65 H. oxyrhynchus 156, 169 D. 'deep' 46, 47, 52 Heterobranchus 12 D. ecclesi 18, 39 Hydrocynus 12 D. greenwoodi 46, 48, 52 Key to the offshore cichlids 20 D. holochromis 46 Labeo mesops 219 D. 'intermediate' 45, 46 Lethrinops 8, 23, 39, 76, 119, 133, D. limnothrissa 16, 18, 38, 45, 46, 51, 152, 201, 214 67 L. albus 87 D. 'macrops' 17, 48, 52, 65 L. altus 14, 16, 56, 83, 85,86 D. 'micrentodon' 39 L. argenteus 201 D. 'similis' 46, 47, 51 L. auritus 91 D. 'white belly' 48, 52, 65 L. 'bighead' 92, 93 Docimodus johnstonii 148, 168 L. 'black chin' 56, 83 D. evelynae 148 L. 'blue-orange' 56, 83 Distichodus 12 L. christyi 14, 16, 56, 86,98 Eclectochromis 202 L. 'dark' 16, 55, 80, 82,84 E. festivus 206 L. 'deep-water albus' 17, 57, 87, 89, 90 E. lobochilus 206 L. 'deep-water altus' 17, 56, 85 E. ornatus 205, 206 L. 'Domira blue' 18, 159, 170 Engraulicypris sardella 13, 19, 38, 66 L. fasciatus 99, 134 Exochochromis 152, 153, 185 Lethrinops cf furcifer 59, 95, 97, 98 E. anagenys 172, 181, 182, 185 L. gossei 16, 17, 57, 87, 89, 90 Fossorochromis 202 L. 'grey' 16, 80 F. rostratus 150, 187, 203, 207 L. lethrinus 13, 14, 88, 99, 176, 201 Genyochromis mento 33 L. longimanus 15, 17, 58, 89, 90, 91 36, 50 L. longipinnis 14, 15, 17, 57, 58, Haplochromis bellicosus 144 85, 87, 88, 91 H. bodyi 191 L. 'loweae' 91 H. fuelleborni 192 L. macracanthus 58, 89, 90, 91 H. gigas 132 L. macrochir 13, 59, 91, 93, 94, 96 H. 'guttatus' 185 L. macrorhynchus 99, 134 H. leuciscus 177 L. 'macrostoma' 59, 92, 93 H. leucogaster 65 L. 'matumbae' 56, 83, 85 H. longimanus 76 L. micrentodon 77, 80, 81, 91, 123- H. longipes 145 125, 160 H. macrognathus 120 L. 'micrentodon Makokola' 55, 77, 79, H. macrorhynchus 208 80, 81, 91

237 L. microdon 16, 17, 55, 76, 77, 78, M. sphaerodon 136, 137, 141 80, 91 M. spilostichus 17, 122, 132, 146, 167 L. mylodon 58, 89 M. 'torpedo' 132, 143, 166, 167, 179 L. m. borealis 89 M. 'torpedo blue' 179 L. m. mylodon 89 Naevochromis 152 L. 'oliveri' 16, 17, 55, 77, 78, 80, 82, N. chrysogaster 186 83, 91 Ncheni 38 L. orbitalis 87 Ndunduma 38 L. cf. parvidens 14, 59, 91, 92, 94, 95- Nimbochromis 202, 218 97 N. fuscotaeniatus 202 L. 'pink head' 13, 59, 91, 92, 93, 94, 96, N. linni 176, 203, 205 123, 125 N. livingstonii 17, 119, 176, 202, 210 L. polli 17, 57, 86, 119, 120 N. maculimanus 204 L. stridei 14, 55, 76, 77, 78, 80, 91 N. pardalis 204 L. 'yellow' 16, 17, 56, 83, 84 N. polystigma 119, 127, 176, 203, 205, L. 'yellow chin' 16, 55, 77, 80, 81, 208 83, 91, 159, 160, 188 N. venustus 176, 204 L. 'yellow tail' 55, 83 Nyassachromis 66, 190 Lichnochromis 133 N. 'argyrosoma blue' 14, 16, 54, 74, 75, L. acuticeps 134 123, 125, 126, 179 Malapterurus 12 N. boadzulu 66 Maravichromis 133 N. conophoros 73 M. 'silver torpedo' 143 N. conophorus 67 M. sp. 'Kande' 145 N. cyclicos 67, 73 Mbuna 32 N. eucinostomus 45, 66, 67, 74 33 N. 'eucinostomus black' 54, 67, 73 M. parallelus 33 N. 'eucinostomus yellow' 54, 67, 73 Mylochromis 126, 129, 133 N. inornatus 67, 73 M. anaphyrmus 16, 17, 135, 136, N. leuciscus 75, 178 137, 141, 165 N. microcephalus 54, 74 M. cf balteatus 138, 165 N. nigritaeniatus 200 M. 'chekopae' 17, 141, 142, 181 N. prostoma 66, 67 M. 'deep' 18, 136, 137, 141, 166 N. thinos 67, 73 M. 'double spot' 142, 181 Opsaridium microcephalus 13, 19 M. epichoralis 141 O. microlepis 19 M. ericotaenia 127, 138, 141, 165 Oreochromis 11, 15, 16, 128, 217 M. formosus 132, 143, 144, 145, 146, O. karongae 28, 49 167 O. lidole 28, 29, 49 M. gracilis 132, 146, 167 O. macrochir 214 M. guentheri 141 O. mossambicus 12, 30, 214 M. incola 141 O. placidus 12, 214 M. labidodon 127, 141 O. saka 29 M. lateristriga 133, 134, 187 O. shiranus 12, 30, 49, 214 M. melanonotus 139, 141, 166 O. squamipinnis 27, 28, 49 M. melanotaenia 140, 141, 166 Otopharynx 152 M. mola 141 O. argyrosoma 67, 73, 75, 152, 160, M. mollis 141 180 M. obtusus 141 O. 'argyrosoma deep' 15, 17, 74, 160, M. plagiotaenia 141, 166 170, 180

238 O. 'argyrosoma red' 13, 74, 171, 178, P. cf. subocularis 126, 163 179 Platygnathochromis 140 O. auromarginatus 15, 157, 170, 177, Pristis spp. 12 178 Protomelas 190, 213 O. 'auromarginatus margrette' 177 P. insignis 196, 197 O. 'auromarginatus stripe' 170, 177, 178, P. 'insignis mumbo' 197 181 P. kirkii 175, 197 O. brooksi 119, 172, 182, 184 P. labridens 175, 197, 198 O. decorus 160, 170, 178, 181 P. macrodon 200 O. heterodon 158, 159, 188, 189 P. marginatus 198, 200 O. 'heterodon Nankumba' 158 P. m. marginatus 199 O. 'modestus makokola' 184 P. m. vuae 199 O. ovatus 196 P. 'paedophage' 151 O. 'productus' 17, 143, 160, 171, P. pleurotaenia 200 178, 179, 181 P. 'red dorsal' 175, 200 O. selenurus 152, 160, 178 P. similis 175, 197, 198 O. speciosus 16, 172, 182, 184 P. triaenodon 175, 200 O. tetraspilus 157, 158, 169, 188, 189 Pseudocrenilabrus philander 12 O. tetrastigma 13, 152, 157, 158, 169, Pseudotropheus 33 188, 189 Ps. 'acei' 36 O. 'yellow fin mloto' 158 Ps. callainos 33 Paedophages 186, 196 Ps. elegans 14, 16, 32, 34, 35, 50 Pallidochromis 38 Ps. lanisticola 17, 32, 34, 35, 50 P. tokolosh 39, 47, 52, 65 Ps. livingstonii 14, 15, 32, 33, 34-36, 50 Petrotilapia 33 Ps. 'livingstonii likoma' 32 Placidochromis 76, 112 Ps. lucerna 33 P. 'acuticeps' 119, 120, 161 Ps. pursus 32, 33 P. 'carnivore' 119, 161 Ps. tropheops complex 33, 36, 84 P. 'green stripe' 122 Ps. 'tropheops olive' 32 P. hennydaviesae 113, 117, 161 Ps. zebra complex 33 P. 'hennydaviesae II' 113 Ps. 'zebra gold' 34 P. 'hennydaviesae III' 116, 161 Rhamphochromis 8, 14, 16, 19, 20, P. 'hennydaviesae IV' 116 38, 65, 190, 214 P. 'hennydaviesae V' 117 R. 'big mouth' 41 P. 'hennydaviesae VI' 117 R. 'bigtooth brown' 40 P. johnstoni 126, 127, 161, 203 R. brevis 40 P. 'Johnstoni Gold' 128 R. esox 39, 50, 143 P. 'johnstoni solo' 128 R. ferox 41, 42 P. 'long' 120, 123, 162 R. 'kolowilo' 44, 51 P. longimanus 13, 74, 94, 124 R. leptosoma 39, 143 P. 'longimanus Chirombo' 126, 163 R. 'long fin yellow' 44, 51 P. 'longimanus Maleri' 126, 163 R. cf. longiceps 19, 44, 50 P. 'longimanus Malombe' 93, 123, 125 R. 'long snout' 41, 50 P. 'longimanus Namiasi' 14, 16, 77, R. lucius 43 123, 125, 126, 163 R. macrophthalmus 40 P. 'longimanus slender' 126 R. pallidus 65 P. 'macrognathus' 119, 120, 161 R. 'shire ferox' 42, 51 P. 'platyrhynchos' 113, 116, 117, R. 'short-tooth brown' 43 120, 121 R. woodi 40

239 Sciaenochromis 113, 117 Sc. ahli 120 Sc. benthicola 121, 162 Sc. 'deep water' 120, 161 Sc. fryeri 121 Sc. psammophilus 120, 121, 162 Sc. 'silver torpedo' 143 Serranochromis robustus 27 Stigmatochromis 152, 153 S. 'guttatus' 65, 172, 181, 183, 185 S. pholidophorus 171, 181 S. pleurospilus 182 S. 'Spilostichus type' 147 S. 'tolae' 183 S. woodi 171, 181, 182, 183 Synodontis njassae 14, 15, 17, 19 Taeniochromis holotaenia 173, 190 Taeniolethrinops 23, 76, 124, 133, 187 T. furcicauda 16, 17, 61, 91, 97, 98, 133 T. 'furcicauda Ntekete' 96, 98 T. laticeps 17, 97, 98, 99, 133, 165 T. 'laticeps itungi' 99 T. praeorbitalis 60, 91, 97, 98, 133 Tilapia rendalli 12, 30 T. sparrmanii 12 Tramitichromis 23, 76, 124, 133 T. brevis 99, 133, 135, 136, 137 T. intermedius 99, 152, 168 T. lituris 60, 92, 93, 94, 96 T. variabilis 97 Trematocranus 152 T. brevirostris 14, 16, 81, 159, 173, 188 T. 'brevirostris deep' 159, 173, 188, 189 T. labifer 152, 154 T. microstoma 152, 158, 169, 188 T. placodon 14, 152 153 Tyrannochromis 190 Utaka 39, 66

240 2 C M Y K Offshore CichlidsofLakeMalawi Offshore Cichlids of Lake Malawi OFFSHORE CICHLIDS Lake Malawi is internationally renowned among evolution- ary biologists and aquarium enthusiasts alike for its immense number of species of cichlid fishes, which display a dazzling of repertoire of structural diversity, feeding specialisations, and bril- liant colours. Almost all of the attention has focused on the rocky shore fish or mbuna, and it has been assumed that the offshore waters and the sandy or muddy shores harbour a relatively low- LAKE MALAWI diversity community dominated by non-cichlid fishes.

This book, the first ever guide to the cichlid fishes of the off- shore waters of Lake Malawi, shows that this habitat harbours a great diversity of little-known cichlid species, which dominate the catches of the commercial and artisanal food fisheries. How they evolved, how they continue to coexist, and how to manage George F. Turner their exploitation for food while conserving these unique spe- cies will be major challenges to scientists and policymakers. Aquarists will be fascinated by the wealth of previously-unknown brilliantly-coloured and bizarrely-shaped species.

Information on distinguishing features, colour, size, distribution and abundance, commercial importance, diet, reproduction, and taxonomy are given for 199 species, 79 of which are pres- ently undescribed. 186 colour and 44 black and white photo- graphs illustrate all species. Also included are figures showing the main distinguishing features of the fish, maps, a glossary of George F. Turner technical terms, a full bibliography and brief discussion of the evolution, ecology and conservation of these fishes.

The author, Dr George Turner, has worked on African cichlid fishes for 13 years and has spent more than 3 years in Malawi. Most of the information collected in this book was obtained while working on the commercial fisheries of the southern lake for the UN Food and Agriculture Organisation from 1990 to 1992. He is presently lecturer in Ecology at the University of Southampton, England, and continues to carry out research on evolution, ecology, behaviour, taxonomy, conservation, and management of African freshwater fishes.

ISBN 3-928457-33-0

C M Y K