The Pennsylvania State University the Graduate School

The Pennsylvania State University

The Graduate School

SYSTEMATICS AND BOWER-BUILDING BEHAVIOR OF THE

TRAMITICHROMIS ECCLES AND TREWAVAS (TELEOSTEI: CICHLIDAE)

FROM THE SOUTHEAST ARM OF LAKE MALAWI, AFRICA

A Thesis in

Ecology

Matthew R. Lisy

Submitted in Partial Fulfillment of the Requirements for the Degree of

Doctor of Philosophy

December 2006

The thesis of Matthew R. Lisy was reviewed and approved* by the following:

Jay R. Stauffer, Jr. Distinguished Professor of Ichthyology Thesis Advisor Chair of Committee

Cecilia Paola Ferreri Associate Professor of Fisheries Management ...

Ke Chung Kim Professor of Entomology and Curator; Director, Center for Biodiversity Research ...

Ganapati P. Patil Distinguished Professor of Mathematical Statistics and Director, Center for Statistical Ecology and Environmental Statistics ...

David Mortensen Professor and Chair Ecology IGDP ... Head of the Intercollege Graduate Degree Program in Ecology

*Signatures are on file in the Graduate School

iii ABSTRACT

The Malaŵians derive 70% of their consumed animal protein from fish from Lake

Malaŵi. It is the goal of the Malaŵi government to preserve and conserve this resource, but effective fishery management plans cannot be developed because species descriptions are still lacking. Part of my research focused on fish identified as Tramitichromis (an important food fish) in the southeast arm of Lake Malaŵi. Based upon suggestive evidence in a collection of works (see bibliography in this thesis) by Konings, Turner, numerous publications by Stauffer along with various other authors, and personal communications of Konings and Stauffer, I hypothesized that there were at least three undescribed species.

Collections of fish from the Pennsylvania State University and the Museum of

Natural History, London were examined. I examined each collection in the laboratory and reexamined the identity of the fish based on the keys in Tramitichromis. A subset of fish was chosen out of each collection and lower pharyngeal bones dissected, and then used to verify the species. Fishes that could not be identified were grouped together based on some phenotypic character(s). Twenty-four morphometric and fourteen meristic data points were collected per fish. Differences in body shape were analyzed using sheared principal components analysis (SPCA) of the morphometric data.

Differences among species were illustrated by plotting the sheared components of the morphometric data against the principal components of the meristic data in order to maximize the amount of separation. If the mean multivariate scores of the clusters formed by the plots were significantly different along one axis, independent of the other

iv axis, a Duncan’s multiple range test (p<0.05) was used to determine which clusters

differed from each other. If not, then a MANOVA, in conjunction with a Hotelling-

Lawley trace, was used (p<0.05).

Collections in the 1920s and 1930s allowed Trewavas to describe many new

species of Lethrinops and place Haplochromis brevis into the Lethrinops based on buccal

dentition. In 1935, Trewavas added one new species, Lethrinops intermedia. In 1989

Eccles and Trewavas placed all Lethrinops that possessed a keel into Tramitichromis.

Tramitichromis is rediagnosed by the presence of a solid diagonal band that runs from the nape to the caudal peduncle, a significantly different body shape (e.g. a shorter caudal peduncle, longer vertical eye diameter, longer lower jaw, and fewer gill rakers on the outer ceratobranchial), and the use of a rock in cone shaped bowers constructed by breeding males. Tramitichromis brevis is retained in the genus. The remaining species currently in Tramitichromis are moved to a new genus. Six previously undescribed species that have variations in their lower pharyngeal bone, body pattern, and/or shape are described.

Bower building is the manifestation of a behavioral trait and is being used to diagnose species. Another research objective was to determine if it was possible to test bower-building behavior in a laboratory setting, and give comments and suggestions for future research. I had also hoped to provide some anecdotal evidence of the heritability of bower building. Comparisons to an analysis of a previous bower building study showing overlap between genetic and bower (behavioral) data were made as well. A new study for three of the new species was conducted which showed a different bower shape

v for each of the three species. This is significant because it supports the use of the bower building behavior as a taxonomic tool and shows correlation between morphological data and bower building (behavioral) data. Comments on the feasibility of laboratory studies of bower building are made where it was determined that an extremely large pool was needed with a high stocking density and a ratio of 7 males to 2 females.

vi TABLE OF CONTENTS

LIST OF FIGURES ...... vii

LIST OF TABLES...... xiii

ACKNOWLEDGEMENTS...... xv

Chapter 1 Introduction...... 1

Chapter 2 Materials and Methods...... 8

Chapter 3 Taxonomic Character Analysis ...... 17

Chapter 4 Taxonomy of Tramitichromis ...... 21

Chapter 5 Discussion and Conclusions...... 117

Literature Cited...... 121

Appendix A Laboratory Feasibility Study...... 130

A.1 Introduction...... 130 A.2 Methods ...... 131 A.3 Results and Discussion ...... 133

Appendix B Tables of Morphometric and Meristic Values...... 143

vii LIST OF FIGURES

Figure 2.1: Known localities of Tramitichromis species, both described and undescribed, in this study...... 9

Figure 2.2: Illustration of the 24 morphometric data points...... 12

Figure 2.3: Illustration of the 14 meristic data points...... 13

Figure 2.4: Schematic illustration showing measurements recorded for bowers constructed by breeding males in the Apetra group (A – width of base; B – slope length; C – outside top diameter; D – inside top diameter; E – height; F – lip length)...... 16

Figure 4.1: Characteristics of Tramitichromis brevis. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) anterior pharyngeal teeth, d) posterior pharyngeal teeth, e) lateral view of lower pharyngeal bone, f) dorsal view of lower pharyngeal bone. Individual pictured is from Cobue, PSU 4089, #6...... 24

Figure 4.2: Location of the collections of Tramitichromis brevis, PSU 4089...... 26

Figure 4.3: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Tramitichromis brevis (N = 24) PSU 4089; and the other described Tramitichromis species type material from the British Museum, T. variabilis (N = 22) BMNH 1930.1.31.14-20; BMNH 1930.1.31.1-2; BMNH 1930.1.31.4-13; BMNH 1930.1.31.3; T. lituris (N = 9) BMNH 1930.1.31.21-23; BMNH 1930.1.31.24-28; BMNH 1930.1.31.45; T trilineata (N = 1) BMNH 1930.1.31.76; T. intermedius (N = 6) BMNH 1935.6.14.2081-2084; BMNH 1935.6.14.2085...... 29

Figure 4.4: Characteristics of Apetra lituris. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. Individual pictured is the Lectotype from Karonga BMNH 1930.1.31.21...... 33

Figure 4.5: Typical characteristics of Apetra lituris lower pharyngeal bone and anterior teeth. Top to bottom: a) lateral view of lower pharyngeal bone, b) anterior pharyngeal teeth. Individual pictured is a Paralectotype from Karonga BMNH 1930.1.31.23...... 34

Figure 4.6: Localities of Apetra lituris: Karonga BMNH 1930..31.21-23; Vua BMNH 1930.1.31.24-28; Mwaya BMNH 1930.1.31.35-44...... 36

viii Figure 4.7: Plot of the sheared second principal components (morphometric data) and the first factor scores (meristic data) of BMNH Apetra lituris: Mwaya (N = 24) BMNH 1930.1.31.35-44; Fort Maguire (N = 1) BMNH 1930.1.31.45; Vua (N = 5) BMNH 1930.1.31.24-28; Karonga (N = 3) BMNH 1930.1.31.21-23. The damaged all female group from Mwaya (N = 13) appear as separate cluster, BMNH 1930.1.31.29-34...... 39

Figure 4.8: Plot of the sheared second principal components (morphometric data) and the first factor scores (meristic data) of Apetra lituris types without the poorly preserved Mwaya group: Mwaya (N = 24) BMNH 1930.1.31.35-44; Fort Maguire (N = 1) BMNH 1930.1.31.45; Vua (N = 5) BMNH 1930.1.31.24-28; Karonga (N = 3) BMNH 1930.1.31.21-23...... 40

Figure 4.9: Characteristics of Apetra intermedia. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) posterior pharyngeal teeth, f) anterior pharyngeal teeth. Individual pictured is from Chembe Village, PSU 4147, #1...... 43

Figure 4.10: Dorsal view of lower pharyngeal bones from Apetra intermedia from (top row, left to right) Chembe Village PSU 4147, #1; Chembe Village PSU 4156, #16; Golden Sands Swamp PSU 4117, #2; Kanjedza Island 4107, #5; and (bottom row, left to right) Kanjedza Island PSU 4081, #1; Kanjedza Island PSU 4081, #3; Kanjedza Island 4101, #4; Kanjedza Island PSU 4101, #8...... 44

Figure 4.11: Localities of BNMH and PSU collections of Apetra intermedia South BMNH 1935.6.14.2081-2084; Monkey Bay BMNH 1935.6.14.2085; Chembe Village PSU 4147, 4144, 4092, 4104, 4156; Golden Sand Swamp PSU 4117; Kanjedza Island PSU 4101, 4081, 4107,4110...... 47

Figure 4.12: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra intermedia type material from the British Museum (N = 6) BMNH 1935.6.14.2081-2084; BMNH 1935.6.14.2085; and populations from Chembe Village (N = 18), PSU 4092, 4104, 4144, 4147, 4156; Golden Sands Swamp (N = 2) PSU 4117; and Kanjedza Island (N = 49) PSU 4081, 4101, 4107, 4110...... 49

Figure 4.13: Plot of the second sheared principle component and the first factor scores of two distant populations of Apetra intermedia: Chembe Village (N = 18), PSU 4092, 4104, 4144, 4147, 4156; Kanjedza Island (N = 49), PSU 4081, 4101, 4107, 4110...... 50

Figure 4.14: Characteristics of Apetra variabilis. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of

ix lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. Individual pictured is the Lectotype from Lake Nyasa South, BMNH 1930.1.31.4...... 53

Figure 4.15: Localities of BMNH and of Apetra variabilis: Monkey Bay BMNH 1930.1.31.3; South BMNH 1930.1.31.4-13. The exact location(s) of “South” is/are unknown...... 55

Figure 4.16: Characteristics of Apetra trilineata. Top to bottom: a) external appearance, b) gill rakers on outer ceratobranchial. The specimen pictured is the holotype, BMNH 1930.1.31.76...... 58

Figure 4.17: Proposed location of Apetra trilineata, BMNH 1930.1.31.76...... 60

Figure 4.18: Characteristics of Apetra linea. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. Individual pictured is the Holotype from Vua, BMNH 1930.1.31.17...... 64

Figure 4.19: Comparison of body patterns (top to bottom) of Apetra linea: a) spots, BMNH 1930.1.31.17 from Vua; and b) a broken non-overlapping oblique line PSU 4145 fish #1 from Fisheries Research Station...... 65

Figure 4.20: Locations of Apetra linea: Vua BMNH 1930.1.31.14-20; Mwanga BMNH 1930.1.31.1-2; Fisheries Research Station PSU 4139, 4140, 4141, 4142, 4145, 4150, 4161...... 67

Figure 4.21: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of the type material of Apetra variabilis (N = 12): Lake Nyasa South BMNH 1930.1.31.4-13; Monkey Bay BMNH 1930.1.31.3; and Apetra linea (N = 10): Vua BMNH 1930.1.31.14- 20; Mwanga BMNH 1930.1.31.1-2...... 69

Figure 4.22: Characteristics of Apetra simula. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. Individual pictured is the Holotype from Otter Point PSU 4187...... 72

Figure 4.23: Localities of Apetra simula: Otter Point PSU 4097, 4187; Golden Sands Swamp PSU 4112, 4113, 4118, 4121, 4122...... 74

Figure 4.24: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra simula (N = 38):

x Otter Point PSU 4097, 4187; Golden Sands Swamp 4112, 4113, 4118, 4121, 4122; Apetra linea (N = 10): Vua BMNH 1930.1.31.14-20; Mwanga BMNH 1930.1.31.1-2; Fisheries Research Station PSU 4139, 4140, 4141, 4142, 4145, 4150, 4161; and Apetra variabilis (N = 12): Lake Nyasa South BMNH 1930.1.31.4-13; Monkey Bay BMNH 1930.1.31.3...... 76

Figure 4.25: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra linea caught at Fisheries Research Station at different depths: 12m (N = 14) (PSU 4139, 4140, 4141, 4142, 4161), and 36-54m (N = 6) (PSU 4145, 4150)...... 78

Figure 4.26: Characteristics of Apetra perjur. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) posterior pharyngeal teeth, f) anterior pharyngeal teeth. Specimen shown is the holotype from Songwe Hill PSU 4162...... 80

Figure 4.27: Localities of Apetra perjur: Songwe Hill PSU 4087, 4096, 4162 and the bar to Fort Maguire BMNH 1930.1.31.45...... 83

Figure 4.28: Lateral view (left) and anterior pharyngeal teeth (right) of (from top to bottom): Apetra lituris - northern localities a) Karonga BMNH 1930.1.31.21-23, b) Vua BMNH 1930.1.31.24-28, c) Mwaya BMNH 1930.1.31.35-44; Apetra perjur - southern localities d) Fort Maguire BMNH 1930.1.31.45, e) Songwe Hill PSU 4087, 4096, 4162...... 85

Figure 4.29: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra perjur (N = 46): BMNH 1930.1.31.45; PSU 4087, 4096, 4162; and Apetra lituris (N = 32): BMNH 1930.1.31.21-23, BMNH 1930.1.31.24-28, BMNH 1930.1.31.35-44. ...86

Figure 4.31: Location of the collection of Apetra meniscosteum: Kanjedza Island PSU 4130, 4134, and 4163...... 91

Figure 4.32: Characteristics of Apetra cryptopharynx. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. The specimen pictured is the holotype from Kanjedza Island PSU 4186...... 94

xi Figure 4.33: The keels of six individuals showing the variability of the shape and length of the keel of Apetra cryptopharynx. Clockwise starting with the top left Kanjedza Island individuals pictured are from collections: a) PSU 4186 holotype, b) PSU 4136 #4, c) PSU 4105 #3, d) PSU 4105 #8, e) PSU 4105 #10, f) PSU 4105 #1...... 95

Figure 4.34: Location of the collection of Apetra cryptopharynx: Kanjedza Island PSU 4080, 4105, 4131, 4136, 4186; Golden Sands Swamp PSU 4093; Songwe Hill 4082, 4085, 4133; Nkhudzi Bay PSU 4115; Otter Point 4083, 4111, 4120...... 97

Figure 4.35: Plot of the third sheared principle components (morphometric data) and the second factor scores (meristic data) of Apetra cryptopharynx (N = 220): Kanjedza Island PSU 4080, 4105, 4131, 4136, 4186; Golden Sands Swamp 4093; Nkhudzi Bay 4115; Otter Point 4083, 4111, 4120; SongweHill 4082, 4085, 4133; and Apetra meniscosteum (N = 20) Kanjedza Island PSU 4130, 4134,4163...... 98

Figure 4.37: Localities of Apetra retrodens: PSU 4084, 4095, 4119, 4146, 4149, 4151, 4152, 4153, 4154, 4156, 4157, 4158, 4159, 4160, 4164...... 104

Figure 4.38: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra retrodens (N = 113): Chembe Village PSU 4084, 4095, 4119, 4156, 4157, 4158, 4159, 4160, 4164; Golden Sands Swamp 4151, 4154; Fisheries Research Station 4146, 4149, 4152, 4153; and Apetra meniscosteum (N = 20): Kanjedza Island PSU 4130, 4134, 4163...... 107

Figure 4.39: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra retrodens (N = 113): Chembe Village PSU 4084, 4095, 4119, 4156, 4157, 4158, 4159, 4160, 4164; Golden Sands Swamp 4151, 4154; Fisheries Research Station 4146, 4149, 4152, 4153; and Apetra cryptopharynx (N = 220): Kanjedza Island PSU 4080, 4105, 4131, 4136, 4186; Golden Sands Swamp 4093; Nkhudzi Bay 4115; Otter Point 4083, 4111, 4120; SongweHill 4082, 4085, 4133...... 108

Figure 4.40: Plot of the second sheared principle components (morphometric data) and the second factor scores (meristic data) of the type material of

xii Apetra retrodens (N = 113): Chembe Village PSU 4119, 4164; Apetra meniscosteum (N = 20): Kanjedza Island PSU 4130, 4134, 4163; and Apetra cryptopharynx (N = 220): Kanjedza Island PSU 4131, 4136, 4186...... 109

Figure 4.41: Comparison of the lateral view of the keels from the holotypees of (left to right), Apetra meniscosteum PSU 4163, Apetra cryptopharynx PSU 4186, and Apetra retrodens PSU 4164...... 109

Figure 4.42: Plot of the third sheared principle components (morphometric data) and the second sheared principle components (morphometric data) of the in situ bower data of Apetra meniscosteum (N = 10) from Kanjedza Island (PSU 4134), Apetra cryptopharynx (N = 15) from Kanjedza Island (PSU 4105), and Apetra retrodens (N = 20) from Chembe Village (PSU 4084, 4095, 4119, 4164)...... 112

Figure 4.43: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of the morphology data of Apetra meniscosteum (N = 11) from Kanjedza Island (PSU 4134), Apetra cryptopharynx (N = 15) from Kanjedza Island (PSU 4105), and Apetra retrodens (N = 35) from Chembe Village (PSU 4084, 4095, 4119, 4164)...... 113

Figure A.1: Schematic illustration showing measurements recorded for bowers constructed by breeding males in the Apetra group (A – width of base; B – slope length; C – outside top diameter; D – inside top diameter; E – height; F – lip length)...... 133

Figure A.2: Examples of the bowers build by male Apetra sp. in the pools. The top picture shows a male towards the beginning of construction, while the bottom picture shows a fully functional bower with fish spawning in it...... 135

Figure A.3: Characteristics of the lab fish (Apetra cryptopharynx). Clockwise from the top: a) external appearance, b) dorsal view of lower pharyngeal bone, c) lateral view of lower pharyngeal bone, d) anterior pharyngeal teeth, e) posterior pharyngeal teeth...... 137

Figure A.4: Diagram of bowers (gray) built in the pool (clear). Two-thirds of the bower was not built, as it would extend beyond the walls of the pool...... 140

Figure A.5: Suggested pond structure and expectant bower placement. Only the center three bowers would be completely useful...... 142

xiii LIST OF TABLES

Table 2.1: Morphometric and meristic measurements that were taken on each fish...11

Table 4.1: Character matrix for Apetra species...... 114

Table B.1: Morphometric and meristic values of the Tramitichromis brevis population from Cobue (N = 24) PSU 4089...... 143

Table B.2: Morphometric and meristic values of Apetra lituris type material, which includes the lectotype (N = 32) from Karonga, BMNH 1930.1.31.21- 23; Vua, BMNH 1930.1.31.24-28; Mwaya, BMNH 1930.1.31.35-44. Morphometric and meristic values of the Apetra lituris holotype BMNH 1930.1.31.21 are also listed...... 144

Table B.3: Morphometric and meristic values of Apetra intermedius populations from Chembe Village (N = 17) PSU 4147, 4144, 4092, 4104, 4156; Golden Sand Swamp (N = 2) PSU 4117; Kanjedza Island (N = 49) PSU 4101, 4081, 4107,4110. Morphometric and meristic values of the Apetra intermedius types are also listed, which includes the lectotype South BMNH 1935.6.14.2081-2084; Monkey Bay BMNH 1935.6.14.2085. The morphometric and meristic values of the Apetra intermedius lectotype BMNH 1935.6.14.2081 are also listed...... 145

Table B.4: TABLE A.3 (concluded)...... 146

Table B.5: Morphometric and meristic values of the Apetra variabilis types, which include the lectotype (N = 12), from Monkey Bay, BMNH 1930.1.31.3; South, BMNH 1930.1.31.4-13. Morphometric and meristic values of the Apetra variabilis lectotype BMNH 1930.1.31.4 are also listed...... 147

Table B.6: Morphometric and meristic values of the Apetra trilineata holotype from an unknown locality BMNH 1930.1.31.76...... 148

Table B.7: Morphometric and meristic values of Apetra linea type material, which includes the holotype (N = 10) Vua BMNH 1930.1.31.14-20 and Mwanga BMNH 1930.1.31.1-2; Fisheries Research Station (N = 20) PSU 4139, 4140, 4141, 4142, 4145, 4150, 4161. Morphometric and meristic values of the holotype BMNH 1930.1.31.17 are also listed...... 149

Table B.8: Morphometric and meristic values of the Apetra simula types, which includes the holotype, are listed (N = 41) from Otter Point PSU 4097, 4118, 4121, 4122, 4187; and Golden Sands Swamp PSU 4112, 4113.

xiv Morphometric and meristic values of the Apetra simula holotype PSU 4187 are also listed...... 150

Table B.9: Morphometric and meristic values of Apetra perjur type material, which includes the holotype, from Songwe Hill (N = 45) PSU 4087, 4096, 4162; and the bar to Fort Maguire (N = 1) BMNH 1930.1.31.45. Morphometric and meristic values of the Apetra perjur holotype PSU 4162 are also listed...... 151

Table B.10: Morphometric and meristic values of Apetra meniscosteum type material, which includes the holotype, (N = 20) from Kanjedza Island PSU 4130, 4134, 4163. Morphometric and meristic values of the Apetra meniscosteum holotype are also listed PSU 4163...... 152

Table B.11: Morphometric and meristic values of Apetra cryptopharynx type material from Kanjedza Island, which includes the holotype, (N = 89) PSU 4080, 4105, 4131, 4136, 4186; Golden Sands Swamp (N = 6) PSU 4093; Songwe Hill (N = 69) PSU 4082, 4085, 4133; Nkhudzi Bay (N = 15) PSU 4115; Otter Point (N = 41) PSU 4083, 4111, 4120. Morphometric and meristic values of the Apetra cryptopharynx holotype are also shown PSU 4186...... 153

Table B.12: TABLE A.11 (concluded)...... 154

Table B.13: Morphometric and meristic values of Apetra retrodens type material from Chembe Village, which includes the holotype, (N = 78) PSU 4084, 4095, 4119, 4156, 4157, 4158, 4159, 4160, 4164; Golden Sands Swamp (N = 11) PSU 4151, 4154; Fisheries Research Station (N = 24) PSU 4146, 4149, 4152, 4153. Morphometric and meristic values of the Apetra retrodens holotype are also shown PSU 4164...... 155

xv ACKNOWLEDGEMENTS

I would like to thank Dr. Jay Stauffer, Jr. for his guidance along the way, and providing me a means to support myself throughout most of my graduate career with teaching and research assistantships. I would especially like to thank you for all the patience and help you gave via email while you were in Africa. You have provided me with many opportunities for intellectual growth and enrichment during my tenure at Penn

State and for that I thank you also. Dr. Ganapati Patil, thank you so much for your statistical guidance. I enjoyed your classes and our talks in your office. Dr. Ke Chung

Kim, I really appreciate your mentorship throughout this process. Your blunt, to the point method really helped steer me in the right direction on numerous occasions. Dr.

Paola Ferreri, thank you for all that you have taught me. I enjoyed working with you in

West Virginia, and on other projects. Your advice both academic and personal was really important and well received.

Many thanks to Jack Yarnell for feeding the fish and helping with the maintenance on them while I was in the field. I do not even know how to say thank you to Timothy Stecko who has been so important to the completion of this work. You have helped at so many points along the way, and asked nothing in return. Penn State is very fortunate to have a person like you on staff, because you helped many with their research, like you did for me. I do not know how you find time for it all. Thank you to Leslie

Leckvarcik for allowing me to work on the minnow project. You taught me a lot and enabled me to support myself for another semester.

xvi A sincere thank you to Mr. Oliver Crimmen, Curator of Reptiles, Amphibia, and

Fish and Mr. James MacLaine, Assistant Curator of Reptiles, Amphibia, and Fish from the Natural History Museum, London, England, for sending me the type specimens of the

Tramitichromis. I could not have done this work without them.

I would also like to thank my grandfather, E. Eugene Peldyak, my parents, Roger and Sally Lisy, and my in-laws, Robert and Maribeth Schwartz for all their support in ways too numerous to list. Claire Schwartz, thank you for double-checking the statistics in the excel program I made. Wesley J. Neal thanks for your help and friendship throughout this process. Thank you to my wife and best friend, Emily, for all your help, support, understanding, and patience while I finished this dissertation. I dedicate this work to you.

Chapter 1

Introduction

Cichlids are found throughout the world, with 70% of them found in Africa

(Greenwood, 1991). Their unique behaviors, morphological adaptations, and rapid speciation, have fascinated ecologists and systematists for many years. With more than

460 described haplochromine cichlid species in Lake Malaŵi alone, all but one being endemic, there are more cichlids in Lake Malaŵi than any other freshwater lake

(Greenwood, 1991; Konings, 2001). With estimates of the total number of species in the lake in excess of 850 (Konings, 2001), many of the species are undescribed (Stauffer et al., 1997b; Turner et al., 2001).

What is worrisome to biologists working in Lake Malaŵi is the ever-present threat of over fishing (Stauffer et al., 1995). The Malaŵians derive 70% of their consumed animal protein from fish from Lake Malaŵi (Stauffer et al., 1995). It is the goal of the Malaŵi government to preserve and conserve this resource (Jay Stauffer, per. comm.), but effective fishery management plans cannot be developed because species descriptions are still lacking. It would be impossible to manage species that are unknown to fishery managers. Distribution maps cannot be developed until scientists know what species are present. Treating many undescribed species as one large group could result in the loss of species diversity, which could have devastating effects on the ecosystem. The lake has already experienced this with the over fishing of Trematocranus placodon and the resultant schistosomiasis outbreak (Stauffer et al., 1997a).

2 To help remedy this problem, my research objectives were to resolve the species status of populations of cichlids within the genus Tramitichromis in the Southeast Arm of

Lake Malaŵi, show evidence to support the use of bower shape as a taxonomic tool by showing congruence between morphology data and bower data, provide anecdotal evidence of the heritability of bower building, and conduct a laboratory feasibility study on bower building behavior in order to provide direction for future research.

The Tramitichromis are locally referred to as “chisawasawa” and are important food fishes. They (all species in this genus were formerly placed in Lethrinops) are comprised of sand dwelling species that sift invertebrates and algae from the substrate

(Konings, 2001). During the breeding season, males aggregate and defend territories

(Stauffer and Kellogg, 1996). This lekking behavior results in the formation of large breeding arenas in which each male constructs a species-specific bower (spawning platform) out of sand (Stauffer and Kellogg, 1996; Kellogg et al., 2000). The populations show site fidelity for the breeding grounds, assortatively mate, and then disappear after breeding is over (Jay Stauffer, per. comm.). The Tramitichromis are diagnosed by the presence of a keel on the lower pharyngeal bone, the use of a figure eight courtship pattern, and the building of a cone-shaped bower by the males (see figures 3 and 4 in

Stauffer et al, 2002).

In practice, a taxonomist recognizes populations of organisms that exist in nature, and such populations can range from the local deme, the sympatric community of potentially interbreeding organisms, to the species taxon (Mayr, 1996; Stauffer and

McKaye, 2001). Before one can delimit species, one has to define what is meant by the term species. Wilson (1992) states that the search for a species concept that accurately

3 represents the diversity of life as the “Holy Grail” of the natural sciences (Stauffer and

McKaye, 2001). With the 22 species concepts listed by Mayden (1997), delimitation of species can be a difficult task (Stauffer et al., 2002). Part of the reason for the debate over a species definition is due to some biologists treating species as epiphenomena (here today, gone tomorrow), whereas others regard species as participants in the evolutionary process (Mayr and Ashlock, 1991; Stauffer et al., 1995).

Wiley (1978: 227) defines the evolutionary species concept as “…a single lineage of ancestor-descendant populations, which maintains its own evolutionary tendencies and historical fate.” The species is therefore a natural entity on an independent evolutionary trajectory, regardless of its mode of reproduction, or being extant or extinct (Stauffer et al., 2002). The problem with the evolutionary species concept is that it relies on a well- resolved phylogeny, and it is non-operational (Mayden, 1997; Stauffer and McKaye,

2001; Stauffer et al., 2002). The Lake Malaŵi cichlids lack a comprehensive and well- supported phylogeny (Stauffer et al., 2002). Thus, I will use the biological, morphological, and phenetic species concepts as surrogate concepts to diagnose the various species of the genus (Stauffer and McKaye, 2001; Stauffer et al., 2002). To detect evolutionary lineages, I will use reproductive isolation (from the biological species concept) and morphological/behavioral differentiation (morphological/phenetic species concepts) (Stauffer et al., 2002). Stauffer et al. (2002) state that reproductive isolation, behavioral traits, and morphological differentiation can be used for species delineation and phylogenetic reconstruction.

With the evolutionary species concept as my theoretical concept, and using the biological, morphological, and phenetic species concepts to delimit species, I need to first

4 clarify a few definitions. It is my belief that each species will have different morphology as they have different histories, roles in the ecosystem, and genetics. Morphological differentiation takes time, which is why shape analysis computer programs will be used to illustrate the minute differences in these recently radiated cichlid species. For the organisms with which I am working, reproductive isolation is a clear indication of species. Sexual selection has played an important role in the speciation of the Malaŵi fish fauna. In the Tramitichromis, this has manifested itself in the form of species- specific bower shapes. It would make sense that management strategies use bower data, as this is the driving force of speciation in this group of fishes (Stauffer, per. comm.).

Species misidentification could actually cause a collapse of a fishery.

On the topic of sympatric species, I consider sympatric species one in which they could come into contact with minimal effort. For example, we theoretically could obtain the same global positioning system (GPS) coordinates for two populations of fish that appear to occur at the same place at the same time, yet they are separated microallopatrically because one occurs in deep water (30 m) and the other in shallow water (less than 10 m). I would still call them sympatric because with minimal effort a fish from one group could swim into the other. Compare the previous example to two allopatric populations, one from each end of the lake. It would be nearly impossible for a fish from one population to swim to the other. Some of the species in this study are sympatric by the definition I have given above, but due to site fidelity for breeding grounds are actually allotopic, which means in different places. Again, these populations may only be a few hundred meters from one another, and individuals from one population could easily swim to the other. I recognized that ranking of allopatric

5 populations is problematic, and, following Stauffer and McKaye (2001), I reasoned that if two or more allopatric populations show the same phenotypic, behavioral, and genetic

(already done, McKaye et al., 1993) differences that are present in sympatric species, that they be described as separate species.

I have indicated that morphology is important tool that I will use to delimit species, but I do not believe that it can be the sole basis for species descriptions. I will use the coupling of the morphological data with other data such as bower shape, or anatomical features, for species descriptions. The lower pharyngeal bone is a highly variable interspecies, but not intraspecies character. Since the pharyngeal jaws are used to process food, it would make sense that the diet, or at least the access to the food source is different between the species based on the lower pharyngeal bone characteristics. The ecological role of each species still needs to be determined in full.

The rapid speciation of the African cichlid flock (including the Tramitichromis genus) is problematic for taxonomists because there is little morphological differentiation, and genetic tests are not able to conclusively separate species (Stauffer et al., 2002). Shape analysis has traditionally been used to delimit species (Stauffer et al.,

1993; Stauffer et al., 1997b). In addition, behavior has been shown to be important for the delimitation of Lake Malaŵi cichlid fish species (Stauffer et al., 1993; Stauffer et al.,

1995; Stauffer et al., 2002), and it may have played a role in sympatric speciation events

(Dominey, 1984; Smith and Todd, 1984; Turner and Burrows, 1995; Stauffer et al.,

2002). Female mate choice based on male behaviors (including bower building) can be a driving force in evolution (Barlow, 1991; Barlow, 1998; Clutton-Brock, 1991; Anderson,

6 1994; Johnsgard, 1994; Hogland and Alatalo, 1995; Stauffer and Kellogg, 1996, Stauffer et al., 2002), and supports the use of behavior in species descriptions.

Behavior is an extremely important and useful characteristic when working with a recently radiated group that has not accumulated morphological differences between species. Bower building is the manifestation of a behavioral trait (Stauffer et al., 1996;

Kellogg et al., 2000; Stauffer et al., 2002). Bower shape is a species-specific trait

(Stauffer et al., 1996) that has been used to diagnose species (Stauffer et al., 1996;

Stauffer and Konings, 2006).

Unfortunately, the behavioral component is missing in all previous descriptions of the fishes in this study. Konings (2001) provides some behavioral data on these species, but it is not linked with any other kind (morphological or genetic). McKaye et al. (1993) demonstrated congruence between bower shape (behavioral characteristic) and allozyme data (genetic data). I wanted to determine if there was also congruence between morphological data and bower shape (behavioral data) for the Tramitichromis as was done for Copadichromis (Stauffer et al., 1993). This would reinforce the use of behavioral data in species delimitation.

All of the type material used in this work came from collections made by C.

Christy during the mid 1920s to mid 1930s, which has been housed in the British

Museum of Natural History (BMNH). His collections and guidance allowed Trewavas

(1931) to diagnose many new species in the genus Lethrinops. Trewavas (1935) subsequently described Lethrinops intermedia. I reexamined Lethrinops intermedia,

Lethrinops brevis, Lethrinops lituris, and Lethrinops variabilis to ensure accurate

7 diagnosis and because there was suggestive evidence that each species was actually composed of more than one.

Eccles and Trewavas (1989) placed the above-mentioned species in a new genus,

Tramitichromis due to the presence of a keel on the lower pharyngeal bone. The remaining fish with a dark stripe from the nape to the caudal base, teeth in the lower jaw

3 to 5 in series, and a densely scaled caudal area were placed in the Taeniolethrinops. All remaining species were left in the Lethrinops. Tramitichromis was diagnosed by the presence of a keel on the lower pharyngeal bone as well as three or more rows of teeth extending to the end of the bone, which is rounded.

Chapter 2

Materials and Methods

Methods for species determination will follow (Stauffer et al., 1993; Stauffer et al., 1997b). Since the Tramitichromis spp. show site fidelity for breeding grounds and assortatively mate, populations are defined as aggregations of males during the breeding season at a particular locality, which have constructed species-specific bowers. In Lake

Malaŵi, a male was observed to breed with multiple females, each of which was collected after leaving the bower; then the male himself was collected. Only males from the same lek and their mates were preserved together. At other populations, only males were collected because of the absence of females. In addition, collections during the early 1980s were not done in this manner. During that time, fish from one area were collected and preserved together, but not with the strict “male and all his mates” technique. Live fish were collected by chasing them into a monofilament net (7 m x 1 m;

1.5 cm mesh) while SCUBA diving (Stauffer et al. 1993). A total of 738 fish was captured in the southeast arm of the lake and in Cobue between 1983 and 2002. Type specimens from the British Museum of Natural History (BMNH) were comprised of lakewide collections and examined to provide comparative references to fishes caught by

Stauffer, which reside in the Pennsylvania State University fish museum (PSU) (Fig 2.1).

= PSU Collection Lake Malawi = BMNH Collection Mwaya Karonga Vua Tanzania Deep Bay Mwanga? Zambia

Nkhata Bay Likoma

Cobue Mozambique Nkolongwe Malawi Fort Maguire

Chembe Village Monkey Bay, Monkey Bay Koma Village Fisheries Research Station Kanjedza Island Mazinzi Bay Golden Sands Swamp Nkhudzi Bay Otter Point Songwe Hill Liwonde

Chigubi Point Fort Johnston

Figure 2.1: Known localities of Tramitichromis species, both described and undescribed, in this study.

I examined each collection in the laboratory and reexamined the identity of the fish based on the keys in Eccles and Trewavas (1989). A subset of fish was chosen out of each jar at random, with the exceptions that some males, females, large, and small fish were in the subgroup. The lower pharyngeal bones of these fishes were dissected, and then used to verify the species. The number of fish examined per jar varied based upon the number of fish in the jar. Approximately 20% of the fish in each jar were observed in this way. If a jar was found to contain more than one species, then all of the fish were examined. Also, any fish with damaged or missing lower pharyngeal bones were eliminated, as this made accurate diagnosis/identification nearly impossible.

10 Twenty-four morphometric and fourteen meristic data points were collected on

738 fish (Table 2.1, Figs. 2.2, 2.3). Due to poor preservation or inaccurate species identification, some were eliminated and 611 fish (295 males, 316 females) remained.

All counts and measurements were made on the left side of the fish, except gill-raker counts. Gill-raker counts require bending the opercular and cutting part of the gular; thus, the right side was used to avoid damaging the measured side of the fish.

Morphometric values in tables were expressed as percent standard length (SL) or percent head length (HL) (Stauffer et al., 1997b).

Table 2.1: Morphometric and meristic measurements that were taken on each fish Morphometric Meristic Standard Length Dorsal Spines Head Length Dorsal Rays Snout Length Anal Spines Post-Orbital Head Length Anal Rays Horizontal Eye Diameter Pelvic Rays Vertical Eye Diameter Pectoral Rays Preorbital Depth Lateral Line Scales Cheek Depth Pored Scales Past Lateral Line Lower Jaw Length Cheek Scales Head Depth Gill Rakers on ceratobranchial Body Depth Gill Rakers on epibranchial Snout to Dorsal Fin Insertion Teeth Outer Row Left Lower Jaw Snout to Pelvic Fin Insertion Teeth Rows in Upper Jaw Dorsal Fin Base Length Teeth Rows in Lower Jaw Anterior Dorsal Fin to Anterior Anal Fin Anterior Dorsal Fin to Posterior Anal Fin Posterior Dorsal Fin to Anterior Anal Fin Posterior Dorsal Fin to Posterior Anal Fin Posterior Dorsal Fin to Ventral Caudal Fin Insertion Posterior Anal Fin to Dorsal Caudal Fin Insertion Anterior Dorsal Fin to Pelvic Fin Insertion

Posterior Dorsal Fin to Pelvic Fin Insertion

Caudal Peduncle Length

Least Caudal Peduncle Length

Figure 2.2: Illustration of the 24 morphometric data points.

Dorsal Rays Dorsal Spines

Lateral Line Scales

Teeth Rows Upper Jaw

Cheek Scales

Teeth Rows Lower Jaw

Pectoral Rays

Anal Rays Anal Rays Pelvic Rays Pored Scales Past Lateral Line Gill Rakers on Epibranchial

Gill Rakers on Ceratobranchial Figure 2.3: Illustration of the 14 meristic data points.

The fish that could not be identified using the key in Eccles and Trewavas (1989) were grouped together based on phenotypic characters. Keel shapes were not analyzed if the lower pharyngeal bone showed distinction. Differences in body shape were analyzed using sheared principal components analysis (SPCA) of the morphometric data

(Humphries et al., 1981; Bookstein et al., 1985; Stauffer, 1991; Stauffer et. al., 1993;

Stauffer et al., 1997b). This analysis restricts the size variation to the first component, thus subsequent components are strictly shape related (Bookstein et al., 1985, Stauffer et al., 1997b), and ordinates factors independently of a main linear ordination (Reyment et al., 1984; Stauffer et al., 1997b). This technique was used by Stauffer and Boltz (1989) to distinguish between two sympatric species of fish from Lake Malaŵi: Metriaclima

14 barlowi McKaye and Stauffer and Metriaclima xanstomachus Stauffer and Boltz

(Stauffer, 1991). Meristic differences were compared using principal components analysis (PCA) (Stauffer and Hert, 1992; Stauffer et al., 1997b). The correlation matrix was factored in all principal component analyses of meristic data, while the covariance matrix was factored in the calculation of all sheared principal components of the morphometric data (Stauffer and Hert, 1992; McKaye et al., 1993; Stauffer et al., 1997b).

Differences among species were illustrated by plotting the sheared components of the morphometric data against the principal components of the meristic data in order to maximize the amount of separation (Stauffer and Hert, 1992; Stauffer et al., 1997b). The second or third sheared principle component scores of the morphometric data (SHRD

PC2 and PC3 respectively) were plotted against the first or second principal component scores of the meristic data (PC 1 or PC 2).

For minimum polygon clusters that overlapped, I determined if they were significantly different. If the mean multivariate scores of the clusters were significantly different along one axis, independent of the other axis, a Duncan’s multiple range test

(p<0.05) was used to determine which clusters differed from each other (Stauffer et al.,

1997b). If, in fact, the clusters were not significantly different along one axis independent of the others, then a MANOVA, in conjunction with a Hotelling-Lawley trace, was used to determine whether the mean multivariate scores of clusters formed by the minimum polygons of the PCA scores were significantly different (p<0.05) (Stauffer et al., 1997b). Polygon clusters of different species may overlap by three quarters and still be significant. Ideally, polygon clusters should minimally overlap if at all.

15 I compared bower shape taken in the field for three of the previously undescribed species to determine if differences in bower shape among these three species supported differences indicated by morphological data. The bowers were measured while using

SCUBA (self contained underwater breathing apparatus) equipment. Bower measurements were taken according to Stauffer et. al. (1993) and include: width of base, slope length, outside top diameter, inside top diameter, and bower height (Fig. 2.4). Two sets of measurements were taken, the second set at 90 degrees from the first. Bower shape was analyzed following Stauffer et al. (1993). Differences in bower shape were analyzed using SPCA (see discussion above) (Humphries et al., 1981; Bookstein et al.,

1985; Stauffer et al., 1993). Differences are illustrated by plotting the sheared components of the data in order to illustrate differences in shape among the bowers

(Stauffer et al., 1993). The clusters formed by each taxa were analyzed using MANOVA

(Stauffer et al., 1993). Differences among dimensions were tested by using a MANOVA in conjunction with Duncan’s multiple range test (Stauffer et al., 1993). The only difference in the analysis is that only shape variables were analyzed; thus the SHRD PC2 was plotted against the SHRD PC3 (Stauffer et al., 1993).

C D

B F E

Chapter 3

Taxonomic Character Analysis

As stated in the introduction, recently radiated groups of fish may not have had the time to accumulate observable morphological differences. For this work, I have employed the principal components analysis to maximize small differences in body shape. When using this method, it becomes extremely important to ensure landmarks for data collection are consistent among fishes.

The description of the morphometric measurements follows (Fig. 2.2). The standard length is from the tip of the snout to the hypural plate as evidenced by the fold of the caudal fin where it meets the body (where the “meat” of the fish stops). The head length is from the tip of the snout to the notch in the opercle. The snout length is from the tip of the snout to the anterior orbit. The post-orbital head length is from the posterior orbit to the notch in the opercle. The horizontal eye diameter is the measurement of the orbit without stretching. Vertical eye diameter is the same except in the vertical direction. The preorbital depth is the area between the anterior orbit and the orbital bone.

The cheek depth is the area from the bottom of the orbit to the ridge formed around the area where the cheek scales end. The lower jaw length is the area between the anterior end of the jaw and the fleshy “v” formed on the ventral side of the fish between the gills.

The head depth is a perpendicular line to the horizontal plane of the fish, the bottom of which is the origin of the “v” described above. The body depth is a line perpendicular to the horizontal plane of the fish with its origin at the dorsal fin origin.

18 The following morphometric measurements are between the following landmarks.

The snout to dorsal-fin origin is the distance from the tip of the snout to the origin of the dorsal fin. The snout to pelvic-fin origin is the distance from the snout to the origin of the pelvic fin. The dorsal-fin base length is the area between the origin and termination of the base of the dorsal fin. Anterior dorsal fin to anterior anal fin, anterior dorsal fin to posterior anal fin, posterior dorsal fin to anterior anal fin, the posterior dorsal fin to the posterior anal fin, the posterior dorsal fin to ventral caudal fin insertion, the posterior anal fin to the dorsal caudal fin insertion, the anterior dorsal fin to pelvic fin insertion, the posterior dorsal fin to the pelvic fin insertion are the distances between the respective points.

The caudal peduncle length is the distance from a vertical line formed between the termination of the dorsal and anal fin bases to a fold made where the caudal fin starts as the body (the meat) of the fish stops. The least caudal peduncle depth is the smallest vertical distance anywhere between the vertical line formed between the termination of the dorsal and anal fin bases to the fold made where the caudal fin starts and the body of the fish stops.

Meristic (Fig. 2.3) include the dorsal spines, which are hard cactus like spines in the dorsal fin. The dorsal rays are soft and start after the dorsal spines stop and continue to the end of the fin. Anal spines are found at the anterior portion of the fin and have the same hard and sharp feel as the dorsal spines. The anal rays start after that, but the last two are counted as one if they have the same origin. Pelvic rays are found on the pelvic fin. Pectoral rays are found on the pectoral fin and do not include the hard outer edge.

19 Lateral line scales are found along the lateral line. They are pored and start behind the head along the upper lateral line. They are counted moving posteriorly, and when the end of the top lateral line is reached, the line is traced down to the lower one and then the count continues. The count stops at the hypural plate. Pored scales past the lateral line are found where the lateral line scales stop and are past the fold formed by the insertion of the caudal fin (i.e., hypural plate). Cheek scales are the rows of scales moving ventrally from the eye.

Gill rakers on the ceratobranchial are the rakers on the lower portion of the gill below the notch (Fig 2.3). The raker that separates the upper and lower limb is not counted. Gill rakers on the epibranchial are above the notch, not including the raker found in the notch.

Teeth in the outer row of the left lower jaw are counted from the midline of the fish moving toward the side. As soon as the tooth row begins to curve behind the other rows, the count stops as different rows are blending together. Teeth rows in the upper jaw are the number of rows from the front to the back. Slight bumps or immature teeth are counted also. Teeth rows in lower jaw refers to the rows of teeth from the anterior portion of the lower jaw moving toward the posterior. The best place to observe the rows is along the midline of the jaw. Small immature teeth as well as bumps of the teeth rows are counted.

The shape of the lateral view of the lower pharyngeal bone is highly diagnostic.

The angle of inclination as well as the depth and length of the keel are important characters. They are different enough between species not to need to be measured.

Judging the angle is sufficient. When viewed dorsally, the number of teeth rows of the

20 lower pharyngeal bone differs between species. The number ranges from two to six.

Care must be taken as these end teeth can be damaged or pushed out of place during the preservation process. One has to trace the rows across at the origins of the teeth.

The anterior teeth on the lower pharyngeal bone can have a cylindrical shape or have a cusp. One species seemed to have a minute cusp. The teeth then can point in any particular direction, which varies depending on species. The posterior lower pharyngeal bone teeth can vary much in the same way, but they either have a cusp or are molariform; suggesting the fish eat snails. The posterior lower pharyngeal bone teeth may point in various directions also.

Courtship behavior consists of the male swimming in a figure 8 pattern. Males also build cone shaped bowers with a depression in the top that serves as a spawning platform. Other genera have circular courtship patterns and a range of bower shapes from flat to multiple mounds. Certain genera build bowers with a rock, without a rock, or on top of rocks. Height of the bower may change, but the shape does not (Fig 2.4). The width of the base is the distance along the bottom. The slope is the side measurement from the base to the rim. The outside diameter is the distance across the top portion of the bower (the bowl). The inside top diameter is the actual bowl or depression diameter.

The lip length is the small area between the top edge of the bower and the bowl formed on the top platform. The height is a line perpendicular to the base length.

Chapter 4

Taxonomy of Tramitichromis

Tramitichromis Eccles and Trewavas

Lethrinops Regan 1921. Regan. 1922. The cichlid fishes of Lake Nyassa. Proc. Zool.

Soc. Lond, (for 1921): 675-727.

Lethrinops Trewavas. 1931. A Revision of the Cichlid Fishes of the Genus Lethrinops,

Regan. Annual Magazine of Natural History, Ser. 10, 7: 133-152.

Tramitichromis Eccles and Trewavas. 1989. Malawian Cichlid Fishes: The

Classification of Some Haplochromine Genera. Lake Fish Movies, Herten,

Germany, pp 335.

Type Species – Tramitichromis brevis (Boulenger) (Fig. 4.1)

Diagnosis – Currently, this genus is monotypic, but Snoeks (2004) suggests that it will eventually include several undescribed species, exhibits two distinct characteristics:

1) a complete single dark lateral band that runs from just below the dorsal fin insertion to the middle of the caudal fin (Fig 4.1a) in conjunction with a keel on the lower pharyngeal bone, and 2) the building, by males during lekking, of a cone shaped bower which contains a rock.

22 Etymology – Tramitichromis, from the Greek, meaning a departure of the pharyngeal jaws from the usual range of structure (Eccles and Trewavas, 1989).

Tramitichromis brevis (Boulenger) (Fig. 4.1)

Tilapia brevis Boulenger, 1915, Catalogue of African Freshwater Fishes III. 526 pp. 351

fig. London, B.M.N.H.

Haplochromis brevis Regan. 1922. The cichlid fishes of Lake Nyassa. Proc. Zool. Soc.

Lond, (for 1921): 675-727.

Lethrinops brevis Boulenger. Trewavas, E. 1931. A Revision of the Cichlid Fishes of

the Genus Lethrinops, Regan. Annual Magazine of Natural History, Ser. 10, 7:

133-152.

Tramitichromis brevis (Boulenger). Eccles, David H., Ethelwynn Trewavas. 1989.

Malawian Cichlid Fishes: The Classification of Some Haplochromine Genera.

Lake Fish Movies, Herten, Germany, pp 335.

Material Examined – PSU 4089, 24 fish, February 18, 2002, Cobue (Figs. 4.1,

4.2).

Diagnosis – Tramitichromis brevis retains a complete dark lateral band that runs from just below the dorsal fin insertion to the middle of the caudal fin (Fig. 4.1a). This trait can be seen on live as well as preserved specimens. Inspection of the lower pharyngeal bone confirms its placement within Tramitichromis, with the “anterior blade

[of the keel] steeply inclined ventrally” (Fig. 4.1e) (Eccles and Trewavas 1989: 256).

The anterior teeth are cylindrical with the ends pointing backwards (Fig. 4.1c). Posterior

23 teeth are enlarged up to two rows anteriorly beyond the last (Fig. 4.1d). I found no evidence of variation, although my samples are based on one collection.

25 Description – Jaws isognathous (Fig. 4.1a); teeth on upper jaw in 2-4 rows; teeth on lower jaw in 4-5 rows; 9-14 teeth in outer row of left lower jaw. Dorsal fin with 15-16 spines and 10-12 rays; pectoral fin with 15-17 rays; anal fin with 3 spines and 8-10 rays.

Lower pharyngeal bone triangular in outline with a deep notch posteriorly (Fig. 4.1f).

Scales along side ctenoid with 31-33 in lateral-line series. First gill arch with 6-8 rakers on the ceratobranchial, 3-4 on the epibranchial with 1 between the epibranchial and ceratobranchial (Table B.1).

Live coloration was not recorded. Preserved pattern consists of a dark lateral band that runs from just below the dorsal fin insertion to the middle of the caudal fin.

Distribution – The “type” material accounts for this species in the northern (Vua) and southern (bar to Fort Maguire) ends of the lake (Fig. 4.2). Trewavas (1931) lists only the Fort Maguire collection as the types, but indicates she used the Vua population for her description (which would make them paratypes). With the additional PSU material from

Cobue, which is close to the middle of the lake, it probably occurs throughout. Konings describes T. brevis as “a common cichlid, which is found all around the lake” (Konings

2001, pg 287).

= PSU Collection Lake Malawi = BMNH Collection

Vua Tanzania

Zambia

Cobue Mozambique

Malawi Fort Maguire

Figure 4.2: Location of the collections of Tramitichromis brevis, PSU 4089.

Discussion – The analysis of T. brevis was limited to a single collection from

Cobue, Mozambique (Fig. 4.2). Unfortunately, I was not able to obtain the T. brevis type specimens from the British Museum to which I could compare as they were not in the shipment of type material, and numerous requests for them did not produce the fish. The

T. brevis specimens at the museum are not labeled as types, which could have caused the confusion on the part of the museum; they would only send specimens labeled as such.

Clearly the species was diagnosed based on BMNH collection 1930.1.31.46-49 and possibly BMNH 1935.6.14.2067-2068 (Trewavas 1931). Either way, I am certain of

27 their proper identification due to the presence of the lateral band and the shape and dentition of the lower pharyngeal bone. Comparison to the type specimens of the other members of the Tramitichromis using the principle components analysis revealed a distinct clustering of T. brevis from all of the other Tramitichromis species when the first principal components of the meristic data are plotted against the sheared second principle components of the morphometric data (Fig. 4.3). The minimum polygon cluster formed by T. brevis is significantly different from the other clusters (p<0.05). The clusters were found to be significantly different along both the SPCA 2 (morphometric data) and the

PC 1 (meristic data) axis independent of each other. The variables that had the highest loadings on the sheared second principal components were caudal peduncle length (-

0.44418), vertical eye diameter (0.37196), and lower jaw length (0.34474); while those with the highest loadings on the first principal components of the meristic data were lower gill rakers (0.34695), cheek scales (0.30843), and lateral line scales (0.28821).

The fish from Cobue were taken from a breeding arena where the males had built cone shaped bowers with a rock in them. Ad Konings, an avid Lake Malaŵi diver and cichlid expert, confirms T. brevis breeding in a cone shaped bower with a rock in it (per comm; see Konings 2001 pg 285 for picture). Neither Stauffer nor Konings has observed

T. brevis breeding in an environment other than one with rocks and sand. Tramitichromis brevis seems to be the only member of the genus to breed with the use of a rock in the bower.

This use of the rocky sand for breeding most likely was a secondary adaptation by

T. brevis. I base this statement on the fact that no other members of the genus use a rock

28 in their bowers, nor do any of the Lethrinops with which this group is closely related.

Although T. brevis may be found at the same locality as other members of its genus, it is separated from them microallopatrically (during spawning) due to its preference for the rocky/sand interface; thus, an effective pre-mating isolation mechanism.

Tramitichromis brevis is the type species for the genus; however, due to the following dissimilarities with the other members currently found within this genus, I have decided to remove them and place them in a new genus. The reasons for this action are summarized below:

1. Tramitichromis brevis is the only species to posses a complete dark lateral band

that runs from just below the dorsal fin insertion to the middle of the caudal fin,

which is often reflective of phylogeny (Eccles and Trewavas, 1989).

2. Tramitichromis brevis clusters separately from the other former Tramitichromis

species in a plot of the sheared PC 2 and PC 1, indicating a different body shape

(e.g. shorter caudal peduncle, longer vertical eye diameter, longer lower jaw,

fewer gill rakers on outer ceratobranchial).

3. Tramitichromis brevis is the only species within the genus in which the males

construct a bower that includes a rock.

0.15

0.1

0.05

T. brevis T. variabilis 0 T. lituris T. trilineata T. intermedius

SPCA 2 (morphometric data) -0.05

-0.1

-0.15 -2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 PC 1 (meristic data)

Figure 4.3: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Tramitichromis brevis (N = 24) PSU 4089; and the other described Tramitichromis species type material from the British Museum, T. variabilis (N = 22) BMNH 1930.1.31.14-20; BMNH 1930.1.31.1-2; BMNH 1930.1.31.4- 13; BMNH 1930.1.31.3; T. lituris (N = 9) BMNH 1930.1.31.21-23; BMNH 1930.1.31.24-28; BMNH 1930.1.31.45; T trilineata (N = 1) BMNH 1930.1.31.76; T. intermedius (N = 6) BMNH 1935.6.14.2081-2084; BMNH 1935.6.14.2085.

Snoeks (2004) indicates some differences between populations of this species, but no formal descriptions were made. This species should be sampled lake wide along with making in situ behavioral observations.

Apetra, n. gen.

30 Lethrinops Regan 1921. Regan. 1922. The cichlid fishes of Lake Nyassa. Proc. Zool.

Soc. Lond, (for 1921): 675-727.

Lethrinops Trewavas. 1931. A Revision of the Cichlid Fishes of the Genus Lethrinops,

Regan. Annual Magazine of Natural History, Ser. 10, 7: 133-152.

Tramitichromis Eccles and Trewavas. 1989. Malawian Cichlid Fishes: The

Classification of Some Haplochromine Genera. Lake Fish Movies, Herten,

Germany, pp 335.

Type Species – Apetra lituris (Trewavas) (Fig. 4.4)

Diagnosis – This genus comprises ten species that have a keel on the lower pharyngeal bone without a complete single dark lateral band that runs from just below the dorsal fin insertion to the middle of the caudal fin, and males that build a cone shaped bower without a rock on open sand during lekking. It differs from the closely related

Tramitichromis in that none of the members exhibit the complete dark lateral band described above. Instead, the species may have one or more broken lines, spots, horizontal elements, or combinations thereof.

Etymology – Apetra, from the Greek, meaning without a rock to indicate the bowers built by male members of this genus, which do not contain a rock like the phenotypically similar Tramitichromis.

Apetra lituris (Trewavas) (Fig. 4.4)

31 Lethrinops lituris Trewavas. 1931. A Revision of the Cichlid Fishes of the Genus

Lethrinops, Regan. Annual Magazine of Natural History, Ser. 10, 7: 133-152.

Tramitichromis lituris (Trewavas) 1989. Eccles, David H., Ethelwynn Trewavas. 1989.

Malawian Cichlid Fishes: The Classification of Some Haplochromine Genera.

Lake Fish Movies, Herten, Germany, pp 335.

LECTOTYPE. – BNMH 1930.1.31.21, adult male, 125.2 mm, Karonga,

Lake Malwai, Malawi, Africa (Fig. 2.5) (I designated this lectotype).

PARALECTOTYPES. – BMNH 1930.1.31.22-23, 2 fish, Karonga;

BMNH 1930.1.31.24-28, 5 fish, Vua; BMNH 1930.1.31.35-44, 24 fish,

Mwaya (Fig. 4.6).

Diagnosis – Apetra lituris is a medium-sized fish attaining a length of 140mm

(Eccles and Trewavas, 1989). The pattern of this species is not very distinctive. It consists of a dark line along the upper lateral line, and may include darker horizontal elements along the “bars”. Various elements of it may or may not be preserved (Fig.

4.4a). This pattern, along with some other traits it shares with the remaining species, are yet to be discussed. The upper edge of the blade of the lower pharyngeal bone is inclined downwards at less than 45o to the plane of the toothed surface (Fig. 4.4d) (Eccles and

Trewavas, 1989). The majority of the anterior teeth do not have a cusp, and the ends are

turned backwards slightly at an angle of up to but not more than 45o (Fig. 4.4f). Outside

of the lectotype, which was the only individual without a damaged lower pharyngeal

32 bone, the anterior teeth of most speciemens are not pointed backward at all and appear to point almost straight up at about 85o (Figs. 4.4e, 4.5a and b). The posterior teeth do have

a cusp, are pointed forwards, and are not enlarged, except for the posterior row (Fig.

4.4e).

Description – Jaws isognathous (Fig. 4.4a); teeth on upper jaw in 4 rows in lectotype, 2-4 rows in paralectotypes; teeth on lower jaw in 5 rows in lectotype, 4-5 rows in paralectotypes; 13 teeth in outer row of left lower jaw in lectotype, 13-16 in

35 paralectotypes. Dorsal fin with 15 spines in lectotype, 15-16 in paralectotypes; 11 rays in lectotype, 10-12 in paralectotypes; pectoral fin with 16 rays in lectotype, 14-16 in paralectotypes; anal fin with 3 spines in both the lectotype and paralectotypes, 9 rays in the lectotype, 8-10 in paralectotypes. Lower pharyngeal bone triangular in outline with a notch in the posterior (Fig. 4.4c). Scales along side ctenoid with 32 in lateral-line series in the lectotype, 31-34 in paralectotypes. First gill arch with 8 rakers on ceratobranchial in lectotype, 7-10 in paralectotypes, 4 on epibranchial in lectotype, 2-4 in paralectotypes,

1 between the epibranchial and ceratobranchial (Table B.2).

Live coloration has not been recorded. Preserved pattern consists of a dark line along the upper lateral line, and may include darker elements along the “bars”. Various elements of it may or may not be preserved. The type material is rather faded, but some elements of the broken lines may be observed.

Distribution – The type material comes from the northern end of the lake at

Karonga, Mwaya, and Vua (Fig. 4.6). It is unknown at this time how far the range of this species extends south. It did not appear in PSU collections from the southern ends of the lake, and may be a northern species.

= PSU Collection Lake Malawi = BMNH Collection Mwaya Karonga Vua Tanzania

Zambia

Mozambique

Malawi Fort Maguire (not A. lituris)

Figure 4.6: Localities of Apetra lituris: Karonga BMNH 1930..31.21-23; Vua BMNH 1930.1.31.24-28; Mwaya BMNH 1930.1.31.35-44.

Discussion – There were not a lot of specimens of this species in the study. The type material, which until now consisted of two species and did not have a lectotype (I declared one), comes from four localities. What puzzles me is Eccles and Trewavas

(1989) indicate that the type material locality was not specified, but they place it in two locations in the southeast arm. From the tags and information catalogued in the British

Museum, the localities of the type material are Karonga, Vua, Mwaya, and Fort Maguire

(Fig. 2.7). Only Fort Maguire is in the southeast arm (this is actually a different species).

In addition, when I looked at the fish from Mwaya, there was one group of all females,

37 and one mixed sex group. Sometimes SPCA will show differences between males and females when just they are plotted, but when they are compared to another species, the males and females cluster together. On my graph, there is a clear separation of the all female group from this locality from the rest of the fish, including the other Mwaya fish, when the first principal components of the meristic data are plotted against the sheared second principle components of the morphometric data (Fig. 4.7). The poorly preserved all female Mwaya group was significantly different (p<0.05) along the PC 1 (meristic data) axis. I do not think this is accurate because that jar of fish was poorly preserved and in poor condition, missing many scales, having torn and damaged fins, and missing spines and post lateral line scales in many cases. All the fish were flimsy in addition to the damage listed above, which probably produced these results. For that reason, I did not use them in the multivariate analysis. The other Mwaya group was better, but it contained 24 individuals instead of the 10 indicated by the BMNH number. They all appeared to be the correct species, so I used them. I have no idea where else they could have come from, and I have found a few other cases (with different species) of more fish in the jar then the number indicates, but they were all members of the same species. One other confounding factor is that there is only one fish from Fort Maguire, and it does not cluster with the group when the first principal components of the meristic data are plotted against the sheared second principle components of the morphometric data (Fig. 4.8).

The plot of the minimum polygon cluster of the Fort Maguire fish was found to be significantly different (p<0.05) along both the SPCA 2 (morphometric data) and PC 1

(meristic data) from all the other clusters. It does, however, cluster with one of the

Karonga fish from the north. The Fort Maguire fish is a separate species (to be discussed

38 diagnosed below). The outlier from Karonga has a lower pharyngeal bone that identifies it as A. lituris, but for some reason it does not cluster with the group. Variables that had the highest loadings on the sheared second principal components were lower jaw length

(0.40724), horizontal eye diameter (0.34942), and cheek depth (0.34540); while those with the highest loadings on the principal components of the meristic data were dorsal rays (0.39201), dorsal spines (-0.33281), and anal rays (0.26013).

This species appears to be from the northern end of the lake as there were not any specimens in the PSU or BMNH collections from the south. An extensive survey of the northern end of the lake to resolve the distribution of this species is needed.

0.15

0.1

0.05 Karonga Fort Maguire Vua Mwaya all female group Mwaya mixed sex group 0 SPCA 2 (morphometric data) 2 (morphometric SPCA

-0.05

-0.1 -2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 PC 1 (meristic data)

Figure 4.7: Plot of the sheared second principal components (morphometric data) and the first factor scores (meristic data) of BMNH Apetra lituris: Mwaya (N = 24) BMNH 1930.1.31.35-44; Fort Maguire (N = 1) BMNH 1930.1.31.45; Vua (N = 5) BMNH 1930.1.31.24-28; Karonga (N = 3) BMNH 1930.1.31.21-23. The damaged all female group from Mwaya (N = 13) appear as separate cluster, BMNH 1930.1.31.29-34.

0.08

0.06

0.04

0.02

Karonga Fort Maguire -0.02 Vua Mwaya -0.04 SPCA 2 (morphometric data) 2 (morphometric SPCA -0.06

-0.08

-0.1

-0.12 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 PC 1 (meristic data)

Apetra intermedia (Trewavas) (Fig. 4.9)

Lethrinops intermedia Trewavas. 1935. A Synopsis of theCichlid Fishes of Lake Nyasa.

Annual Magazine of Natural History, Ser. 10, 16: 65-118.

41 Tramitichromis intermedius (Trewavas). 1989. Eccles, David H., Ethelwynn Trewavas.

1989. Malawian Cichlid Fishes: The Classification of Some Haplochromine

Genera. Lake Fish Movies, Herten, Germany, pp 335.

Material Examined – LECTOTYPE BMNH 1935.6.14.2081, Southwest Arm and

Fort Johnston; PARALECTOTYPES BMNH 1935.6.14.2082-2084, 4 fish, Southwest

Arm and Fort Johnston; PARALECTOTYPE BMNH 1935.6.14.2085, Monkey Bay;

Other material examined: PSU 4092, 8 fish, April 9, 1983, Chembe Village, Cape

Maclear; PSU 4104, 5 fish, March 25, 1995, Chembe Village Swamp; PSU 4144, 3 fish,

September 8,. 1983, Chembe Village Swamp; PSU 4147, 1 fish, September 8, 1983,

Chembe Village Swamp; PSU 4156, 1 fish, April 18, 1984, Chembe Village Swamp;

PSU 4117, 2 fish, September 6, 1983, Golden Sands Swamp, Cape Maclear; PSU 4081,

10 fish, April 23, 1991, Chirombo Bay, Kanjedza Island; PSU 4101, 9 fish, April 22,.

1991, Chirombo Bay, Kanjedza Island; PSU 4107, 12 fish, February 26, 1991, Chirombo

Bay, Kanjedza Island; PSU 4110, 18 fish, February 26, 1991, Chirombo Bay, Kanjedza

Island (Fig. 4.11).

Diagnosis – Apetra intermedia is distinguished from A. lituris by the possession of three dorsolateral spots, a small decurved keel compared to a steeply inclined one, forward facing anterior pharyngeal bone teeth with a cusp compared to cylindrical backward facing teeth, and molariform posterior pharyngeal teeth which are absent in A. lituris. Apetra intermedia is easily distinguishable from the rest of the species in this genus by the presence of three dark dorsolateral spots in live and preserved specimens

42 (Fig. 4.9a). Breeding males in full color might not exhibit the dorsolateral spots due to the color pigments covering up the melanophores that cause the pattern. The first is located under the insertion of the dorsal fin, the second is positioned beneath the first third of the dorsal fin, and the third is found around the insertion of the last few rays of the dorsal fin. The identity of this species can be confirmed by examination of the lower pharyngeal bone. According to Eccles and Trewavas (1989), the anterior blade of the bone is “decurved,” (Fig. 4.9d) and it has a most similar appearance with T. trilineata. A comparison with the drawing of the side view of the lower pharyngeal bone in

Trewavas’s 1935 work shows an exaggeration of the keel, which is actually much more conservative (Fig 4.9, see Trewavas, 1935, Fig. 11). The anterior teeth have a cusp, and the ends are pointed forward (Fig. 4.9f). I did not observe any variation in the anterior teeth with regard to type or position. The posterior teeth are molariform (Fig 4.9c), suggesting that it feeds on snails, and the number of enlarged teeth/rows seems to vary with the individual (Fig. 4.10).

Description – Jaws isognathus (Fig. 4.9a); teeth on upper jaw in 3 rows in lectotype and paralectotypes, 2-4 in some specimens from Chembe Village and Kanjedza

Island; teeth rows on lower jaw in 4 rows in lectotype, 4-5 in paralectotypes, as few as 2 in some specimens from Chembe Village, as few as 3 in some specimens from Kanjedza

Island; 14 teeth in outer row of left lower jaw in lectotype, 12-16 in paralectotypes, as many as 19 in some specimens from Chembe Village, 10-21 for specimens from

Kanjedza Island. Dorsal fin with 15 spines in the lectotype and paralectotypes, 14-16 in some specimens from Chembe Village and Kanjedza Island, 10 rays in the lectotype, and

45 10-11 rays in the paralectotypes, 9-13 in some specimens from Chembe Village; pectoral fin with 15 rays in the lectotype, and 14-16 in the paralectotypes, 17 in some individuals from Kanjedza Island; anal fin with 3 spines in all the types, 9 rays in the lectotype, 8-9 in the paralectotypes, as many as 10 in some specimens from Kanjedza Island. Lower pharyngeal bone triangular in outline (Figs. 4.9c, 4.10). Scales along side ctenoid with

32 in lateral-line series in the lectotype, 31-32 in paralectotypes, as many as 33 in some specimens from Chembe Village and Kanjedza Island, as many as 34 in one individual from Golden Sands Swamp . First gill arch with 8 rakers on the ceratobranchial in the lectotype, 7-10 in paralectotypes, as many as 12 in some specimens from Chembe

Village, as many as 15 in some specimens from Kanjedza Island, 4 on the epibranchial in lectotype, 2-4 in paralectotypes, with 1 between the epibranchial and ceratobranchial

(Table B. 3).

Live coloration was recorded for males at Kanjedza Island, PSU 4107. Males had sides with a white ground coloration with green, blue, and yellow highlights; 7 light blue/gray vertical bars; three spots which cannot be seen on males on bowers; dark dorsally fading to white ventrally. On the head, the interorbits and dorsal to eye region was a dark gray; ventral to eye was a florescent blue/green with a bright orange gular.

The dorsal fin was dark gray with a white marginal bar and orange lappets; the membranes between the rays had orange spots. The caudal fin was blue with yellow and orange vermiculations. The anal fin was pink with 15-20 beige ocelli. The pelvic fins had gray spines and rays with some orange melanophors. The pectoral fins were clear.

46 Preserved specimens have a series of three dark dorsolateral spots.

Distribution – The lectotype came from the southwest arm to Fort Johnston as did most of the paralectotypes except for one from Monkey Bay in the southeast arm (Fig

4.11). The PSU fish came from Golden Sands Swamp and Chembe Village Swamp, which are found on Cape Maclear, as well as Kanjedza Island, which is in the southeast arm of the lake. An additional specimen (not in this study) is catalogued at the British

Museum from Nkhata Bay, which is on the west side of the northern half of the lake (see

Fig. 2.1). Most likely this species is found lakewide.

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi

Chembe Village Monkey Bay

Kanjedza Island Southwest Arm Golden Sands Swamp

Fort Johnston

Discussion – From the cotypes of Apetra [Tramitichromis] intermedia Eccles and

Trewavas (1989) declared a lectotype. I noted one of the type specimens was “different,” and when I examined their work, I saw we were in concordance as they indicate the fish with the standard length of 91mm was of a different species. They cited the tooth pattern as their clue as to the different species status, while I noted differences in its external morphology.

48 The locality for the type material is simply listed as “South,” (including the lectotype), and Monkey Bay (Fig 4.11). It is unfortunate that there is nothing more specific than “South” as it makes it impossible to go observe and/or collect topotypes.

The alternative would have been to use the Monkey Bay specimen, but naming an individual from a group is advantageous, as they are more likely to be the same species as opposed to two different populations. The fish used in this study from the PSU collections came from Golden Sands Swamp, Fisheries Research Station, Chembe

Village, and Kanjedza Island.

Occasionally the type material may consist of more than one species. Upon review of the Apetra intermedia from the British Museum, they all appear to be the same species, or at least I did not observe any phenotypic differences (outside of the 91mm SL individual discussed above). When the types are compared to all the other collections of

A. intermedia, all the fish cluster together except for two outlying individuals from

Chembe Village when the first principal components of the meristic data are plotted against the sheared second principle components of the morphometric data (Fig. 4.12).

These fish are identified as A. intermedia, but are aberrant in their snout length and preorbital depth by having much shorter measurements for these characters than similar sized fish. The minimum polygon clusters formed by the different populations are not significantly different (p<0.05).

0.25

0.2

0.15

0.1 Lectotype Paralectotypes 0.05 Chembe Village Golden Sands Swamp Kanjedza Island 0 SPCA 2 (morphometric data)

-0.05

-0.1

-0.15 -4 -3 -2 -1 0 1 2 3 4 PC 1 (meristic data)

Many times distant populations can appear to be different on the SPCA graphs, but when the intermediate populations are plotted, the two distinct shapes of the populations grade into one another (Bowers and Stauffer, 1993; Stauffer et al., 1997). I did not observe this with this species. The Chembe Village and Kanjedza Island populations were tested because they were distant from each other and there were a good number of each population. I did not observe any phenotypic differences between them, and I wanted to see if the SPCA graph confirmed this. When they were plotted, there was no separate clustering of the two separate populations, although there were the same

50 two outliers noted in earlier graphs (Fig. 4.13). The minimum polygon clusters formed by the two populations were not significantly different (p<0.05). This would indicate the two populations belong to the same species, and that there are no phenetic differences.

0.25

0.2

0.15

0.1

Chembe Village 0.05 Kanjedza Island

0 SPCA 2 (morphometric data) 2 (morphometric SPCA

-0.05

-0.1

-0.15 -4 -3 -2 -1 0 1 2 3 4 PC 1 (meristic data)

One of the most interesting discoveries I made involving these fish is that they do not seem to vary in their anterior pharyngeal teeth. Many of the other species have some sort of variation in these. Either this species does not have the genetic plasticity to allow such variation (doubtful), or this trait is under strong selective pressure (probable). These anterior teeth must be strongly tied to its diet, and any variation in them would mean

51 difficulty eating. There did seem to be some variation in the shape of the keel when viewed from the lateral. I was not able to correlate this with any other differences, however.

Apetra variabilis (Trewavas) (Fig 4.14)

Lethrinops variabilis Trewavas. 1931. A Revision of the Cichlid Fishes of the Genus

Lethrinops, Regan. Annual Magazine of Natural History, Ser. 10, 7: 133-152.

Tramitichromis variabilis (Trewavas) 1989. Eccles, David H., Ethelwynn Trewavas.

1989. Malawian Cichlid Fishes: The Classification of Some Haplochromine

Genera. Lake Fish Movies, Herten, Germany, pp 335.

LECTOTYPE. – BMNH 1930.1.31.4, adult male, 104.4 mm, Lake Nyasa South

[former name of Lake Malaŵi], Malaŵi, Africa (Fig. 4.14)(I designated this lectotype).

PARALECTOTYPES. – BMNH 1930.1.31.5-13, 10 fish, Lake Nyasa Souh;

BMNH 1930.1.31.3, 1 fish, Monkey Bay (Fig. 4.15).

Diagnosis – Apetra variabilis is distinguished from A. lituris by a steeply inclined keel (at least 45 degrees) and the swollen anterior of the lower pharyngeal bone with six rows of teeth (compared to four). It is distinguished from A. intermedia by the lack of three dorsolateral spots, lack of molariform teeth on the posterior lower pharyngeal bone, and a steeply inclined keel (45 degrees) compared to the small decurved keel in A.

52 intermedia. The body pattern consists of an incomplete oblique band from the nape, intersecting the lateral line below the posterior part of the spiny dorsal (Eccles and

Trewavas, 1989) (Fig. 4.14a). The pattern appears to contain a single break in the solid line, with a ventral anterior shift at the break as the line continues toward the posterior of the fish (Fig. 4.14a; see Fig. 2B of Trewavas, 1931). It may be possible to observe the pattern of dots noted by Trewavas (see Trewavas, 1931, Fig. 2A). The lower pharyngeal bone has the steepest angled keel with regard to the bone itself; being inclined at least 45 degrees (Figs. 4.14d). The anterior teeth are long and cylindrical, curving backwards

(Fig 4.14e). The posterior teeth have a cusp, and the ends are pointed forward along the back row, previous rows pointed backward (Fig. 4.14f). The teeth seem to transition from the anterior types, to the posterior types (Fig 4.14d and f). The anterior end of the keel is also rather wide, having six teeth rows compared to two to five in other species

(Fig. 4.14c). The anterior end appears to be swollen, maintaining the large number of teeth rows, where in the other species, the teeth rows reduce as the bone tapers anteriorly.

Trewavas shows similarity with T. brevis in regards to the swollen anterior of the bone, but I did not observe this character in T. brevis (Trewavas, 1931).

Figure 4.14: Characteristics of Apetra variabilis. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. Individual pictured is the Lectotype from Lake Nyasa South, BMNH 1930.1.31.4. 54 Description – Jaws isognathous (Fig. 4.14a); teeth on upper jaw in 4 rows in lectotype, 2-4 in paralectotypes; teeth in lower jaw in 4 rows in both lectotype and paralectotypes; 14 teeth in outer row of left lower jaw in lectotype, 12-15 in paralectotypes. Dorsal fin with 15 spines in lectotype, 14-16 in paralectotypes, 11 rays in lectotype, 10-12 in paralectotypes; pectoral fin with 15 rays in lectotype, 14-16 in paralectotypes; anal fin with 3 spines in the lectotype and paralectotypes, 9 rays in lectotype, 8-9 in paralectotypes. Lower pharyngeal bone triangular in outline with a notch in the posterior and curved inward laterally. Scales along side ctenoid with 32 in lateral-line series in lectotype, 32-33 in paralectotypes. First gill arch with 9 rakers on the ceratobranchial in lectotype, 7-10 in paralectotypes, 3 on the epibranchial in lectotype, 3-4 in paralectotypes, 1 between epibranchial and ceratobranchial (Table B.5).

Live coloration was not recorded. Preserved pattern consists of a broken lateral band, but may have a series of dots depending on the level of stress on the fish immediately before preservation, quality of preservation, or duration of preservation.

The pattern appears to contain a single break in the solid line, with a ventral shift at the break as the line continues toward the posterior of the fish.

Distribution – The type material comes from Lake Nyasa South and Monkey Bay

(Fig. 4.15). It is not known how far the range of this species extends at this time, as it did not appear in any other collection.

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi

Monkey Bay

South?

Figure 4.15: Localities of BMNH and of Apetra variabilis: Monkey Bay BMNH 1930.1.31.3; South BMNH 1930.1.31.4-13. The exact location(s) of “South” is/are unknown.

Discussion – In this case, the type material was comprised of two different species

(only one shown in map above). This has been a recognized problem for some time though the two species/two phenotypes issue had not been reconciled until now

(Trewavas 1931, Eccles and Trewavas 1989, Konings 2001). The resulting species may still exhibit both patterns (see Trewavas, 1931, Fig. 2), and more likely than not these patterns are only strictly adhered to in sympatry with similar species. I chose to designate the populations from the south as the Apetra variabilis even though they did not have

56 clear locality information. This species is illustrated in Trewavas, 1931, Fig. 2B, and shows the pattern of a broken lateral band. Eccles and Trewavas (1989) state that lectotypes were declared where needed; yet they did not declare one for A. variabilis. I can only assume that this is due to the possibility of separate species status between the two forms. I designated the lectotype (BMNH 1930.1.31.4). Other species that were part of the Apetra variabilis complex are discussed below.

Apetra trilineata (Trewavas) (Fig. 4.16)

Lethrinops trilineata Trewavas. 1931. A Revision of the Cichlid Fishes of the Genus

Lethrinops, Regan. Annual Magazine of Natural History, Ser. 10, 7: 133-152.

Tramitichromis trilineata (Trewavas) 1989. Eccles, David H., Ethelwynn Trewavas.

1989. Malawian Cichlid Fishes: The Classification of Some Haplochromine Genera.

Lake Fish Movies, Herten, Germany, pp 335.

Material Examined – HOLOTYPE BMNH.1930.1.31.76, Unknown locality

(Figs. 4.16, 4.17).

Diagnosis – Apetra trilineata is distinguished from A. lituris by the presence of a cusp on the anterior lower pharyngeal bone teeth compared to the cylindrical teeth of A. lituris. It is distinguished from A. intermedia by the lack of three dorsolateral spots and lack of molariform teeth on the posterior lower pharyngeal bone. A. trilineata is distinguished from A. variabilis by having two rows of teeth on the anterior lower pharyngeal bone compared to the swollen anterior lower pharyngeal bone with six rows

57 of teeth in A. variabilis. The pattern of this fish is very similar to A. lituris (Fig. 4.16a).

It is possible, although difficult, to distinguish A. trilineata and A. lituris based on external characteristics. It is difficult, however, to distinguish A. trilineata from other similar species (to be described) in this study based on external appearance. Although gill rakers on the outer ceratobranchial are at times helpful, they are not always reliable enough to be diagnostic. Apetra trilineata is described as having a keel inclined less than

45o to the plane of the toothed surface. The anterior teeth are said to have “a minute

anterior cusp and the ends turned backwards” (Eccles and Trewavas, 1989). The inner

posterior teeth are not to be enlarged. None of the over 700 (only some of which may be

this species) fish I have examined fit this description. Sometimes, some of the anterior

teeth are facing forwards, but this is not indicated in the description. Also, some fish

exhibit all the classic characters of A. trilineata, but there are enlarged posterior teeth, which would prohibit their inclusion in this species based on its formal description.

Figure 4.16: Characteristics of Apetra trilineata. Top to bottom: a) external appearance, b) gill rakers on outer ceratobranchial. The specimen pictured is the holotype, BMNH 1930.1.31.76.

Description – Jaws isognathous (Fig. 4.16a); teeth on upper and lower jaw in 4 rows; 13 teeth in outer row of left lower jaw. Dorsal fin with 16 spines, 11 rays; pectoral fin with 15 rays; anal fin with 3 spines, 10 rays. Lower pharyngeal bone missing. Scales along the side ctenoid with 33 in lateral-line series. First gill arch with 9 rakers on the

59 ceratobranchial, 4 on epibranchial, 1 between epibranchial and ceratobranchial (Table

B.6).

Distribution – The description of this species is based on one specimen from an unknown locality (Eccles and Trewavas, 1989). If we trace Christy’s path of collection in Lake Malaŵi in 1930, it would put him in the southern part of Lake Malaŵi, and I believe the tag on the holotype says “Monkey Bay,” but none of the authors have indicated this locality in their work (Fig. 4.17). I suspect the problem, however, is that there was a lot of collecting done at that time, and fish were often sorted after collection.

Sometimes long trawls were done, and the collectors were not sure from where the specimens came. A lot of the fish collected at this time were simply labeled “South.” In all, the type locality for A. trilineata remains a mystery. A major confounding problem with this species is that the lower pharyngeal bone, so crucial for proper identification, is missing from the holotype.

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi LOCALITY UNKNOWN

Monkey Bay

South?

Figure 4.17: Proposed location of Apetra trilineata, BMNH 1930.1.31.76.

Discussion – Without the lower pharyngeal bone, comparisons cannot be made.

From my research I have found that sometimes slight differences in the shape and dentition of this bone will separate species. Normally this would not present a difficult problem because many species during this time period were described using a group of fish (syntypes), without designating a holotype, for species descriptions. Unfortunately, there were no other fish of this species collected at the same time, or any other. Using strictly external morphology is not possible either, as this species looks like other Apetra, and SPCA plots cannot distinguish it (Fig. 4.3). This still would not present an impossible situation, except the type locality is in question. Trewavas described this

61 species in 1931 from one specimen collected by Christy in 1930 (Trewavas, 1931; Eccles and Trewavas, 1989). She indicated (1989) that the type locality of this fish is unknown.

A tag on the fish said Monkey Bay, but this may not be accurate (see above). This made it impossible to collect or observe topotypes to help delimit this species more clearly.

Because of the above stated reasons, I am recommending that A. trilineata be restricted to the holotype currently cataloged into the Natural History Museum (London)

(BMNH.1930.1.31.76). A comparable recommendation was made by Stauffer et al.

(1993) when they confronted a similar problem while working on the Copadichromis eucinostomus group.

Apetra linea, n. sp. (Fig. 4.18)

HOLOTYPE – BMNH 1930.1.31.17, adult female, 95.10 mm, Vua, Lake

Malaŵi, Malaŵi, Africa (Fig. 4.18).

PARATYPES - BMNH 1930.1.31.14-16, 18-20, 7 fish, Vua; BMNH

1930.1.31.1-2, 2 fish, Mwanga, Tanzania; PSU 4139, 3 fish, September 7, 1983, Fisheries

Research Station, Cape Maclear; PSU 4140, 4 fish, September 7, 1983, Fisheries

Research Station, Cape Maclear; PSU 4141, 1 fish, September 7, 1983, Fisheries

Research Station, Cape Maclear; PSU 4142, 5 fish, September7, 1983, Fisheries Research

Station, Cape Maclear; PSU 4145, 5 fish, April 4, 1984, Fisheries Research Station, Cape

Maclear; PSU 4150, 1 fish, April 4, 1984, Fisheries Research Station, Cape Maclear;

62 PSU 4161, 1 fish, September 7 1983, Fisheries Research Station, Cape Maclear (Fig.

4.20).

Diagnosis – Apetra linea is distinguished from A. lituris by a steeply inclined keel

(more than 45 degrees compared to less than 45 degrees), the presence of a swollen anterior lower pharyngeal bone with six rows of teeth compared to four rows of teeth, and a broken or spotted line on the dorsolateral portion of the fish compared to a dark line along the upper lateral line with other horizontal elements. It is distinguished from A. intermedia by the lack of three dorsolateral spots, lack of molariform teeth on the posterior lower pharyngeal bone, and a steeply inclinced keel at more than 45 degrees compared to a small decurved keel. Apetra linea is distinguished from A. variabilis by the broken or spotted line on the dorsolateral portion of the fish compared to the incomplete band with a break and ventral anterior shift in A. variabilis. It differs from A. trilineata by the swollen six rows of anterior teeth on the lower pharyngeal bone compared to the two rows in A. trilineata, and long cylindrical anterior lower pharyngeal bone teeth in A. linea and anterior teeth with a minute cusp in A. trilineata. Apetra linea is the second largest of all the Apetra species (up to 142 mm SL). The body pattern consists of a dark lateral band that runs from the nape to caudal base. It may be a broken non-overlapping line or an oblique series of spots (Fig. 4.18a, 4.19) (Eccles and

Trewavas, 1989). The lower pharyngeal bone is similar to A. variabilis (see description above). The angled keel with regard to the bone itself, is inclined at least 45 degrees, if not more (Figs. 4.18d). The anterior teeth are long and cylindrical, curving backwards

(Fig 4.18e). On some anterior teeth, a cusp was present, but this does not seem to be the

63 norm. The posterior teeth have a cusp, and the ends are pointed forward along the back row, previous rows pointed backward (Fig. 4.18f). The teeth seem to change from the anterior types, to the posterior types (Fig 4.18d and f). The anterior end of the keel is also rather wide, having six teeth rows compared to two to five in most other species

(Fig. 4.18c).

Description – Jaws isognathous (Fig. 4.18a); teeth on upper jaw in 4 rows in holotype, 2-4 rows in paratypes; teeth on lower jaw in 4 rows in the holotype and paratypes; 15 teeth in outer row of left lower jaw in holotype, 10-17 in paratypes. Dorsal fin with 16 spines in holotype, 14-16 in paratypes, 11rays in holotype, 10-12 in paratypes; pectoral fin with 15 rays in holotype, 15-16 in paratypes; anal fin with 3 spines

66 in the holotype and paratypes, 9 rays in holotype, 8-10 in paratypes. Lower pharyngeal bone triangular in shape with a notch in the posterior and curved inward laterally (Fig.

4.18c). Scales along side ctenoid with 33 in lateral-line series in lectotype, 31-34 in paratypes. First gill arch with 8 rakers on the ceratobranchial in holotype, 5-9 in paratypes; 4 on epibranchial in holotype, 2-4 in paratypes; 1 between epibranchial and ceratobranchial (Table B.7).

Live coloration not recorded. Preserved pattern consists of an oblique series of spots from the nape to the caudal base; it may posses a broken non-overlapping line instead (Eccles and Trewavas 1989).

Distribution – The type material comes from Vua and Mwanga in the northern end of the lake and Fisheries Research Station in the southern end (Fig. 4.20).

= PSU Collection Lake Malawi = BNMH Collection

Vua Tanzania Mwanga?

Zambia

Mozambique

Malawi

Fisheries Research Station

Figure 4.20: Locations of Apetra linea: Vua BMNH 1930.1.31.14-20; Mwanga BMNH 1930.1.31.1-2; Fisheries Research Station PSU 4139, 4140, 4141, 4142, 4145, 4150, 4161.

Discussion – Trewavas (1931) stated that she noticed that the A. variabilis contained two different patterns, the spotted form was more common in the north, while the broken line form came from the south (Trewavas, 1931; Eccles and Trewavas, 1989).

She was not able to find any other differences at those times. The types, which at the time of my research contained more than one species (A. variabilis and A. linea), came from the southern (Monkey Bay, Lake Nyasa [the former name of Lake Malaŵi] South) and northern (Vua and Mwanga) ends of the lake (Figs. 4.09, 4.18). Between the BMNH collections, there were four areas of the lake collected; two in the north, and two in the

68 south. Many times different patterns are reflective of different species, and historically patterns have been used many times to separate species (Eccles and Trewavas, 1989).

Naturally when I read about the difference in body pattern between northern and southern

T. variabilis populations I hypothesized that the two groups might be separate species. I was not very optimistic about it though as Trewavas stated (1931: 139) “I have been unable to correlate the difference of color-pattern [in T. variabilis] with any other differences.” The pattern (dorsolateral spots in the north, and oblique line in the south) observed by Trewavas in 1931, which was recapitulated in Eccles and Trewavas in 1989, seems to be explained by my study. What I discovered, was that the result of the plot of the first principal components of the meristic data and the sheared second principle components of the morphometric data show that the fish from the north (A. linea) and the fish from the south (A. variabilis) cluster distinctly (with a little overlap) suggesting separate species (Fig. 4.21). The minimum polygon clusters formed by the two species were significantly different along both the SPCA 2 (morphometric data) and PC 1

(meristic data) axes independent of each other. Variables that had the highest loadings on the sheared second principal components were cheek depth (-0.55767), distance between the posterior dorsal fin insertion and anterior anal fin insertion (0.28440), the dorsal fin base length (0.27787), and the head depth (-0.27448); while those with the highest loadings on the principal components of the meristic data were gill rakers on the first ceratobranchial (0.31701), dorsal spines (0.28297), and anal rays (0.18249). This, on top of the difference in pattern, led me to declare a lectotype for the A. variabilis from the southern collections and declare a new species (A. linea) for the northern collections. In addition, I was able to find A. linea in the southern end of the lake at Fisheries Research

69 Station, which was from PSU collections not available to Trewavas in 1931, or Eccles and Trewavas in 1989.

0.12

0.1

0.08

0.06

0.04

0.02 A. variabilis A. linea 0

-0.02 SPCA 2 (morphometric data) SPCA 2 (morphometric

-0.04

-0.06

-0.08

-0.1 -2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 PC 1 (meristic data)

What is particularly interesting is that the Monkey Bay specimen of A. variabilis did not cluster with the Lake Nyasa South fish (Fig. 4.21, bottom right corner of graph).

I would have liked to investigate this further, but the lower pharyngeal bone, so diagnostically important, was missing from this specimen. The fish was rather faded and

70 therefore the pattern could not be observed accurately. Also, it was the only fish from that locality so the question cannot be answered at this time. Another collection of this species from Monkey Bay should help resolve the issue and is suggested for future research. It is possible that this is a different species. For my analysis, I will rely on the

Lake Nyasa South A. variabilis for comparisons.

Etymology – The name linea from the Latin, meaning a line to note the dorsolateral band, which may be either spotted or broken.

Apetra simula, n. sp. (Fig. 4.22)

HOLOTYPE. – PSU 4187, adult male, 144.0 mm, First Beach North of Otter

Point, Lake Malawi, Malawi, Africa, 9.1 m, July 6, 1991 (Fig. 4.22).

PARATYPES PSU 4097, 6 fish, July 6, 1991, Otter Point; PSU 4112, 5 fish,

September 6, 1983, Golden Sands Swamp; PSU 4113, 5 fish, September 6, 1983, Golden

Sands Swamp; PSU 4118, 12 fish, April, 6, 1983, Otter Point; PSU 4121, 8 fish, April 7,

1983, Otter Point; PSU 4122, 1 fish, April 6, 1983, Otter Point (Fig. 4.23).

Diagnosis – Apetra simula is distinguished from all other species by the presence of 25 % of the outer anterior lower pharyngeal bone teeth turned toward the midline. In addition, it is distinguished from A. lituris and A. trilineata by the presence of a swollen anterior lower pharyngeal bone with six rows of teeth compared to four in A. lituris and

71 two in A. trilineata ; it also has an extremely large keel inclined downward at more than

45 degrees compared to a smaller keel inclined downward at less than 45 degrees in A. lituris and A. trilineata. It is distinguished from A. intermedia by the lack of molariform posterior lower pharyngeal bone teeth, its body pattern lacks the three dorsolateral spots

(small dark horizontal sections instead), and has an extremely large keel on the lower pharyngeal bone inclined at more than 45 degrees compared to a small decurved keel. It is distinguished from A. variabilis and A. linea by the extremely large keel and the body pattern containing small dark sections compared to an incomplete band with a break and ventral anterior shift in A. variabilis and a broken or spotted line in A. linea. Apetra simula is the largest of all the Apetra species (up to 145 mm SL). The body pattern is slightly different from that described for A. variabilis and A. linea. It consists of an incomplete oblique band from the nape, intersecting the lateral line below the posterior part of the spiny dorsal (Fig. 4.23a). These fish do not contain the single break in the solid line as seen in A. variabilis (Fig. 4.18a; see Fig. 2B of Trewavas, 1931), nor do they have a series of spots like in some A. linea (Fig. 4.18a). This species contains some dark horizontal elements dorsolaterally intermediate between a spotted or solid line appearance (see Fig. 4.21). The patterns seen in A. variabilis and A. linea can also be observed (Figs. 4.09a, 4.18a, 4.19). The lower pharyngeal bone has a very large keel when viewed laterally (Fig. 4.22d). The anterior teeth are cylindrical with the ends curving toward the posterior. Also, at least 25 percent of the outer two or more rows of anterior teeth are turned toward the midline (Fig. 4.22c). This is unique to this species.

Posterior teeth have a cusp with the ends pointing backward except for the last row (Fig.

4.22f).

73 Description – Jaws isognathous (Fig. 4.22a); teeth on upper jaw in 4 rows in holotype, 2-4 in paratypes, teeth in lower jaw in 5 rows in the holotype, 4-5 in paratypes,

10 teeth in outer row of left lower jaw in holotype, 10-17 in paratypes. Dorsal fin with 16 spines in holotype, 13-17 in paratypes, 11 rays in holotype, 10-13 in paratypes; pectoral fin with 16 rays in holotype, 14-17 in paratypes; anal fin with 3 spines in the holotype and paratypes, 9 rays in holotype, 8-10 in paratypes. Lower pharyngeal bone triangular in outline with a deep notch posteriorly with the lateral sides curving inward (Fig. 4.22c).

Scales along side ctenoid with 34 in lateral-line series in holotype, 30-34 in paratypes.

First gill arch with 8 rakers on the ceratobranchial in holotype, 6-13 in paratypes, 4 on the epibranchial in the holotype, 3-5 in paratypes, 1 between the epibranchial and ceratobranchial (Table B.8).

Males have a yellowish-blue body with faint blue bars, the dorsal surface being darker than the sides. Dorsal fin blue-black with golden-orange spots on membrane, white outer margin with orange lappets. Caudal fin blue with brown vermiculations, outer edge tinged with orange. Anal fin gray with 15 yellow ocelli. Pelvic fin gray with yellow leading edge. Pectoral fin gray. Head iridescent blue-green, gular region dark.

Females body whitish with flecks of yellow, dorsal surface dark olive green.

Dorsal fin whitish with orange-brown spots on membrane, orange lappets. Caudal fin gray with orange-brown spots. Anal fin clear with 9 yellow ocelli, yellow leading edge.

Pelvic fin yellow proximally, clear distally, white leading edge. Pectoral fin clear. Head yellowish, black opercular spot, gular region yellow.

74 Preserved pattern consists of dark horizontal elements of a broken lateral band, but may have a series of dots.

Distribution – These fish were found at Otter Point and Golden Sands Swamp

(Fig. 4.23). It is unknown how far their range extends, as they did not appear in any other

PSU or BMNH collection. They may be endemic to the listed localities.

= PSU Collection Lake Malawi = BNMH Collection

Tanzania

Zambia

Mozambique

Malawi

Golden Sands Swamp Otter Point

Figure 4.23: Localities of Apetra simula: Otter Point PSU 4097, 4187; Golden Sands Swamp PSU 4112, 4113, 4118, 4121, 4122.

75 Discussion – An unexpected discovery I made while working with the

Tramitichromis [Apetra] variabilis complex was that the type material consisted of two species: A. variabilis and A. linea (see discussions above). When I plotted the first principal components of the meristic data and the sheared second principle components of the morphometric data, the PSU collections that were identified as the A. variabilis complex (now A. simula) from the southern portion of the lake, and the BMNH southern group (now A. variabilis), produced two separate clusters, distinct from each other and the bipolar group (A. linea) (Fig. 4.24). The minimum polygon clusters formed by the three species were significantly different (p<0.05) along the SPCA 2 (morphometric data) axis. Variables that had the highest loadings on the sheared second principal components were preorbital depth (-0.38626), snout length (-0.37056), and distance between the posterior dorsal insertion to the pelvic fin origin (0.36640); while those with the highest loadings on the principal components of the meristic data were dorsal rays (0.34106), pectoral fin rays (0.30191), dorsal spines (-0.27725), and anal rays (0.27097). The two southern groups differ predominantly along the morphometric (shape; SHRD PC2) axis, so I wondered if these differences are due to comparing the BNMH specimens, which were old, and the PSU material, which was not. But if this were true, I would expect to see this pattern every time I compared old and new specimens, which I have not. Unless the types have been badly damaged in some way, the type material and the fresh material cluster together when you have the same species. This evidence, along with the pattern differences and lower pharyngeal bone differences reflects separate species status.

0.15

0.1

0.05

A. variabilis 0 A. linea A. simula SPCA 2 (morphometric data)

-0.05

-0.1

-0.15 -4 -3 -2 -1 0 1 2 3 PC 1 (meristic data)

Figure 4.24: Plot of the second sheared principle components (morphometric data) and the first factor scores (meristic data) of Apetra simula (N = 38): Otter Point PSU 4097, 4187; Golden Sands Swamp 4112, 4113, 4118, 4121, 4122; Apetra linea (N = 10): Vua BMNH 1930.1.31.14-20; Mwanga BMNH 1930.1.31.1-2; Fisheries Research Station PSU 4139, 4140, 4141, 4142, 4145, 4150, 4161; and Apetra variabilis (N = 12): Lake Nyasa South BMNH 1930.1.31.4-13; Monkey Bay BMNH 1930.1.31.3.

Apetra variabilis, A. linea, and A. simula are phenetically similar, yet distinct species. All three species are found in the southern portion of the lake, and many of the localities are very close together. Apetra linea from Fisheries Research Station, and A. simula from Otter Point and Golden Sands Swamp are practically sympatric. The maps

77 (Figs. 4.10, 4.20, 4.23) make the localities appear farther apart than they actually are.

These three sites are rather close geographically and most likely do not have very different environmental conditions. Also, the A. variabilis from the Lake Nyasa South group may have come from Cape Maclear, the region where the other three sites are located, or it may have came from the Southeast arm of the lake; I have no way of knowing for sure. They were collected around the same time as the Monkey Bay fish, so it is more likely than not that they came from the southeast arm. They just as well might have come from the southwest arm of the lake, and be sympatric with A. simula. My next thought was that the type specimens and the A. linea from Fisheries Research Station fish were caught at different depths, which produced different body forms. A comparison using only the A. linea Fisheries Research Station fish, which were from two collections, one caught at 12m (PSU 4139, 4140, 4141, 4142), and one caught at 36-54m (PSU 4145,

4150), did not reveal any separate clustering (Fig 4.25). The minimum polygon clusters observed by the two depth groups were not significantly different (p<0.05). This means that the fish from different depths do not have different body forms, at least for A. linea, and differences observed in Fig. 4.24 are not due to depth as all were captured around 15-

50 m, each species having some deep and shallow water collections.

0.08

0.06

0.04

0.02

0 Collection at 12 meters Collection at 36-54 meters -0.02

-0.04 SPCA 2 (morphometric data)

-0.06

-0.08

-0.1 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 PC 1 (meristic data)

In regards to the pattern, it is most likely the closer geographically these species are to one another the more the species specific patterns come into play.

Etymology – The name simula from the Latin, meaning a likeness or imitation to note its apparent similarities with A. linea and A. variabilis.

Apetra perjur, n. sp. (Fig. 4.26)

79 HOLOTYPE. – PSU 4162, adult male, 110.0 mm, Songwe Hill, Lake Malaŵi,

Malaŵi, Africa, 6-8 m, January 21, 1997 (Fig. 4.26).

PARATYPES – PSU 4087, 24 fish, January 21, 1997, Songwe Hill; PSU 4096,

19 fish, March 27, 1996 Songwe Hill; BMNH 1930.1.31.45, 1 fish, Bar to Fort Maguire

(Fig. 4.27).

Diagnosis – Apetra perjur is distinguished from A. lituris by the anterior lower pharyngeal bone teeth which are curved backward at around 90 degrees compared to a range of straight up at 85 degrees or curved back no more than 45 degrees. It is distinguished from A. intermedia by the dark line and other horizontal elements compared to three dorsolateral spots, a keel inclined downward at no more than 45 degrees compared to a small decurved keel, and the absence of molariform teeth on the posterior lower pharyngeal bone. It is distinguished from A. trilineata by the cylindrical anterior lower pharyngeal bone teeth compared to the anterior teeth with a minute cusp.

It is distinguished from A. variabilis, A. linea, and A. simula by the presence of four rows of anterior lower pharyngeal bone teeth compared to the six rows and swollen appearance of the anterior lower pharyngeal bone in the other three. This species has characteristics rather similar to Apetra lituris (see diagnosis above) with a major exception (Figs. 4.11,

4.26). This species has a lower pharyngeal bone and lower pharyngeal teeth in the same configuration as A. lituris (see above), except that it is distinguished by the anterior teeth being mostly turned backward at more than 45o, and most approach 90o (Fig. 4.26f).

81 Description – Jaws isognathous (Fig. 4.26a); teeth on upper jaw in 3 rows in holotype, 2-4 in paratypes; teeth in lower jaw in 4 rows in holotype, 3-5 in paratypes; 14 teeth in outer row of left lower jaw in holotype, 9-15 in paratypes. Dorsal fin with 15 spines in holotype, 14-16 in paratypes, 12 rays in holotype, 10-13 in paratypes; pectoral fin with 16 rays in holotype, 14-17 in paratypes; anal fin with 3 spines in both the holotype and paratypes, 9 rays in holotype, 8-10 in paratypes. Lower pharyngeal bone triangular in outline with a broad notch posteriorly (Fig. 4.26c). Scales along the side ctenoid with 34 in lateral-line series in holotype, 32-35 in paratypes. First gill arch with 7 rakers on the ceratobranchial in holotype, 6-12 in paratypes, 3 on epibranchial in holotype, 3-4 in paratypes, 1 between the epibranchial and ceratobranchial (Table B.9).

Live coloration for the males consists of lateral scales being yellow outlined in blue. There is a blue/black blotch about 21 lines under the dorsal spines. There is a faint black lateral band from the dorsal rays to the caudal fin. The ventral portion of the fish is white. The head is dark gray between the eyes. The ventral portion of the head to the eyes has blue, green, and yellow highlights. The preorbital is blue. The gular is yellow with black on the posterior half. The dorsal fin has blue/green membranes with yellow spots/vermiculations. There is a black submarginal band, a white marginal band, and yellow lappets. The caudal fin is blue with yellow vermiculations. The anal fin has a proximal portion (4/5) that is blue and a distal portion (1/5) that is yellow. There are 10-

15 ocelli throughout the fin. The pelvic fins have a white leading edge with the tip of the first ray black. The fin also has vermiculations with a faint yellow cast. The pectoral fin was clear.

82 Females laterally have the top 1/3 of their body containing blue/green highlights with the ventral 2/3 portion being white. The ventral portion of the fish is white. The head is gray dorsally and white below the eye to the ventral portion of the fish. The gular is white. The dorsal fin is clear with orange lappets. The caudal fin is clear with yellow just on the dorsal and ventral two rays. The anal fin is clear with yellow distally. Pelvic and pectoral fins are clear.

Preserved pattern consists of a dark line along the upper lateral line, and includes darker elements along the “bars”.

Distribution –There are two localities know to contain this species. The first is

Songwe Hill (PSU 4087, 4096, 4162) and the seconds is the bar to Fort Maguire (BMNH

1930.1.31.45) (Fig. 4.27).

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi Fort Maguire

Songwe Hill

Figure 4.27: Localities of Apetra perjur: Songwe Hill PSU 4087, 4096, 4162 and the bar to Fort Maguire BMNH 1930.1.31.45.

Discussion – Konings (1995; 2001), Turner (1996), and Stauffer (per. comm.) have stated that Apetra lituris [formerly Tramitichromis lituris] might be a complex of many species. What I have discovered seems to lend credence to their thoughts. When I examined the lower pharyngeal bone of this species, I found two basic types. The first has anterior teeth that are cylindrical and most are turned backwards slightly at an angle of up to but not more than 45o (A. lituris). The second type, had cylindrical anterior

teeth, but most were turned backward at 45 to 90o (A. perjur). Also, the differences in

lower pharyngeal bone tooth structure corresponded to the locality at which they were

84 collected – less than 45o in the north, 45-90o in the south (Fig. 4.28). This led to my discovery that the type material was actually comprised of two species; A. lituris which

occurs in the north (Fig. 2.14) and A. perjur which occurs in the south (Fig. 4.27). Then,

when I plotted the first principal components of the meristic data and the sheared second

principle components of the morphometric data, I find that A. lituris clusters separately

from A. perjur (Fig. 4.29). In addition, the A.perjur from the British Museum collection clusters with the other A. perjur specimens from the PSU collections. There is one A. perjur individual that has the lower pharyngeal bone characteristics of the species, but for whatever reason does not cluster with the group. The plot of the minimum polygon clusters formed by the two species found significant differences (p<0.05) along both the

SPCA 2 (morphometric data) and PC 1 (meristic data) axis independent of each other.

Variables that had the highest loadings on the sheared second principal components were preorbital depth (-0.57363), cheek depth (-0.44853), and distance between the insertion of the posterior of the dorsal fin and the pelvic fin origin (0.28239); while those with the highest loadings on the principal components of the meristic data were dorsal rays (-

0.33559), dorsal spines (0.31102), teeth rows on the upper jaw (0.21411), and teeth rows on the lower jaw (0.21253).

0.15

0.1

0.05

A. lituris LECTOTYPE A. lituris PARALECTOTYPES 0 A. perjur HOLOTYPE A. perjur PARATYPES A. perjur PARATYPE BMNH

SPCA 2 (morphometric data) -0.05

-0.1

-0.15 -2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 3 PC 1 (meristic data)

Etymology – The name perjur from the Latin, meaning lying or false to note its apparent similarities with A. lituris.

Apetra meniscosteum, n. sp. (Fig. 4.30)

HOLOTYPE. – PSU 4163, adult male, 87.6 mm, Chirombo Bay, Kanjedza Island,

Lake Malaŵi, Malaŵi, Africa, 3 m, January 15, 1989 (Fig. 4.30).

87 PARATYPES. – PSU 4130, 9 fish, January 12, 1989, Chirombo Bay, Kanjedza

Island; PSU 4134, 10 fish, January 15, 1989, Chirombo Bay, Kanjedza Island (Fig. 4.31).

Diagnosis – Apetra meniscosteum is distinguished from A. lituris and A. perjur by the anterior lower pharyngeal bone teeth having a cusp compared to cylindrical teeth, some of the anterior lower pharyngeal bone teeth turning forward while other are back compared to a range of all straight up to 45 degrees back in A. lituris and curved backward at 90 degrees in A. perjur, and a cresecent shaped keel compared to a keel inclined downwards at 45 degrees in A. lituris and A. perjur, and a small breeding adult size (up to 87.08 mm compared to up to 125.21 mm in A. lituris and 123.58 in A. perjur.

It is distinguished from A. intermedia by a dark line with other horizontal elements compared to three dorsolateral spots, a cresent shaped keel compared to a decurved keel, and two rows or less compared to all of the posterior lower pharyngeal bone teeth being molariform. It is distinguished from A. variabilis, A. linea, and A. simula by the having two rows of anterior lower pharyngeal bone teeth compared to the six rows and swollen appearance, the crescent shaped keel compared to a steeply inclined one at more than 45 degrees, and a cusp on the anterior lower pharyngeal bone teeth compared to cylindrical teeth. It is distinguished from A. trilineata by the having some of the anterior lower pharyngeal bone teeth pointed forward with others backward compared to all backward.

Apetra meniscosteum is phenotypically similar to A. lituris, A. perjur, A. variabilis, and

A. simula, (Figs. 4.09a, 4.11a, 4.22a, 4.26a, 4.30a). Of the two new species found at

Kanjedza Island, it had the largest keel and anterior-ventral projection of the blade of the lower pharyngeal bone (Fig. 4.30d). There seems to be some variability in the blade

88 shape, as some look less curved. The anterior teeth have a cusp, and some ends point forward, others back (Fig. 4.30e). The posterior teeth are enlarged a few rows from the end, but this trait varies among the fish in the group (Fig. 4.30c, f).

Figure 4.30: Characteristics of Apetra meniscosteum. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, f) posterior pharyngeal teeth. The specimen pictured is the holotype from Kanjedza Island PSU 4163. 90 Description – Jaws isognathous (Fig. 4.30a); teeth on upper jaw in 3 rows in holotype, 3-4 in paratypes; teeth on lower jaw in 4 rows in holotype, 4-5 in paratypes; 9 teeth in outer row of left lower jaw in holotype, 7-12 in paratypes. Dorsal fin with 16 spines in holotype, 15-17 in paratypes, 11 rays in holotype, 10-12 rays in paratypes; pectoral fin with 15 rays in holotype, 14-15 in paratypes; anal fin with 3 spines in both the holotype and paratypes, 8 rays in holotype, 8-10 in paratypes. Lower pharyngeal bone triangular in outline with a slight notch posteriorly (Fig. 4.30c). Scales along side ctenoid with 34 in lateral-line series in holotype, 33-35 in paratypes. First gill arch with

11 rakers on the ceratobranchial in holotype, 10-12 in paratypes, 4 on epibranchial in holotype, 3-5 in paratypes, 1 between the epibranchial and ceratobranchial (Table B.10).

Live coloration of males consists of the lateral side with a red blotch behind the opercle and under the pectoral fin. The side was yellow with scales outlined in blue and turns darker dorsally. The head is blue with green flecks. The cheek is gray. The gular is a blue-gray color. There is also a dark blue opercular spot. The dorsal fin has a white marginal band with white lappets. The submarginal band is a diffuse gray/black. The proximal 2/3 is blue with yellow/orange spots. The caudal fin has blue membranes with yellow/orange vermiculations with the posterior margin clear. The anal fin is a black/gray with a red marginal band. The pelvic fins are gray/black. The pectoral fins are clear.

The females are silvery. They are darker dorsally and white ventrally.

91 Preserved pattern consists of a dark dorsolateral line or lines which may or may not consist of some other vertical elements.

Distribution – These fish are endemic to Kanjedza Island in the southeastern arm of Lake Malaŵi in Africa (Fig. 4.31).

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi

Kanjedza Island

Figure 4.31: Location of the collection of Apetra meniscosteum: Kanjedza Island PSU 4130, 4134, and 4163.

92 Discussion – Both Stauffer and Konings (per. comm.) have indicated that

Kanjedza Island has been continuously overfished during the past few years.

Etymology – The name meniscosteum from the Greek meaning “crescent- shaped” (menisc) and “bone” (osteum) to reflect the crescent-shaped keel on the lower pharyngeal bone.

Apetra cryptopharynx, n. sp. (Fig. 4.32)

HOLOTYPE. – PSU 4186, adult male, 82.7 mm, Chirombo Bay, Kanjedza Island,

Lake Malaŵi, Malaŵi, Africa, 4.3-5.5 m, February, 1987 (Fig. 4.32).

PARATYPES. – PSU 4131, 46 fish, Feb. 1987; 4136, 13 fish, Feb. 1987,

Chirombo Bay, Kanjedza Island; PSU 4093, 6 fish, April 9, 1983, Golden Sands Swamp;

PSU 4082, 25 fish, January 5, 1991, Songwe Hill; PSU 4085, 23 fish, January 20, 1994,

Songwe Hill; PSU 4133, 21 fish, February 1, 1991, Songwe Hill; PSU 4115, 15 fish,

February 15, 1987, Island off Nkhudzi Bay; PSU 4083, 29 fish, July 16, 1991, First sand beach north of Otter Point; PSU 4111, 6 fish, July 16, 1991, First sand beach north of

Otter Point; PSU 4120, 6 fish, April 7, 1983, Otter Point (Fig. 4.34).

Diagnosis – Apetra cryptopharynx is distinguished from A. lituris, A. perjur, A. variabilis, A. linea, and A. simula by the anterior lower pharyngeal bone teeth having a cusp with the ends pointed forward compared to cylindrical with the ends pointing

93 backward. Its keel is not inclined as steeply (less than 45 degrees) as in A. variabilis, A. linea, and A. simula (more than 45 degrees). It is distinguished from A. intermedia by the keel, which is inclined and long with the top and bottom portions of the keel being parallel to each other compared to a small decurved keel, the pattern consists of a dark line along the upper lateral line with other horizontal elements compared to three dorsolateral spots. It is distinguished from A. trilineata by a distinct cusp on the anterior lower pharyngeal bone teeth compared to a minute cusp, and the variable enlargement of the posterior lower pharyngeal bone teeth which A. trilineata lacks. It is distinguished from A. mensicosteum by the long parallel top and bottom portion of the keel compared to a variable shorter crescent shaped keel, most of the anterior lower pharyngeal bone teeth turned forward compared to some forward with others backward on the same bone.

This species was very difficult to detect, and has probably been a source of confusion for much time. The outward appearance does not offer any assistance in its identification as it appears as A. meniscosteum with darker elements along the bars (see discussion above)

(Fig. 4.32a). The lateral view of the lower pharyngeal bone shows a slight downward projection to the keel (Fig. 4.32d). The angle of this projection and the distance it projects can vary with the individual (Fig 4.33). The anterior teeth have a cusp, with the ends mostly forward (Fig. 4.32e). The posterior teeth may be enlarged, and the number of enlarged rows varies with the individual (4.32c, f).

Figure 4.33: The keels of six individuals showing the variability of the shape and length of the keel of Apetra cryptopharynx. Clockwise starting with the top left Kanjedza Island individuals pictured are from collections: a) PSU 4186 holotype, b) PSU 4136 #4, c) PSU 4105 #3, d) PSU 4105 #8, e) PSU 4105 #10, f) PSU 4105 #1.

Description – Jaws isognathous (Fig. 4.32a); teeth on upper jaw in 2 rows in holotype, 2-5 in paratypes; teeth on lower jaw in 4 rows in holotype and 4-5 in paratypes;

15 teeth in outer row of left lower jaw in holotype, 8-19 in paratypes. Dorsal fin with 15 spines in holotype, 13-17 in paratypes, 12 rays in holotype, 9-12 in paratypes; pectoral fin with 15 rays in holotype, 13-16 in paratypes, anal fin with 3 spines in holotype, 2-4 in paratypes. Lower pharyngeal bone triangular in outline with a broad notch posteriorly

(Fig. 4.32c). Scales along side ctenoid with 33 in lateral-line series in holotype, 31-35 in

96 paratypes. First gill arch with 11 rakers on the ceratobranchial in the holotype, 8-15 in paratypes, 5 on the epibranchial in holotype, 3-7 in paratypes, 1 between the epibranchial and ceratobranchial (Tables B.11, B.12).

Live coloration was not recorded except for the male having either an orange gular or a yellow gular.

Preserved pattern consists of some degree of a fragmented dark dorsolateral band which may include some darker vertical elements.

Distribution – The type specimens in this study came from Kanjedza Island in the southeast arm of Lake Malaŵi (Fig. 4.34). They were also found at Otter Point, Nkhudzi

Bay, Songwe Hill, and Golden Sands Swamp of Cape Maclear.

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi

Kanjedza Island

Golden Sands Swamp Nkhudzi Bay Otter Point Songwe Hill

Discussion – I compared Apetra cryptopharynx to Apetra meniscosteum and plotted the second principal components of the meristic data against the sheared third principle components of the morphometric data (Fig. 4.35). Outside of a few individuals, the clusters did not overlap supporting separate species status. The outliers were reexamined to ensure proper identification. They had the lower pharyngeal bone characteristics of A. cryptopharynx, but were found, for whatever reason, in the A. mensicsoteum cluster. The plot of the minimum polygon clusters formed by the two species was significantly different (p<0.05) along both the SPCA 3 (morphometric data)

98 and the PC 2 (meristic data) axes independent of each other. The variables that had the highest loadings on the sheared third principle components were preorbital depth (-

0.45690), vertical eye diameter (0.43835), and horizontal eye diameter (0.39558); while those with the highest loadings on the second principle components of the meristic data were lateral line scales (0.42760), lower gill rakers (-0.32225), and cheek scales (-

0.31661).

0.15

0.1

0.05

A. meniscosteum 0 A. cryptopharynx SPCA 3 (morphometric data) 3 (morphometric SPCA -0.05

-0.1

-0.15 -3 -2 -1 0 1 2 3 4 PC 2 (meristic data)

99 Etymology – The name cryptopharynx, from the Greek, meaning hidden or concealed (crypto) and throat (pharynx). This is in reference to it being a cryptic species, where only the lower pharyngeal bone can distinguish it, but differences are slight, therefore this species may be concealed to the untrained eye.

Apetra retrodens, n. sp. (Fig. 4.36)

HOLOTYPE. – PSU 4164, adult male, 102.1 mm, Chembe Village, Cape

Maclear, Lake Malaŵi, Malaŵi, Africa, 2.1-3 m, March 17, 1985 (Fig. 4.36).

PARATYPES. – PSU 4119, 14 fish, March 17, 1985, Chembe Village, Cape

Maclear; PSU 4157, 7 fish, April 15, 1984, Chembe Village; PSU 4158, 18 fish, April 18,

1984, Chembe Village; PSU 4159, 1 fish, April 18, 1984, Chembe Village; PSU 4160, 9 fish, April 18, 1984, Chembe Village; PSU 4156, 19 fish, April 18, 1984, Chembe

Village; PSU 4084, 8 fish, March 21, 1995, Chembe Village; PSU 4095, 12 fish, March

21, 1995, Chembe Village; PSU 4151, 3 fish, April 9, 1984, Golden Sands Swamp; PSU

4154, 8 fish, April 9, 1984, Golden Sands Swamp; PSU 4146, 5 fish, April 8, 1984,

Fisheries Research Station; PSU 4149, 5 fish, April 8, 1984, Fisheries Research Station;

PSU 4153, 2 fish, April 9, 1984, Fisheries Research Station; PSU 4152, 12 fish, April 9,

1984, Fisheries Research Station (Fig. 4.37).

Diagnosis – Apetra retrodens is distinguished from all other species by the presence of two types of anterior lower pharyngeal bone teeth, some with a cusp and some cylindrical on the same bone (A. lituris, A. perjur, A. variabilis, A. linea, and A.

100 simula have only cylindrical; A. intermedia, A. meniscosteum, and A. cryptopharynx have a cusp; A. trilineata has a minute cusp). It can also be distinguished from all other species except A. trilineata (for which the bone was missing) based on the keel on the lower pharyngeal bone which is short and blunt, and dorsoventrally tall compared to inclined downwards less than 45 degrees in A. lituris and A. perjur, decurved in A. intermedia, inclined downward at more than 45 degrees in A. variabilis, A. linea, and A. simula, crescent shaped in A. mensicosteum, long and thin with the top and bottom portions parallel in A. cryptopharynx. It is distinguished from A. intermedia by the presence of a dark line along the upper lateral line with other dark horizontal elements compared to three dorsolateral spots. Its external appearance is once again of little use in comparison with many species, as this species contain dark elements along the bars (Fig.

4.36a). The lateral view of the lower pharyngeal bone shows a unique shape where the blade itself is rather short and blunt, and dorsoventrally tall (Fig. 4.36d). The anterior lower pharyngeal bone teeth of this species are highly variable. They may have a cusp

(Fig. 4.36e), or may be cylindrical (Fig. 4.36f) – even on the same bone. Also, the number of teeth turned backwards distinguishes the species. This, however, was variable.

There may be as little as one-quarter of the anterior teeth or almost all the anterior teeth turned backwards. The posterior teeth were, in general, not enlarged, although one or two rows of enlargement was not too rare (Fig. 4.36g).

101

Figure 4.36: Characteristics of Apetra retrodens. Clockwise from the top: a) external appearance, b) gill rakers on outer ceratobranchial, c) dorsal view of lower pharyngeal bone, d) lateral view of lower pharyngeal bone, e) anterior pharyngeal teeth, left side f) anterior pharyngeal teeth, right side, g) posterior pharyngeal teeth. The specimen pictured is the holotype from Chembe Village PSU 4164. 102 Description – Jaws isognathous (Fig. 4.36a); teeth on upper jaw in 3 rows in holotype, 2-4 rows in paratypes, teeth on lower jaw in 4 rows in holotype, 4-5 in paratypes; 12 teeth in outer row of left lower jaw in holotype, 9-19 in paratypes. Dorsal fin with 15 spines in holotype, 14-17 in paratypes, 11 rays in holotype, 9-13 in paratypes; pectoral fin with 16 rays in holotype, 14-17 in paratypes; anal fin with 3 spines in holotype and paratypes, 10 rays in holotype, 8-10 in paratypes. Lower pharyngeal bone triangular in outline with a broad notch posteriorly. Scales along side ctenoid with 33 in lateral-line series in holotype, 30-36 in paratypes. First gill arch with 10 rakers on the ceratobranchial in holotype, 6-15 in paratypes, 2 on epibranchial in holotype, 2-6 in paratypes, 1 between the epibranchial and ceratobranchial (Table B.13).

Live coloration of the types is as follows. Males have their lateral side with a black diffuse stripe from the dorsal to the lateral line. There is another stripe between the first stripe and the dorsal fin base. They have blue highlights on yellow ground color and seven faint diffuse vertical bands. The nape has an orange blotch. The head has a cheek and preorbital that is blue. The interorbital area is dark gray. The gular ranges in color from yellow to orange with black micromelanophores. The pelvic fins have black micromelanophores over yellow with the leading edge white. The pectoral fins are clear.

The anal fin proximally has a greenish/blue sheen with bright yellow ocelli. Distally it is black with the anterior rays having orange tips. The dorsal fin has black membranes with orange spots. Posteriorly the membranes change from black to blue. There is a white marginal band with orange lappets. The caudal fin has a blue membrane with yellow vermiculations.

103 Females have a lateral side that is dark dorsally and fades to white ventrally. .

They have the same two stripes as the males along with the seven vertical bands. The head has a white cheek and preorbital, dark gray interorbital area, and a yellow-white gular. The anal fin is clear with anterior membranes of spines yellow. The distal third of the ray membranes is yellow-orange. The dorsal fin is clear with yellow orange spots.

There are orange tips of the spines. The pelvic fin is yellow. The pectoral fin is clear.

The preserved pattern is not too distinguishing. Some, all, or none of the various two horizontal stripes and seven vertical bars may be seen.

Distribution – The type material comes from Chembe Village of Cape Maclear,

Lake Malaŵi, Africa (Fig. 4.37). This species was also found at Golden Sands Swamp of

Cape Maclear, and Fisheries Research Station of Cape Maclear.

104

= PSU Collection Lake Malawi = BMNH Collection

Tanzania

Zambia

Mozambique

Malawi

Chembe Village

Fisheries Research Station

Golden Sands Swamp

Figure 4.37: Localities of Apetra retrodens: PSU 4084, 4095, 4119, 4146, 4149, 4151, 4152, 4153, 4154, 4156, 4157, 4158, 4159, 4160, 4164.

Discussion – When the first principal components of the meristic data were plotted against the sheared second principle components of the morphometric data for A. retrodens and A. meniscosteum, they appear as two separate clusters (Fig. 4.38). There are two A. retrodens individuals that appear in the A. meniscosteum cluster. Their lower pharyngeal bone characteristics were reexamined. It appears that they were correctly identified, but for whatever reason they cluster away from the rest. The minimum polygon clusters formed by these two species was significantly different (p<0.05) along both the SPCA 2 (morphometric data) and PC 1 (meristic data) axes independent of each

105 other. This indicates that as well as the differences that I noted in the lower pharyngeal bones, there are shape differences between the two species also. The variables that had the highest loadings on the sheared second principle components were snout length (-

0.40164), caudal peduncle length (0.38931), and horizontal eye diameter (-0.35344); while those with the highest loadings on the first principal components of the meristic data were dorsal spines (0.25555), lower gill rakers (0.24390), and teeth rows in upper jaw (0.23540).

When the first principal components of the meristic data were plotted against the sheared second principle components of the morphometric data for A. retrodens and A. cryptopharynx, the two clusters meet and overlap slightly, with the same two A. retrodens outliers from the last graph in the cluster of A. cryptopharynx (Fig. 4.39). The minimum polygon clusters formed by the two species were significantly different (p<0.05) along both the SPCA 2 (morphometric data) and PC 1 (meristic data) axes independent of each other. The variables that had the highest loadings on the sheared second principal components were snout length (-0.45005), caudal peduncle length (0.27135), and lower jaw length (-0.25721); while those with the highest loadings on the principal components of the meristic data were pectoral rays (-0.28998), dorsal rays (-0.28517), and lower gill rakers (0.27925).

The type material of all three species was compared together when the second principal components of the meristic data were plotted against the sheared second principle components of the morphometric data (Fig. 4.40). Three separate clusters can

106 be observed, with two A. cryptopharynx individuals spilling over into the A. meniscosteum cluster. The minimum polygon clusters formed by the three species was significantly different (p<0.05) along both the SPCA 2 (morphometric data) and the PC 2

(meristic data) axes independent of each other. The variables that had the highest loadings on the sheared second principal components were caudal peduncle length

(0.41329), vertical eye diameter (-.38017), and horizontal eye diameter (-0.37851); while those with the highest loadings on the second principal components of the meristic data were lateral line scales (0.45410), dorsal spines (0.32697), and teeth rows in the upper jaw (0.28149). In summary, a comparison to the lateral views of the lower pharyngeal bones of A. meniscosteum, A. cryptopharynx, and A. retrodens shows their unique species-specific shape differences (Fig. 4.41) which are corroborated by lower pharyngeal bone anterior tooth differences (described above) and body shape differences illustrated in the SPCA plots shown above.

107

0.25

0.2

0.15

0.1

0.05 A. meniscosteum A. retrodens 0 SPCA 2 (morphometric data) 2 (morphometric SPCA -0.05

-0.1

-0.15

-0.2 -3 -2 -1 0 1 2 3 4 PC 1 (meristic data)

108

0.2

0.15

0.1

0.05

0 A. cryptopharynx A. retrodens -0.05 SPCA 2 (morphometric data) 2 (morphometric SPCA -0.1

-0.15

-0.2

-0.25 -3 -2 -1 0 1 2 3 PC 1 (meristic data)

109

0.2

0.15

0.1

0.05

A. meniscosteum 0 A. cryptopharynx A. retrodens SHRD PC2

-0.05

-0.1

-0.15

-0.2 -3 -2 -1 0 1 2 3 4 Factor 2

Figure 4.40: Plot of the second sheared principle components (morphometric data) and the second factor scores (meristic data) of the type material of Apetra retrodens (N = 113): Chembe Village PSU 4119, 4164; Apetra meniscosteum (N = 20): Kanjedza Island PSU 4130, 4134, 4163; and Apetra cryptopharynx (N = 220): Kanjedza Island PSU 4131, 4136, 4186.

Figure 4.41: Comparison of the lateral view of the keels from the holotypees of (left to right), Apetra meniscosteum PSU 4163, Apetra cryptopharynx PSU 4186, and Apetra retrodens PSU 4164.

110 Apetra meniscosteum, A. cryptopharynx, and A. retrodens are extremely similar in their outward appearance. The trouble still remains, albeit to a lesser degree, as dissection of the lower pharyngeal bone and morphometric and meristic comparison to the types is the only reliable way to diagnose these species. My work will help immensely with this problem, as now there are formal descriptions. Future researchers should rely heavily on the locality data provided in this work as a clue into species diagnosis for all of the new species described in this dissertation.

Stauffer (per. comm.) has indicated that these fishes show site fidelity for their breeding grounds. In addition, only one species in this genus is found on an arena at one time. One very important discovery that I have made is there seems to be an occasional fish of a different species collected at the same place and at the same time. This happened in the type material of Apetra intermedia for example (see Eccles and

Trewavas, 1989 and above). Occasionally I saw this happen in the PSU collections too.

There are a number of reasons that the “wrong species” could end up on the arena of another. It could be that it is just passing by and came down for a closer look to see if they found the correct place. The second reason could be that the fish made a mistake and went to the wrong place. The third possibility is that they have nowhere else to go as their species has been overfished in that area, which limits their choices for mates. We know that in the aquarium these fish readily hybridize (and produce viable offspring) due to premating isolating mechanisms being broken down and/or lack of a choice. They are

111 possibly trying to breed, although most likely will be unsuccessfully due to the choice of the other species. Further investigation of these hypotheses is needed.

Examination of the bower data taken in the field for Apetra meniscosteum from

Kanjedza Island (PSU 4134), Apetra cryptopharynx from Kanjedza Island (PSU 4105), and Apetra retrodens from Chembe Village (PSU 4084, 4095, 4119, 4164) showed separate clustering when the second principal components of the morphometric data were plotted against the third principal components of the morphometric data (Fig. 4.42). The minimum polygons formed by the three species were significantly different (p<0.05) along both the SPCA 2 (morphometric data) and SPCA 3 (morphometric data) axes independent of each other. In order to ensure proper comparison, I analyzed the morphological data for these exact populations used in the bower data. When I plotted the first principal components of the meristic data against the sheared second principal components of the morphometric data I found that the three species clustered separately

(Fig. 4.43). The minimum polygon clusters formed by the three species were significantly different (p<0.05) along the SPCA 2 (morphometric data) axis. Once again

I have validated the use of bower measurements as a taxonomic tool, and have shown its congruence with morphological data.

112

0.25

0.2

0.15

0.1

0.05

0 A. meniscosteum A. cryptopharynx A. retrodens -0.05

SPCA 3 (morphometric data) -0.1

-0.15

-0.2

-0.25

-0.3 -0.5 -0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 0.4 0.5 SPCA 2 (morphometric data)

113

0.25

0.2

0.15

0.1

A. meniscosteum 0.05 A. cryptopharynx A. retrodens

0 SPCA 2 (morphometric data) 2 (morphometric SPCA

-0.05

-0.1

-0.15 -3-2-10123 PC 1 (meristic data)

Etymology – The name retrodens, from the Latin, meaning backward (retro) tooth

(dens) and refers to the backward pointing anterior teeth on the lower pharyngeal bone.

114

Table 4.1: Character matrix for Apetra species. Species Pattern Posterior pharyngeal teeth

A. lituris dark line along upper lat. line, other horizontal elements not enlarged A. intermedia three dorsolateral spots molariform many A. variabilis incomplete band, break with ventral anterior shift cusp pointed backward A. trilineata dark horizontal elements, variable not enlarged A. linea line broken, or spotted cusp pointed backward A. simula small dark horizontal sections cusp pointed backward A. perjur dark line along upper lat. line, other horizontal elements not enlarged A. meniscosteum dark line along upper lat. line, other horizontal elements few rows enlarged varies A. cryptopharynx dark line along upper lat. line, other horizontal elements may be enlarged A. retrodens dark line along upper lat. line, other horizontal elements occasional enlargement

Species Shape of lower pharyngeal bone Rows of anterior pharyn. teeth

A. lituris inclined downwards not more than 45 degrees 4 rows A. intermedia decurved, small keel, small angle 3 rows A. variabilis steepest keel inclined at least 45 degrees swollen 6 rows A. trilineata unknown unknown A. linea steepest keel inclined at least 45 degrees swollen 6 rows A. simula extremely large keel swollen 6 rows A. perjur inclined downwards not more than 45 degrees 4 rows A. meniscosteum cresent shaped 2 rows A. cryptopharynx ~ 30 deg long top and bottom parallel, variable 2 rows A. retrodens short and blunt, dorsoventrally tall 2 rows

Species Anterior pharyngeal teeth Anterior pharyn. tooth direction

A. lituris most cylind back up to 45deg, most up at 85d A. intermedia cusp forward A. variabilis long cylindrical backward A. trilineata minute cusp backward A. linea long cylindrical backward A. simula long cylindrical, 25% outer two rows turned to midline backward A. perjur cylindrical backward at around 90 deg A. meniscosteum cusp some forward, some backward A. cryptopharynx cusp most forward A. retrodens cusp and cylindrical same bone 25% to most turned backward

115 Key to species of Apetra

1a. Three dorsolateral spots on body …..…………………………..………A. intermedia

b. Other markings on body…………….………………………………………...………2

2a. Dorsal view of lower pharyngeal bone “U” shaped with third row from the anterior

with six teeth………………………………………………………………………3

b. Dorsal view of lower pharyngeal bone “V” shaped with third row from the anterior

with less than 6 teeth………………………………………………………………5

3a. 25 % percent or more of the anterior teeth on the lower pharyngeal bone turned

toward the midline……………………………...…...……………………A. simula

b. Less than 25 % of the anterior teeth on the lower pharyngeal bone turned toward the

midline………………………………………………………………….…………4

4a. Pattern consists of a broken line with a ventral anterior shift at the

break………………………………………………………………...…A. variabilis

b. Pattern consists of a single horizontal element, which may consist of a broken line or

series of spots, but with no shift or overlapping portions…………….…….A. linea

5a. 25 % or more of the anterior teeth on the lower pharyngeal bone turned toward the

posterior…………………………………………………………………………...6

b. Less than 25 % of the anterior teeth on the lower pharyngeal bone turned toward the

posterior…………………………………………………………………………...8

6a. Anterior lower pharyngeal bone teeth turned toward the posterior at more than 45o

with most approaching 90o…………………………………..……………A. perjur

b. Anterior lower pharyngeal bone teeth turned toward the posterior at 45o or less…....7

116 7a. All of the anterior teeth turned toward the posterior, cheek depth 28-38 % SL

………………………………………………..…………………………….…..…A. lituris

b. Some of the anterior teeth turned toward the anterior while others turned toward the posterior, cheek depth 20-30 % SL ………………………………………...…A. retrodens

8a. Horizontal eye diameter 36-45 % HL, vertical eye diameter 34-43 % HL

………………………………………………………………………..….A. cryptopharynx

b. Horizontal eye diameter 31-40 % HL, vertical eye diameter 31-38 % HL…………..9

9a. Snout length 29-38 % HL, head length 27-30 % SL………...………A. meniscosteum

b. Snout length 42 % HL, head length 33 % SL………………....…………..A. trilineata

Chapter 5

Discussion and Conclusions

Tramitichromis was diagnosed by Eccles and Trewavas (1989) by the presence of a keel on the lower pharyngeal bone as well as three or more rows of teeth extending to the end of the bone, which is rounded. I have observed the latter characteristic only in A.

[T.] variabilis, A. linea, and A. simula. With the PSU collections, I found fish with a keel that had an affinity for (phenotypic similarity to) certain species, but could not be identified using the dichotomous key provided by Eccles and Trewavas (1989). In addition, certain fish had observable morphological differences in their lower pharyngeal bone but were identified as the same species using Eccles and Trewavas (1989). What this meant is that there were unknown and/or cryptic species in the lake that were not diagnosed.

In summary, Tramitichromis is now a monotypic genus. Some preliminary evidence suggests it may contain more species, but no formal descriptions have been made (Snoeks, 2004). The new genus, Apetra, contains ten species. Apetra lituris is the type species. The type material of A. lituris contained two species, one that retains the name for which I designated a lectotype, and the other A. perjur. Apetra meniscosteum,

A. cryptopharynx, and A. retrodens are three new species that are most similar to A. lituris, but smaller. I also had bower data for these fishes that reinforced the species descriptions. Another species, A. trilineata, was restricted to the individual that resides in the jar at the Natural History Museum, London, as its lower pharyngeal bone was

118 missing, and the description was based on one specimen from an unknown locality. No other fish were identified as this species in my study from any of the other collections in the Natural History Museum, London or the PSU fish museum. The A. variabilis type material contained two species, one which retains the name and for which I designated a lectotype, and A. linea. I discovered a third species, A. simula, which was very similar to the previous two. Apetra intermedia showed variations in the amount of posterior pharyngeal tooth enlargement, but no other differences were observed. Collection localities were mapped for all of the above species.

My suggestion for anyone studying the Tramitichromis and Apetra is to preserve specimens from each locality and then dissect the lower pharyngeal bone. It is the only way to accurately identify the species being observed. Examination of the literature revealed many times when the specimens had been misidentified. Subsequent versions of

Malawi cichlid books or later examination of the fish used reveals species identity corrections. Even in my feasibility study, I thought I had Lethrinops when it was actually

Apetra. Behavior is an accurate way to distinguish the genera, but many of the species listed above can be easily confused for one another if diagnosis is based solely on external appearances.

In the past, it has been shown that females are rather choosy when selecting a male with which to spawn. The female will approach a breeding arena, circle with numerous males, but only breed with a few (Stauffer and Kellogg, 1996). It has been suggested that these females are choosing the best males with which to spawn (Stauffer and Kellogg, 1996; Kellogg et al., 2000). Now I do not doubt that this is happening, but most likely, the females are looking to see if the males are the same species first. Bowers

119 are one of the preliminary characteristics females use for mate selection (Stauffer et al.

2002). This would explain the fact that two of the newly discovered species that live sympatricly at Kanjedza Island also have distinct bowers. The females may be cueing in on other subtle interspecific differences as well such as appearance, sounds, courtship dance, etc., all of which have been shown to be important. Similar observations can be made for A. variabilis, A. linea, and A. simula around localities near Cape Maclear.

Usually one species of fish spawns in each breeding arena (lek). Sometimes, other species in other genera spawn in the same lek, but they are not very closely related.

It has generally been assumed that there is only one species of Tramitichromis or Apetra spawning in each lek. What I have discovered, is that there many be more than one species of Apetra spawning in the same lek. This will change many assumptions scientists working on Lake Malawi cichlids have made. No longer can a few sample individuals be taken from one site, because there may be many species there.

I noted an occasional fish that was the “wrong species” found at a particular arena. It could be that the fish came in for a closer look to see if the correct species was present, the fish made a mistake and went to the wrong place, or the fish did not have a choice as its population could have been over-fished. This issue is in need of research, and the frequency of this problem needs to be quantified.

Apetra intermedius did show some variation in the shape of the keel when viewed from the lateral, but I was not able to find any other differences. Some literature has indicated that there may be separate species (Konings, 2001; Turner, 1996), but I was unable to find any correlations between keel shape and anything else. It is possible that I did not have enough fish in the collections, or not enough collections from different

120 localities. Nevertheless, this species seems to be rather widespread in the lake, so it is worth investigating any observed variations.

Laboratory studies need to be done to show the heritability of bower shapes. I have identified ways in which this can be done, and have shown suggestive evidence of this. The next step in the process would be to hybridize the bower builders to see if the shapes built are intermediate, or follow Mendelian principles.

I have shown congruence between bower data and morphology data. Future work on bower shapes in the field should be measured according to the diagrams in chapter 3.

In addition, I think an additional characteristic should be measured: from the center of one bower, to the center of the surrounding bowers. It could be that different species have different spaces that they defend and therefore place their bowers a certain distance apart. Bower measurements need to be obtained for A. lituris, A. intermedia, A. variabilis, A. trilineata (if it can be found), A. linea, A. simula, and A. perjur.

Now that species have been diagnosed, fisheries managers can accurately identify populations and develop range maps. Population sizes for each species need to be obtained as well as sustainable harvest estimates. Eventually management plans can be developed lake wide.

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124 strategies and tactics (eds. R. J. Wooton and G. W. Potts). Academic

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and Fortune Shonhiwa. 1993. Sexual Selection and Genetic Relationships

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to Control Vectors of Parasitic Disease: Schistosomiasis in Lake Malawi –

A Case History. BioScience 47: 41-49.

127 Stauffer, Jay R., N. J. Bowers, K. A. Kellogg, and K. R. McKaye. 1997b. A

Revision of the Blue-black Pseudotropheus zebra (Teleostei: Cichlidae)

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Aquariculture and Aquatic Sciences: Cichlid Research: State of the Art. 9:

1-16.

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128 Trewavas, E. 1931. A Revision of the Cichlid Fishes of the Genus Lethrinops,

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Appendix A

Laboratory Feasibility Study

A.1 Introduction

Bower building is the manifestation of a behavioral trait (Stauffer et al., 1996;

Kellogg et al., 2000; Stauffer et al., 2002). The problem with behavior is that it can be learned or innate. The question of whether or not bower building, and therefore the associated species-specific bower shape, is heritable needs to be answered. The purpose of this portion of my research was to determine if it was possible to test this in a laboratory setting, and give comments and suggestions for future research. What confounded the problem I was investigating is that cichlids, especially African cichlids from Lake Malaŵi, are known to rearrange tank furniture (small rocks, gravel, sand, decorations). This includes moving rocks, and piling up the substrate in some places, while exposing the floor of the aquarium in others. Even rock-dwelling fish that do not have access to movable substrates in nature do this in the aquarium. Simply put, to see sand piled up or moved would not mean that a bower has been constructed and therefore is not significant. Only sand piled up in a cone shape, and that shape strictly maintained by its owner, would indicate that a bower has been constructed. A courtship dance of the bower building male trying to lure a female to mate with him confirms the structure’s use as spawning platform (Stauffer et al., 1996). Bowers are one of the preliminary characteristics females use for mate selection (Stauffer et al. 2002). I therefore

131 hypothesized that different species will have different shaped bowers, and that bower building is a heritable trait.

A.2 Methods

In order to ensure that the fish in this experiment did not learn how to build bowers from their fathers, the fry were pulled from the mother’s mouth before she released them into the tank. In addition, the tanks that the parent fish were kept in did not have a large enough area or depth of sand for a bower to be built in case the mother temporarily released the fry to feed. If it was observed that the female had temporarily released the fry, then that batch of fish was not used. The fry were raised in isolation from the parents where they could not see the parental tank. Once the fry reached sexual maturity, they were used in the experiment. The fish that were used in this experiment were F3 and F4 Apetra cryptopharynx, but were labeled Lethrinops sp.

A sand substrate about 25.4 cm deep, evenly spaced along the bottom was placed in three 1514-liter circular tubs. Filtration equipment was housed outside the tank and siphon and return line hoses were located on the insides of the tank. Water was changed as needed, the same amount being changed in each tub (usually 25%). The fish were fed all the Tetra Cichlid Sticks and Tetra Cichlid Flake they could consume in 5 min.

The bower-base diameter in this species is roughly 74.47 cm (Kellogg et. al.,

2000). The pool base was 162.5 cm. Cichlids generally partition available space, provided they each have enough, so there should have been room for three bowers in each tub if the fish partitioned the area equally (Stauffer, per. com.). The fish were kept

132 in these pools until the females were brooding eggs, or no more work was done on the bowers. A period of about two weeks should have been sufficient for this to happen, but

I left the fish in the tanks for a few months (McKaye, 1991). At that time any bowers were measured and photographed (if possible). In between trials, the sand was smoothed to eliminate the previous males’ bowers and have the layer of sand even along the bottom of the tank (McKaye et. al., 1990).

Initially, two experimental groups using F3 fish were set up with three males and four females per pool. After this first run was over, the fish were euthanized, and another trial was done, except that this time, I used F4 fish. Two experimental groups containing

three males and six females were used as I thought more females would stimulate more

bower building.

During the time of these experiments, another tub was established with three

males and four females. The fish were allowed to breed in the tub, but the fry were not

removed, but allowed to observe their parents’ behaviors. The parents were euthanized at

the same time as the second trials started in the other two pools. The fish in this pool

were used to see if there is a difference between fish raised with the parents vs. fish raised

in isolation.

Methods for data collection and analysis followed Stauffer et al. (1993). Bower

measurements were taken according to Stauffer et. al. (1993) and include: width of base,

slope length, outside top diameter, inside top diameter, and bower height (Fig. A.1). The

same measurements were taken for both the lab and in situ bowers (see Ch. 2 methods).

133

C D

B F E

A.3 Results and Discussion

Some problems that are associated with performing laboratory experiments of this magnitude were discovered. First of all, I tried to produce a natural setting in the lab, but that is essentially impossible. The pools were big, but they still restricted the movement of the fish. They did not have the vast expanses that they would have had in the lake.

134 Secondly, there were no predators or natural prey items. Third, in the wild the breeding arenas are huge, and males have countless males with which they must compete. In the lab setting, one or two aggressive males can dominate a tank, or in this case, even the large pools. The males may not need to put a lot of time and effort into building the bowers to mate. Female choice is also limited. Finally, seasonal cues and cycles are not present, so artificial means, like abundant food and large water changes, were done in order to stimulate spawning.

For the heritability portion of the question, I have provided the answer. The baby fish were third and fourth generation captive bred fish raised in isolation from their parents. The fish were not allowed to observe their parents, and the parental tanks did not have the space for the construction of a full bower. This should have removed any possibility of learning. When I placed the fish into the 1514-liter pools, they did in fact build bowers (Fig. A.2). Although it is hard to see from the photos, the bowers are the characteristic cone-shaped bowers of the genus Apetra.

135

136 To see cone-shaped bowers was surprising, as the fish I was using were labeled as

Lethrinops c.f. parvidens in these experiments. When I also noted a “figure eight” courtship pattern, I began to wonder whether or not I actually had Lethrinops. Inspection of the lower pharyngeal bone on the euthanized fish revealed that I was dealing with a member of the genus Apetra due to the presence of a keel as well as the correct dentition

(Fig A.3c-e). The exact species was not determined as the importance of the feasibility study was to see if the fish would/could build bowers.

137

138 I actually discovered the correct ratio of males to females needed to produce strict bowers by chance. In the first round of tests, I used groups of three males and four females in two pools. The first pool in the first trial produced one poorly built bower.

The other two males in the tank did not seem to be interested in breeding either because they were not physiologically able to do so, or because the third male treated the whole pool as his territory, so they never got the chance. The dominant fish did not seem to need the bower to attract females to spawn, as he just defended the whole pool.

The second pool produced two less poorly built bowers. The third male did not breed for one of the above two reasons. The competition between the two breeding males did seem to produce bower shapes that more closely resembled the cone shaped bowers.

There was rigorous fighting, and they finally did coexist, each claiming one-third of the tub. Females were observed to breed with both fish.

In the second round of tests (replication of the first), the same results were obtained for one pool. In the other, however, I had a hard time sexing the small fish and ended up placing seven males and two females together. This happened to produce the best results. Three of the males built strict cone-shaped bowers. The shape was maintained throughout the experiment, but the height did increase. The reason I believe it worked so well is because the males had to compete for space. If one did not defend its bower, then another male could take its place, just like in the wild. Also, by having seven males in the tank, it increased the chances that three would be ready to breed at all times as well as decreasing the ability of one male dominating the pool. In addition, there were only two females, which meant the males had to work very hard to attract a mate.

139 Although I did get the fish to build bowers, and thereby show suggestive evidence of the heritability of bower building, I wanted to have quantitative data to back up my observations. I tried to measure the bowers, but ran into a problem. Because the pools are round, the fish built their bowers with one edge along the side (Fig. A.4). This prevented me from taking two sets of measurements, the second at 90 degrees to the first.

I could not, however, take accurate measurements of the first set due to the curving of the outside of the pool wall. When the complete base width was made, I was left with about one-third of the bower actually being built. What would have been needed, was the fish to build a bower in the center of the pool, with none of it touching the pool sides.

140

Figure A.4: Diagram of bowers (gray) built in the pool (clear). Two-thirds of the bower was not built, as it would extend beyond the walls of the pool.

In order to get this and similar lab experiments to work, I would recommend a ratio of seven males to two females. A much larger set up would be required. An actual swimming pool with a fish safe liner would work well giving the fish enough space to spread out. My concern, though, is that in order to maximize their distance from other males, they would still choose to build along the wall. A square shaped pond at least 3m by 3m would be better, as at least one dimension would be usable when the experimenter measured the bowers provided the fish did not start too close to the pond wall (Fig. A.5).

The goal would be to have the fish start building the bowers in the center of the pond. In

141 the lake, the male in this position has the most fertilizations (Kellogg et al., 2000). It could be that the fish learn which location is best, or the pool set up makes them feel too uncomfortable to build bowers in the center. The depth of sand used in this study was perfect, as none of the males dug down to the bottom of the pool. I would first condition the fish in tanks and get them used to captivity, prepared food, humans, and water changes. I would not expose them to any substrate. After I felt the fish were comfortable and used to the daily routine, I would then place them in the pond at the 7:2 ratio of males to females. Once bower building began, I would stop the experiment two or three weeks after, and then measure the bowers. Repeating the experiment would be beneficial, but if enough males are used (more than twenty bowers measured), then it could be terminated at that point.

142

Figure A.5: Suggested pond structure and expectant bower placement. Only the center three bowers would be completely useful.

Appendix B

Tables of Morphometric and Meristic Values

Table B.1: Morphometric and meristic values of the Tramitichromis brevis population from Cobue (N = 24) PSU 4089.

Tramitichromis brevis Cobue M ean Standard Range Deviation min max St andard lengt h, mm 88.8 16.3 63.3 110.2 Head lengt h, mm 29.9 5.3 20.9 36.6 Percent o f st and ard leng t h Head length 33.7 0.9 32.4 35.7 Snount to dorsal-fin origin 39.8 1.3 37.6 43.4 Snount to pelvic-fin origin 40.4 1.1 37.7 42.9 Dorsal-f in base lengt h 55.1 1.6 51.4 57.7 Anterior dorsal to anterior anal 49.3 2.1 45.4 52.4 Post erior dorsal t o post erior anal 15.9 0.7 14.4 17.3 Anterior dorsal to posterior anal 59.4 1.6 56.6 61.7 Posterior dorsal to anterior anal 31.4 1.0 30.0 33.3 Post erior dorsal t o vent ral caudal 19.8 0.9 18.3 21.6 Post erior anal t o dorsal caudal 22.5 0.8 21.2 23.9 Anterior dorsal to pelvic-fin origin 39.4 1.3 36.8 42.2 Posterior dorsal to pelvic-fin origin 53.7 1.2 51.1 55.8 Caudal peduncle lengt h 16.5 1.0 14.4 18.0 Least caudal peduncle dept h 12.2 0.4 11.5 12.8 Body depth 37.2 1.2 35.3 40.0 Percent head leng t h Horizont al eye diamet er 39.7 3.4 35.3 47.5 V ert icle eye diamet er 38.3 2.8 34.7 43.1 Snout length 38.6 2.5 32.7 41.8 Postorbital head length 41.0 2.1 35.4 44.7 Preorbital dept h 21.8 2.0 18.8 25.8 Lower-jaw lengt h 39.8 1.4 37.3 42.3 Cheek depth 29.5 1.9 25.7 32.8 Head depth 97.4 4.9 89.7 108.6 Counts M ode %Freq. Range Lat eral-line scales 32 70.8 31 33 Pored scales posterior to lateral line 2 75.0 1 3 Scale rows o n cheek 3 9 5.8 2 3 D o r s al - f i n s p i nes 15 9 1. 7 15 16 Dorsal-fin rays 11 83.3 10 12 A nal-f in sp ines 3 100.0 3 3 Anal-fin rays 9 45.8 8 10 P ec t o r al - f i n r ay s 16 70 . 8 15 17 Pelvic-fin rays 5 100.0 5 5 Gill rakers on first ceratobranchial 7 54.2 6 8 Gill raker on first epibranchial 4 58.3 3 4 Teeth in outer row of left lower jaw 11 29.2 9 14 Teeth rows on upper jaw 3 62.5 2 4 Teet h ro ws o n lo wer jaw 4 6 6.7 4 5

144

Table B.2: Morphometric and meristic values of Apetra lituris type material, which includes the lectotype (N = 32) from Karonga, BMNH 1930.1.31.21-23; Vua, BMNH 1930.1.31.24-28; Mwaya, BMNH 1930.1.31.35-44. Morphometric and meristic values of the Apetra lituris holotype BMNH 1930.1.31.21 are also listed.

Apetra lituris Lectotype Types Mean Standard Range Deviation min max Standard length, mm 125.2 107.3 12.4 83.0 132.9 Head length, mm 40.9 35.4 3.8 27.2 42.3 Percent o f st and ard leng t h Head length 32.7 33.0 0.8 31.3 34.7 Snount to dorsal-fin origin 40.2 39.4 1.2 37.1 42.7 Snount to pelvic-fin origin 40.6 40.3 1.2 38.5 44.1 Do rsal-f in b ase leng t h 56 .8 54.2 1.4 50 .8 56.8 Anterior dorsal to anterior anal 51.8 48.7 1.4 46.5 51.8 Post erior d orsal t o p ost erio r anal 17.5 16.2 0 .6 15.3 17.5 Anterior dorsal to posterior anal 61.4 58.2 1.3 55.9 61.4 Post erior dorsal t o ant erior anal 32.2 29.8 1.0 27.7 32.2 Posterior dorsal to ventral caudal 23.0 20.9 0.8 19.4 23.0 Post erior anal t o dorsal caudal 24.5 23.2 0.7 21.6 24.5 Anterior dorsal to pelvic-fin origin 40.3 36.6 1.4 34.5 40.3 Posterior dorsal to pelvic-fin origin 51.0 49.7 1.4 45.9 52.2 Caudal peduncle lengt h 17.9 18.0 0.8 16.2 20.2 Least caudal peduncle dept h 11.8 11.5 0.4 10.5 12.1 Body depth 38.6 35.4 1.1 33.5 38.6 Percent head leng t h Horizont al eye diamet er 31.4 35.6 1.9 31.4 39.4 V ert icle eye diamet er 29.3 32.8 1.7 29.3 35.3 Snout length 44.3 40.5 1.9 36.2 44.3 Postorbital head length 45.2 43.4 1.4 40.4 46.5 Preorbital depth 27.5 26.1 1.5 21.6 28.7 Lower-jaw lengt h 38.9 38.9 1.9 34.9 42.3 Cheek depth 35.3 31.9 2.3 26.1 37.6 Head depth 100.5 92.6 3.6 84.0 100.5 C ount s M ode %Freq. Range Lat eral-line scales 32 33 53.1 31 34 Pored scales posterior to lateral line 1 1 68.8 0 3 Scale rows o n cheek 4 4 56.3 3 4 Dorsal-fin spines 15 16 68.8 15 16 Dorsal-fin rays 11 11 59.4 10 12 Anal-fin spines 3 3 100.0 3 3 Anal-fin rays 9 9 84.4 8 10 Pectoral-fin rays 16 16 81.3 14 16 Pelvic-fin rays 5 5 100.0 5 5 Gill rakers on first ceratobranchial 8 9 46.9 7 10 Gill raker on f irst epibranchial 4 3 56.3 2 4 Teeth in outer row of left lower jaw 13 14 40.6 13 16 Teet h ro ws o n up p er jaw 4 3 71.9 2 4 Teet h ro ws o n lo wer jaw 5 4 71.9 4 5

145

Apetra intermedius Lectotype Types Chembe V illage, Cape M aclear Golden Sands Swamp, Cape M aclear M ean StandardRange M ean StandardRange M ean Standard Range Deviation min max Deviation min max Deviation min max St and ard leng t h, mm 123.3 101.5 2 2.9 6 3.7 123 .3 91.3 17.3 6 1.1 113.3 105.8 7.6 100.5 111.2 Head length, mm 39.0 32.4 6.9 21.2 39.0 29.1 5.9 18.8 37.0 34.1 3.6 31.5 36.7 Percent of standard length Head length 31.6 32.1 0.7 31.5 33.4 31.7 1.1 29.0 34.2 32.2 1.1 31.4 33.0 Snount to dorsal-fin origin 37.3 38.5 0.7 37.3 39.4 39.3 4.1 36.4 54.9 38.0 2.0 36.6 39.4 Snount to pelvic-fin origin 39.3 39.7 1.2 38.8 42.0 39.5 1.4 37.3 43.1 38.0 1.0 37.3 38.6 Dorsal-f in b ase leng t h 57.7 56.0 2.6 51.3 58.7 54.7 1.7 51.0 57.1 56.1 0.3 55.9 56.3 Anterior dorsal to anterior anal 51.3 50.5 2.5 46.2 53.8 49.5 1.5 47.1 52.0 49.0 0.7 48.5 49.5 Po st erior do rsal t o p ost erio r anal 17.4 16.8 1.0 15.5 18.1 16.2 0 .8 14.7 17.6 15.0 0.2 14.8 15.2 A nt erior dorsal t o p ost erio r anal 6 1.3 59.9 2.9 55.5 63.5 59.2 1.4 56.8 61.8 60.0 0.4 59 .7 60.3 Po st erior do rsal t o ant erior anal 3 1.7 30.8 1.6 2 8.5 3 2.3 3 0.6 1.1 28.9 32.3 30.3 0.1 30 .3 30.4 Posterior dorsal to ventral caudal 21.2 20.7 0.5 19.9 21.2 21.0 0.9 20.2 23.4 21.4 0.7 20.9 21.8 Po st erior anal t o do rsal caudal 2 4.1 23.4 0.7 22 .6 24.1 23.1 0 .8 21.6 24 .7 23.2 0.2 23 .0 23.3 Anterior dorsal to pelvic-fin origin 39.5 38.4 2.2 35.0 41.5 37.7 1.4 35.5 40.0 37.1 0.5 36.7 37.5 Po st erior do rsal t o p elvic-f in origin 52.8 51.4 1.6 48 .6 52.8 52.3 1.3 50 .1 54.9 51.4 0.4 51.1 51.7 Caudal p eduncle leng t h 18.7 18.0 0.8 16.7 19.2 17.4 1.2 15.8 20 .7 17.7 0.3 17.4 17.9 Least caudal peduncle dep t h 12.2 11.5 0.5 10.7 12.2 11.7 0 .5 10.8 12.5 11.7 0.0 11.6 11.7 Body depth 36.4 36.1 1.2 34.6 38.1 35.4 1.1 33.7 37.3 35.0 1.1 34.2 35.7 Percent head lengt h Horizontal eye diameter 32.3 33.4 3.4 30.1 39.0 36.5 3.0 32.4 43.2 35.0 0.4 34.7 35.3 Verticle eye diameter 30.7 32.9 2.8 30.2 37.5 35.4 2.8 32.0 41.6 32.7 1.5 31.6 33.7 Snout length 41.7 40.1 2.8 36.3 42.5 38.9 3.5 30.8 42.7 39.1 0.5 38.8 39.5 Po st orbit al head lengt h 42 .7 43.0 0.9 4 1.9 44.1 41.2 1.8 37.7 44.0 43.5 0.7 43 .0 44.0 Preorbital depth 26.2 23.3 2.5 19.2 26.2 21.0 2.5 16.5 24.3 22.4 1.2 21.5 23.2 Lower-jaw length 39.3 38.0 1.9 35.3 40.1 39.9 1.0 37.9 41.8 40.8 0.3 40.6 41.0 Cheek depth 33.5 30.5 3.4 25.8 33.5 29.5 3.5 21.3 33.8 32.7 0.7 32.3 33.2 Head depth 101.8 98.3 7.8 84.2 105.6 99.3 5.1 89.4 105.1 96.5 0.5 96.2 96.8 Counts M ode %Freq.Range M ode %Freq.Range M ode %Freq. Range Lat eral-line scales 32 32 6 6.7 3 1 32 3 2 50.0 31 33 # N/ A 0.0 32 34 Po red scales po st erior t o lat eral line 1 1 50 .0 0 2 2 44.4 0 4 # N/ A 0.0 1 2 Scale ro ws on cheek 3 3 8 3.3 3 4 3 72.2 2 4 3 100.0 3 3 D o r s al - f i n s p i nes 15 15 10 0 . 0 15 15 15 6 6 . 7 14 16 15 10 0 . 0 15 15 D o r s al - f i n r ay s 10 11 6 6 . 7 10 11 10 4 4 . 4 9 13 11 10 0 . 0 11 11 A nal-f in spines 3 3 100.0 3 3 3 100 .0 3 3 3 100.0 3 3 Anal-fin rays 9 9 83.3 8 9 9 88.9 8 9 9 100.0 9 9 P ec t o r al - f i n r ay s 15 15 6 6 . 7 14 16 15 77. 8 15 16 15 10 0 . 0 15 15 Pelvic-fin rays 5 5 100.0 5 5 5 100.0 5 5 5 100.0 5 5 Gill rakers on first ceratobranchial 8 8 50.0 7 10 8 55.6 7 12 8 100.0 8 8 Gill raker on f irst epibranchial 4 3 50 .0 2 4 3 77.8 2 4 3 100.0 3 3 Teeth in outer row of left lower jaw 14 14 50.0 12 16 15 27.8 13 19 15 100.0 15 15 Teet h rows o n up per jaw 3 3 100.0 3 3 2 50.0 2 4 3 100.0 3 3 Teet h rows o n lower jaw 4 4 8 3.3 4 5 4 77.8 2 5 4 100.0 4 4

146

Table B.4: TABLE A.3 (concluded)

Apetra intermedius Kanjedza Island M ean Standard Range Deviation min max Standard length, mm 93.4 30.6 60.8 141.5 Head length, mm 29.8 9.4 19.6 44.6 Percent of standard length Head length 32.0 1.0 30.1 34.8 Snount to dorsal-fin origin 38.4 1.1 35.9 41.7 Snount to pelvic-fin origin 38.8 1.0 36.4 41.0 Dorsal-fin base length 54.8 2.4 50.3 60.4 A nt erior dorsal t o ant erior anal 50.0 2.1 45.7 54.6 Posterior dorsal to posterior anal 16.7 0.6 15.5 18.5 A nt erior dorsal t o post erior anal 59.7 2.3 56.2 65.2 Posterior dorsal to anterior anal 30.5 1.5 27.8 34.4 Posterior dorsal to ventral caudal 21.1 0.9 18.9 23.4 Post erior anal t o dorsal caudal 23.2 1.0 21.6 25.6 Anterior dorsal to pelvic-fin origin 37.7 2.0 34.0 42.5 Posterior dorsal to pelvic-fin origin 52.2 1.9 47.7 55.8 Caudal peduncle length 17.7 0.8 16.1 19.3 Least caudal peduncle dept h 11.7 0.6 10.3 13.2 Body depth 35.8 1.5 32.9 38.9 Percent head leng t h Horizontal eye diameter 36.2 4.3 29.4 44.3 Verticle eye diameter 34.7 3.6 28.8 40.3 Snout length 37.5 3.2 32.5 42.7 Post orbit al head lengt h 41.2 2.2 37.2 47.6 Preorbital depth 22.4 3.6 16.2 28.3 Lower-jaw length 39.7 1.5 36.0 42.9 Cheek depth 29.8 4.5 22.8 38.2 Head depth 97.0 7.9 84.8 110.0 Counts M ode %Freq. Range Lat eral-line scales 32 61.2 30 33 Pored scales posterior to lateral line 2 69.4 0 2 Scale ro ws o n cheek 3 85.7 2 4 D o r s al - f i n s p i nes 15 77. 6 14 16 Dorsal-fin rays 11 61.2 10 11 Anal-fin spines 3 100.0 3 3 Anal-fin rays 9 81.6 8 10 P ec t o r al - f i n r ay s 15 79 . 6 14 17 Pelvic-fin rays 5 100.0 5 5 Gill rakers on f irst cerat obranchial 7 34.7 7 15 Gill raker on first epibranchial 3 81.6 2 4 Teeth in outer row of left lower jaw 16 18.4 10 21 Teet h ro ws on upp er jaw 3 61.2 2 4 Teet h ro ws on lo wer jaw 4 95.9 3 5

147

Apetra variabilis Lectotype Types Mean Standard Range Deviation min max St andard lengt h, mm 104.4 9 5.3 9 .8 8 2.1 117.1 Head length, mm 35.2 31.5 3.5 27.6 39.6 Percent o f st and ard leng t h Head length 33.7 33.0 0.8 31.6 34.3 Snount to dorsal-fin origin 37.2 37.9 1.7 34.7 41.0 Snount to pelvic-fin origin 42.9 41.5 1.5 39.7 43.9 Do rsal-f in b ase leng t h 53 .8 53.0 1.0 51.8 54.7 Anterior dorsal to anterior anal 50.1 48.5 1.2 47.1 51.0 Post erior d orsal t o p ost erio r anal 17.3 16.2 0 .6 15.5 17.3 Anterior dorsal to posterior anal 59.2 57.7 1.1 56.0 59.2 Post erior dorsal t o ant erior anal 30.6 29.2 0.7 28.3 30.6 Posterior dorsal to ventral caudal 21.1 21.1 0.6 20.1 22.2 Post erior anal t o dorsal caudal 23.5 23.2 0.5 22.0 23.9 Anterior dorsal to pelvic-fin origin 36.7 36.1 0.9 34.1 37.7 Posterior dorsal to pelvic-fin origin 49.0 48.9 0.7 47.9 50.3 Caudal peduncle lengt h 17.4 18.1 0.6 17.2 19.7 Least caudal peduncle dept h 11.4 11.2 0.3 10.6 11.7 Body depth 35.8 35.2 0.9 33.8 37.1 Percent head leng t h Horizont al eye diamet er 32.9 35.4 1.6 32.9 37.6 V ert icle eye diamet er 31.2 33.6 1.6 31.2 36.8 Snout length 41.8 39.9 1.4 36.8 41.8 Postorbital head length 46.2 44.7 2.4 41.5 49.3 Preorbital dept h 24.8 23.1 1.3 20.6 24.8 Lower-jaw length 32.5 35.6 1.7 32.5 38.2 Cheek depth 32.1 29.0 1.6 26.1 32.1 Head depth 88.8 93.2 4.0 88.7 102.5 C ount s M ode %Freq. Range Lat eral-line scales 32 32 83.3 32 33 Pored scales posterior to lateral line 1 1 58.3 0 2 Scale rows o n cheek 3 3 75.0 3 4 Dorsal-fin spines 15 15 75.0 14 16 Dorsal-fin rays 11 11 75.0 10 12 Anal-fin spines 3 3 100.0 3 3 Anal-fin rays 9 9 66.7 8 9 Pectoral-fin rays 15 15 75.0 14 16 Pelvic-fin rays 5 5 100.0 5 5 Gill rakers on first ceratobranchial 9 9 58.3 7 10 Gill raker on first epibranchial 3 3 91.7 3 4 Teeth in outer row of left lower jaw 14 13 50.0 12 15 Teeth rows on upper jaw 4 3 66.7 2 4 Teeth rows on lower jaw 4 4 100.0 4 4

148

Table B.6: Morphometric and meristic values of the Apetra trilineata holotype from an unknown locality BMNH 1930.1.31.76.

Apetra trilineata Holot ype

Standard length, mm 103.7 Head length, mm 34.5 Percent of st andard lengt h Head length 33.3 Snount to dorsal-fin origin 39.0 Snount to pelvic-fin origin 41.0 Dorsal-fin base length 54.7 Anterior dorsal to anterior anal 48.1 Posterior dorsal to posterior anal 15.7 A nt erior dorsal t o post erior anal 58.7 Post erior dorsal t o ant erior anal 30.5 Posterior dorsal to ventral caudal 20.0 Post erior anal t o dorsal caudal 22.6 Anterior dorsal to pelvic-fin origin 36.1 Posterior dorsal to pelvic-fin origin 50.4 Caud al p ed uncle leng t h 17.6 Least caudal peduncle dept h 11.3 Body depth 34.1 Percent head leng t h Ho rizo nt al eye d iamet er 3 3.9 V ert icle eye diamet er 32.0 Snout length 41.9 Post orbit al head lengt h 42.4 Preorbit al dept h 20.7 Lower-jaw length 35.7 Cheek dep t h 2 7.1 Head depth 89.2 C ount s Lat eral-line scales 33 Pored scales posterior to lateral line 1 Scale ro ws o n cheek 3 Dorsal-f in spines 16 Dorsal-fin rays 11 A nal-f in spines 3 Anal-fin rays 10 Pectoral-fin rays 15 Pelvic-fin rays 5 Gill rakers on first ceratobranchial 9 Gill raker on first epibranchial 4 Teeth in outer row of left lower jaw 13 Teet h ro ws on up per jaw 4 Teet h ro ws on lower jaw 4

149

Apetra linea Holot ype Types Fisheries Research Station M ean StandardRange M ean Standard Range Deviation min max Deviation min max Standard length, mm 95.1 97.0 20.7 57.0 129.8 103.9 13.2 84.1 142.0 Head length, mm 29.8 30.4 6.3 17.8 39.1 33.7 4.0 27.7 44.0 Percent of st andard lengt h Head length 31.3 31.4 0.7 30.1 32.9 32.4 0.7 30.8 33.6 Snount to dorsal-fin origin 38.0 38.0 0.9 36.1 38.9 37.6 1.0 35.5 39.5 Snount to pelvic-fin origin 39.4 39.1 1.3 37.0 41.1 39.7 1.1 37.1 41.1 Dorsal-fin base length 54.7 54.1 1.0 52.4 55.1 55.2 1.2 52.6 57.6 Anterior dorsal to anterior anal 49.8 48.2 1.6 44.6 50.4 50.0 1.5 47.2 52.9 Posterior dorsal to posterior anal 16.3 15.9 0.4 15.4 16.5 16.1 0.8 14.0 17.1 Anterior dorsal to posterior anal 59.9 58.2 1.3 55.3 59.9 59.0 2.5 50.1 61.9 Posterior dorsal to anterior anal 30.4 30.2 0.9 28.8 31.6 30.6 1.1 28.4 32.2 Posterior dorsal to ventral caudal 21.1 21.0 0.7 20.1 22.3 21.8 0.7 21.0 23.5 Posterior anal to dorsal caudal 23.2 23.4 0.5 22.7 24.3 23.8 0.7 22.5 25.1 Anterior dorsal to pelvic-fin origin 35.5 35.3 1.4 32.8 37.2 37.6 1.2 35.7 40.5 Posterior dorsal to pelvic-f in origin 50.2 49.8 1.7 46.4 52.2 51.3 1.2 49.1 53.6 Caudal peduncle length 17.1 18.1 0.9 17.1 19.3 17.6 0.6 16.3 18.4 Le as t c a ud a l p e d un c l e d ep t h 11. 2 11. 5 0 . 5 10 . 8 12 . 3 12 . 0 0 . 5 11. 0 12 . 9 Body depth 34.2 34.1 1.4 31.3 35.9 34.9 0.8 33.0 36.3 Percent head lengt h Horizontal eye diameter 36.5 36.6 2.5 33.6 42.2 36.4 1.4 32.9 38.7 Verticle eye diameter 36.4 35.4 2.4 32.6 39.0 35.9 1.6 33.1 38.9 Snout length 38.8 40.0 3.0 33.9 44.9 40.8 1.6 38.5 43.9 Postorbital head length 44.8 44.9 1.6 42.4 47.5 43.3 1.9 40.0 47.2 Preorbital depth 25.3 24.1 2.6 19.6 27.6 22.3 1.7 19.6 26.4 Lower-jaw length 36.8 36.6 0.8 35.3 37.3 38.2 2.0 34.1 41.5 Cheek depth 28.2 27.6 3.4 19.9 33.0 31.5 2.3 25.9 33.9 Head depth 91.5 91.7 3.9 85.4 98.0 98.7 4.0 90.3 106.6 Counts M ode %Freq.Range M ode %Freq. Range Lateral-line scales 33 33 60.0 32 33 34 60.0 31 34 Pored scales posterior to lateral line 1 1 60.0 1 2 1 50.0 0 3 Scale rows on cheek 3 4 60.0 3 4 3 85.0 3 4 D o r s a l - f i n s p i n es 16 16 7 0 . 0 15 16 15 70 . 0 14 16 Dorsal-fin rays 111160.010121165.01012 Anal-fin spines 3 3 100.0 3 3 3 100.0 3 3 A na l - f i n r ay s 9 9 9 0 . 0 9 10 9 8 5. 0 8 10 Pectoral-fin rays 15 15 60.0 15 16 15 60.0 15 16 Pelvic-fin rays 5 5 100.0 5 5 5 100.0 5 5 Gill rakers on first ceratobranchial 8 9 40.0 7 9 7 40.0 5 8 Gill raker on f irst epibranchial 4 4 90.0 3 4 3 90.0 2 4 Teeth in outer row of left lower jaw 15 15 40.0 11 17 11 30.0 10 17 Teeth rows on upper jaw 4 4 50.0 3 4 3 70.0 2 4 Teeth rows on lower jaw 4 4 100.0 4 4 4 100.0 4 4

150

Apetra simula Holot ype Types Mean Standard Range Deviation min max Standard length, mm 144.0 111.1 16.4 79.0 145.4 Head lengt h, mm 42 .9 3 4.7 5.3 24 .2 4 5.0 Percent o f st and ard leng t h Head length 29.8 31.2 1.0 28.4 32.7 Snount to dorsal-fin origin 34.6 37.0 0.9 34.6 39.0 Snount to pelvic-fin origin 38.1 39.5 1.4 36.8 42.2 Do rsal-f in b ase leng t h 57.3 55.7 1.0 54.1 57.8 Anterior dorsal to anterior anal 51.6 51.2 1.2 48.7 54.3 Posterior dorsal t o posterior anal 17.6 16.8 0.6 15.5 18.5 Anterior dorsal to posterior anal 62.4 60.5 1.1 58.3 62.4 Post erior dorsal t o ant erior anal 33.0 32.1 0.8 30.0 33.7 Post erior d orsal t o vent ral caud al 22 .6 2 0.9 1.0 19.2 2 2.9 Post erior anal t o dorsal caudal 24.8 23.2 0.9 21.5 25.9 Anterior dorsal to pelvic-fin origin 38.5 39.2 1.4 35.8 42.6 Posterior dorsal to pelvic-fin origin 54.2 54.5 1.4 51.2 56.1 Caudal peduncle lengt h 18.7 17.2 1.1 15.2 19.9 Least caudal peduncle dept h 11.9 11.7 0.5 10.4 12.8 Body depth 36.6 36.6 1.4 33.5 39.5 Percent head leng t h Horizont al eye diamet er 30.8 36.0 3.0 30.5 41.9 V ert icle eye diamet er 31.6 35.2 2.5 31.1 41.9 Snout length 41.3 39.0 2.4 33.8 43.4 Postorbital head length 46.9 42.6 2.4 39.0 48.1 Preorbital dept h 26.9 22.9 2.3 17.9 27.0 Lower-jaw length 38.1 38.1 1.9 33.4 44.3 Cheek depth 34.8 30.2 3.1 22.4 36.7 Head depth 106.9 100.3 4.4 88.7 110.2 C ount s M ode %Freq. Range Lat eral-line scales 34 33 47.4 30 34 Po red scales p ost erior t o lat eral line 2 2 57.9 0 3 Scale rows o n cheek 3 3 8 9 .5 2 4 Dorsal-fin spines 16 15 73.7 13 17 Dorsal-fin rays 11 12 50.0 10 13 Anal-fin spines 3 3 100.0 3 3 Anal-fin rays 9 9 68.4 8 10 Pectoral-fin rays 16 16 57.9 14 17 Pelvic-fin rays 5 5 100.0 5 5 Gill rakers on first ceratobranchial 8 9 31.6 6 13 Gill raker on f irst epibranchial 4 3 52.6 3 5 Teeth in outer row of left lower jaw 10 15 28.9 10 17 Teeth rows on upper jaw 4 3 63.2 2 4 Teet h ro ws o n lo wer jaw 5 4 71.1 4 5

151

Apetra perjur Holot ype Types Mean Standard Range Deviation min max Standard length, mm 110.0 79.9 17.8 58.0 123.6 Head length, mm 36.1 25.9 5.6 18.8 38.8 Percent o f st and ard leng t h Head length 32.8 32.5 1.1 30.0 35.2 Snount to dorsal-fin origin 37.1 38.5 1.1 36.1 40.9 Snount to pelvic-fin origin 37.2 39.5 1.6 37.2 43.7 Do rsal-f in b ase leng t h 56 .4 53.9 2.1 50 .3 58.1 Ant erior dorsal to ant erior anal 49.9 47.9 1.9 44.4 51.7 Posterior dorsal to posterior anal 16.1 16.2 0.8 14.4 17.5 Anterior dorsal to posterior anal 59.7 58.2 2.1 55.0 62.9 Post erior dorsal t o ant erior anal 31.4 30.7 1.5 28.5 34.1 Post erior d orsal t o vent ral caud al 20 .7 2 0.7 0 .8 19.1 2 2.3 Post erior anal t o dorsal caudal 22.7 23.0 0.9 21.4 25.1 Anterior dorsal to pelvic-fin origin 38.3 36.5 2.1 32.7 40.4 Posterior dorsal to pelvic-fin origin 55.7 52.2 2.0 49.5 56.7 Caudal peduncle lengt h 17.1 17.6 1.0 15.1 19.7 Least caudal peduncle dept h 11.4 11.6 0.5 10.6 13.2 Body depth 35.9 34.6 1.6 31.5 37.8 Percent head leng t h Horizont al eye diamet er 34.4 39.8 3.1 32.1 44.8 V ert icle eye diamet er 34.2 37.6 2.7 31.0 43.3 Snout length 37.5 35.9 3.0 31.9 45.9 Postorbital head length 41.6 39.9 2.1 35.3 45.2 Preorbital dept h 24.8 20.2 2.8 15.9 25.7 Lower-jaw length 41.7 38.4 2.1 32.3 42.0 Cheek dept h 31.9 25.9 4.1 16.9 37.9 Head depth 101.7 94.0 5.8 84.8 108.0 C ount s M ode %Freq. Range Lat eral-line scales 34 33 52.2 32 35 Pored scales posterior to lateral line 0 2 60.9 0 2 Scale rows o n cheek 3 3 76 .1 2 4 Dorsal-fin spines 15 15 93.5 14 16 Dorsal-fin rays 12 12 69.6 10 13 Anal-fin spines 3 3 100.0 3 3 Anal-fin rays 9 9 80.4 8 10 Pectoral-fin rays 16 16 63.0 14 17 Pelvic-fin rays 5 5 100.0 5 5 Gill rakers on first ceratobranchial 7 8 28.3 6 12 Gill raker on f irst epibranchial 3 4 52.2 3 4 Teeth in outer row of left lower jaw 14 12 34.8 9 15 Teeth rows on upper jaw 3 2 56.5 2 4 Teet h ro ws o n lo wer jaw 4 4 9 5.7 3 5

152

Apetra meniscosteum Holot ype Types Mean Standard Range Deviation min max Standard length, mm 87.6 76.8 9.2 63.6 90.3 Head length, mm 25.6 22.0 2.5 18.7 26.1 Percent o f st and ard leng t h Head length 29.3 28.7 0.9 27.1 30.0 Snount to dorsal-fin origin 34.4 34.5 1.2 32.6 36.9 Snount to pelvic-fin origin 36.3 36.5 0.8 34.7 38.0 Do rsal-f in b ase leng t h 57.3 56.4 1.7 52 .9 59.1 Ant erior dorsal to ant erior anal 47.7 48.4 1.5 45.7 50.9 Posterior dorsal to posterior anal 15.7 15.5 0.6 14.0 16.5 Anterior dorsal to posterior anal 58.9 59.6 2.0 55.2 62.9 Post erior dorsal t o ant erior anal 29.9 29.4 1.1 27.7 31.3 Post erior d orsal t o vent ral caud al 19.7 2 1.1 0 .8 19.5 2 2.7 Post erior anal t o dorsal caudal 24.6 23.6 0.7 22.7 24.8 Anterior dorsal to pelvic-fin origin 34.1 34.4 1.3 32.3 36.6 Posterior dorsal to pelvic-fin origin 52.7 53.4 1.4 50.9 56.1 Caudal peduncle lengt h 19.8 18.9 0.8 17.1 20.4 Least caudal peduncle depth 10.4 10.4 0.5 9.5 11.5 Body depth 32.4 31.9 0.9 30.5 34.1 Percent head leng t h Horizont al eye diamet er 33.6 36.0 2.7 30.7 40.0 V ert icle eye diamet er 31.6 34.3 2.1 30.6 38.1 Snout length 34.7 33.7 2.4 29.2 37.8 Postorbital head length 40.7 41.1 1.5 38.0 43.9 Preorbital dept h 21.6 20.5 1.4 18.3 23.3 Lower-jaw lengt h 39.6 39.9 1.1 38.2 41.7 Cheek dept h 25.9 25.5 1.9 22.9 29.8 Head depth 98.5 95.3 5.8 86.4 107.5 C ount s M ode %Freq. Range Lat eral-line scales 34 34 70.0 33 35 Po red scales p ost erior t o lat eral line 2 2 55.0 0 2 Scale rows o n cheek 2 2 55.0 2 3 Dorsal-fin spines 16 16 65.0 15 17 Dorsal-fin rays 11 11 70.0 10 12 Anal-fin spines 3 3 100.0 3 3 Anal-fin rays 8 9 80.0 8 10 Pectoral-fin rays 15 15 60.0 14 15 Pelvic-fin rays 5 5 100.0 5 5 Gill rakers on first ceratobranchial 11 11 80.0 10 12 Gill raker on f irst epibranchial 4 4 50.0 3 5 Teeth in outer row of left lower jaw 9 10 35.0 7 12 Teeth rows on upper jaw 3 3 90.0 3 4 Teeth rows on lower jaw 4 4 90.0 4 5

153

Apetra cryptopharynx Holotype Kanjedza Island Golden Sands Swamp Songwe Hill Mean StandardRange Mean StandardRange Mean Standard Range Deviation min max Deviation min max Deviation min max Standard length, mm 82.7 68.9 9.5 58.2 88.3 84.6 23.7 58.2 108.0 71.6 8.5 58.7 89.5 Head length, mm 25.5 20.7 3.2 16.6 26.8 27.4 9.6 17.7 37.4 21.9 2.3 17.9 26.5 Percent of standard length Head length 30.9 30.1 0.9 28.0 32.9 31.8 2.6 28.8 35.2 30.7 1.3 27.2 34.8 Snount to dorsal-fin origin 37.3 37.0 1.0 34.5 39.7 37.3 1.3 35.2 39.0 37.3 1.4 33.9 41.6 Snount to pelvic-fin origin 40.2 37.8 1.2 35.5 41.8 41.7 3.9 38.3 47.9 38.1 1.3 35.7 42.6 Dorsal-fin base length 55.4 55.2 1.4 51.2 58.6 53.4 1.6 51.7 56.0 55.9 1.5 51.9 58.3 Anterior dorsal to anterior anal 49.0 49.5 1.3 45.5 53.0 48.9 1.0 47.4 50.4 49.1 1.8 46.0 52.9 Posterior dorsal to posterior anal 16.0 16.5 0.7 14.6 18.6 16.3 1.2 14.6 17.9 16.5 0.6 15.0 18.2 Anterior dorsal to posterior anal 61.1 59.7 1.3 55.5 62.5 57.9 1.4 56.5 60.0 60.0 1.6 55.7 63.4 Posterior dorsal to anterior anal 31.0 30.7 0.9 27.5 32.8 30.4 0.8 29.3 31.3 31.1 1.2 28.4 33.5 Posterior dorsal to ventral caudal 20.4 20.4 0.8 18.7 22.4 20.4 0.7 19.5 21.2 20.3 0.8 17.8 22.4 Posterior anal to dorsal caudal 22.9 23.1 0.9 20.2 25.8 23.3 1.9 20.2 24.8 23.0 0.9 21.0 25.5 Anterior dorsal to pelvic-fin origin 37.8 35.8 1.2 33.0 39.5 37.1 1.6 34.5 38.8 35.8 1.3 33.1 38.7 Posterior dorsal to pelvic-fin origin 53.3 53.2 1.3 49.7 56.3 51.4 1.5 49.6 53.3 52.9 1.7 48.8 56.0 Caudal peduncle length 16.8 17.6 1.1 15.4 20.7 17.5 2.0 14.8 20.0 17.5 1.0 15.5 20.2 Least caudal peduncle depth 11.2 11.3 0.5 9.8 12.6 11.5 0.5 11.0 12.2 11.4 0.5 10.2 12.8 Body depth 34.2 33.7 1.0 31.5 36.3 34.7 1.4 33.1 36.7 33.6 1.1 31.5 36.0 Percent head length Horizontal eye diameter 36.6 40.7 2.2 36.1 44.6 36.7 3.7 32.8 40.8 41.0 2.2 36.8 48.0 Verticle eye diameter 35.4 38.5 2.2 32.8 42.5 35.7 4.7 30.4 40.3 38.2 2.3 34.1 44.9 Snout length 37.5 33.3 2.7 28.5 41.5 35.9 5.4 28.7 42.2 34.1 1.9 29.6 37.4 Postorbital head length 39.3 39.4 1.7 34.9 43.9 38.4 1.7 36.3 41.3 39.7 2.6 28.4 44.2 Preorbital depth 19.3 18.8 1.7 15.7 23.1 18.6 2.0 15.4 21.0 19.5 1.5 16.3 23.4 Lower-jaw length 34.7 37.4 1.7 33.4 40.7 36.4 1.4 33.8 37.4 36.9 1.9 30.8 40.8 Cheek depth 22.9 23.4 1.8 20.0 28.4 23.6 3.8 19.1 28.0 23.4 2.5 19.2 30.2 Head depth 102.6 95.3 3.9 87.2 109.1 96.0 3.3 91.1 99.5 93.4 5.4 84.3 105.8 Counts Mode %Freq.Range Mode %Freq.Range Mode %Freq. Range Lateral-line scales 33 32 48.3 31 34 32 50.0 31 33 33 49.3 31 35 Pored scales posterior to lateral line 1 2 60.7 0 3 2 100.0 2 2 2 46.4 0 3 Scale rows on cheek 3 3 68.5 1 4 3 66.7 2 3 3 66.7 1 3 Dorsal-fin spines 15 16 52.8 13 17 15 66.7 15 16 16 58.0 14 16 Dorsal-fin rays 12 11 44.9 9 12 11 66.7 10 12 11 66.7 10 12 Anal-fin spines 3 3 98.9 3 4 3 100.0 3 3 3 100.0 3 3 Anal-fin rays 10 9 66.3 8 10 9 66.7 8 10 9 53.6 8 10 Pectoral-fin rays 15 14 56.2 13 16 15 50.0 14 15 15 66.7 14 16 Pelvic-fin rays 5 5 100.0 5 5 5 100.0 5 5 5 100.0 5 5 Gill rakers on first ceratobranchial 11 13 28.1 8 15 12 50.0 10 13 11 43.5 9 15 Gill raker on first epibranchial 5 5 56.2 4 6 5 33.3 3 7 5 63.8 3 6 Teeth in outer row of left lower jaw 15 14 25.8 10 17 12 50.0 8 14 14 24.6 9 18 Teeth rows on upper jaw 2 3 51.7 2 4 3 83.3 2 3 2 55.1 2 4 Teeth rows on lower jaw 4 4 97.8 4 5 4 100.0 4 4 4 97.1 4 5

154

Table B.12: TABLE A.11 (concluded)

Apetra cryptopharynx Nkhudzi Bay Otter Point Mean StandardRange Mean Standard Range Deviation min max Deviation min max Standard length, mm 74.0 9.3 60.6 84.7 81.8 5.4 68.4 89.3 Head length, mm 22.7 3.1 18.7 27.3 23.9 1.5 20.7 26.6 Percent of standard length Head length 30.6 1.1 28.5 32.4 29.3 0.9 27.2 30.9 Snount to dorsal-fin origin 36.5 1.1 33.8 38.5 36.1 1.1 32.8 39.2 Snount to pelvic-fin origin 38.8 1.3 37.2 41.3 37.1 0.9 35.0 38.7 Dorsal-fin base length 54.1 1.3 51.7 56.4 56.9 1.3 54.2 60.5 Anterior dorsal to anterior anal 48.6 1.0 47.2 50.3 50.8 1.4 47.6 53.4 Posterior dorsal to posterior anal 16.3 0.6 15.0 17.1 16.7 0.7 14.9 18.0 Anterior dorsal to posterior anal 59.0 1.1 57.0 60.5 61.0 1.2 58.3 63.9 Posterior dorsal to anterior anal 30.5 0.9 29.0 31.7 31.8 1.0 29.8 33.6 Posterior dorsal to ventral caudal 20.9 0.8 19.2 22.4 20.9 0.7 19.2 22.8 Posterior anal to dorsal caudal 23.5 0.8 22.5 25.2 23.4 0.9 21.3 25.1 Anterior dorsal to pelvic-fin origin 36.1 1.4 33.8 37.9 37.4 1.3 34.6 40.6 Posterior dorsal to pelvic-fin origin 52.7 0.9 51.2 54.2 54.6 1.3 52.3 57.9 Caudal peduncle length 18.1 1.0 17.0 19.6 18.2 0.9 16.1 19.7 Least caudal peduncle depth 11.4 0.5 10.3 12.2 11.2 0.4 10.5 12.0 Body depth 33.6 0.9 31.6 34.6 34.7 1.0 32.8 37.7 Percent head length Horizontal eye diameter 40.2 2.4 34.9 44.3 39.8 1.9 34.8 43.4 Verticle eye diameter 37.3 2.1 34.4 41.5 37.8 2.2 32.7 42.4 Snout length 34.8 2.4 32.2 40.4 33.8 1.5 31.2 38.9 Postorbital head length 40.4 1.8 36.6 43.1 40.2 1.5 36.5 42.9 Preorbital depth 19.1 1.4 16.4 21.4 19.9 1.2 16.5 22.2 Lower-jaw length 35.5 2.2 32.1 38.1 38.9 1.6 34.3 41.6 Cheek depth 23.5 2.1 20.9 27.4 25.3 1.4 22.6 28.1 Head depth 95.9 4.5 89.0 102.0 101.1 5.0 88.2 110.8 Counts Mode %Freq.Range Mode %Freq. Range Lateral-line scales 32 60.0 32 34 32 46.3 31 34 Pored scales posterior to lateral line 2 73.3 0 2 2 39.0 0 3 Scale rows on cheek 3 86.7 2 3 3 87.8 2 4 Dorsal-fin spines 15 80.0 15 16 16 68.3 14 17 Dorsal-fin rays 11 66.7 10 12 10 61.0 9 12 Anal-fin spines 3 93.3 2 3 3 100.0 3 3 Anal-fin rays 9 60.0 9 10 9 80.5 8 9 Pectoral-fin rays 15 80.0 14 16 14 75.6 13 15 Pelvic-fin rays 5 100.0 5 5 5 100.0 5 5 Gill rakers on first ceratobranchial 10 46.7 9 12 12 36.6 9 14 Gill raker on first epibranchial 5 46.7 3 5 4 53.7 4 6 Teeth in outer row of left lower jaw 12 33.3 8 15 14 36.6 12 19 Teeth rows on upper jaw 3 53.3 2 4 3 73.2 2 4 Teeth rows on lower jaw 4 86.7 4 5 4 92.7 4 5

155

Apetra retrodens Holotype Chembe Village, Cape Maclear Golden Sands Swamp, Cape Maclear Fisheries Research Station, CM Mean StandardRange Mean StandardRange Mean Standard Range Deviation min max Deviation min max Deviation min max Standard length, mm 102.1 79.1 17.3 53.5 110.9 74.4 18.3 58.1 108.1 78.4 14.8 59.1 104.2 Head length, mm 32.6 25.9 5.8 17.4 37.0 24.9 6.8 19.0 36.1 25.7 5.1 19.1 34.9 Percent of standard length Head length 31.9 32.7 1.1 28.2 35.1 33.3 1.6 31.2 36.3 32.8 1.6 28.0 35.7 Snount to dorsal-fin origin 37.7 38.6 1.4 34.1 42.3 38.3 1.7 36.0 41.5 37.9 1.8 33.0 41.8 Snount to pelvic-fin origin 39.9 40.5 1.4 37.0 44.9 41.1 2.4 38.7 45.8 39.7 1.9 36.3 44.0 Dorsal-fin base length 54.5 52.9 1.8 49.4 57.0 51.9 1.1 50.7 53.5 53.4 1.7 50.0 56.4 Anterior dorsal to anterior anal 50.2 47.4 1.6 43.7 51.5 46.9 0.9 45.4 48.2 46.9 1.1 45.1 48.9 Posterior dorsal to posterior anal 16.1 15.3 0.7 13.8 17.0 16.0 0.9 14.7 17.0 16.2 0.8 14.3 17.6 Anterior dorsal to posterior anal 59.7 57.1 1.5 53.4 60.2 56.3 0.8 55.3 57.7 57.3 1.3 54.4 59.9 Posterior dorsal to anterior anal 30.3 29.3 1.0 27.0 31.7 29.5 0.5 28.3 29.9 30.2 1.1 27.4 32.3 Posterior dorsal to ventral caudal 20.2 20.0 0.9 17.6 22.4 19.9 0.8 18.3 20.7 20.6 0.8 19.3 22.2 Posterior anal to dorsal caudal 22.7 22.2 0.9 20.0 24.4 22.1 0.8 20.5 23.2 22.8 1.1 20.9 24.5 Anterior dorsal to pelvic-fin origin 37.4 35.6 1.6 32.4 39.5 35.2 1.1 33.7 36.9 35.7 1.5 31.9 38.2 Posterior dorsal to pelvic-fin origin 51.2 51.2 1.5 48.1 54.3 50.9 1.6 48.0 53.0 51.6 1.3 48.9 53.9 Caudal peduncle length 16.4 16.5 1.1 14.0 19.2 16.2 0.8 14.0 17.1 17.1 1.1 14.4 19.0 Least caudal peduncle depth 11.2 11.0 0.5 9.7 12.3 11.2 0.4 10.8 11.9 11.3 0.5 10.4 12.5 Body depth 34.6 33.8 1.1 30.9 36.2 33.3 0.9 32.0 35.5 33.1 1.1 30.0 34.9 Percent head length Horizontal eye diameter 36.7 38.8 3.3 33.2 47.3 38.9 3.7 32.4 42.8 38.2 2.6 33.6 42.2 Verticle eye diameter 33.6 36.3 3.0 31.8 45.9 36.0 3.1 31.0 39.9 36.1 2.2 32.6 40.7 Snout length 39.0 37.5 2.9 30.1 43.3 36.5 4.5 32.0 45.1 36.7 2.6 30.5 41.2 Postorbital head length 37.8 38.5 1.7 35.4 42.4 37.6 1.7 34.4 39.7 38.3 2.5 30.2 41.0 Preorbital depth 21.5 19.5 1.9 15.6 24.3 18.5 1.8 15.8 23.0 20.8 2.4 17.4 25.0 Lower-jaw length 39.6 38.7 2.5 31.4 43.8 38.5 1.9 35.2 41.4 39.5 2.0 36.6 44.2 Cheek depth 29.0 25.2 2.6 20.2 30.9 23.0 1.7 20.2 25.8 24.9 3.5 20.3 33.3 Head depth 102.4 91.5 4.9 75.9 102.4 87.0 4.7 79.6 95.0 88.6 4.4 79.9 97.3 Counts Mode %Freq.Range Mode %Freq.Range Mode %Freq. Range Lateral-line scales 33 33 43.6 30 36 32 72.7 31 34 33 29.2 31 34 Pored scales posterior to lateral line 2 2 52.6 0 3 2 63.6 0 3 2 54.2 0 2 Scale rows on cheek 2 3 71.8 2 4 3 63.6 2 3 3 70.8 2 4 Dorsal-fin spines 15 15 80.8 14 17 15 72.7 14 16 15 58.3 14 17 Dorsal-fin rays 11 11 52.6 9 12 11 63.6 11 12 11 45.8 10 13 Anal-fin spines 3 3 100.0 3 3 3 100.0 3 3 3 100.0 3 3 Anal-fin rays 10 9 75.6 8 10 9 72.7 9 10 9 62.5 8 10 Pectoral-fin rays 16 16 61.5 15 17 16 54.5 14 16 16 54.2 14 17 Pelvic-fin rays 5 5 100.0 5 5 5 100.0 5 5 5 100.0 5 5 Gill rakers on first ceratobranchial 10 8 21.8 6 15 10 63.6 10 12 8 33.3 7 14 Gill raker on first epibranchial 2 4 53.8 2 5 5 63.6 4 6 4 50.0 3 6 Teeth in outer row of left lower jaw 12 12 20.5 9 19 12 36.4 10 12 12 20.8 10 17 Teeth rows on upper jaw 3 3 64.1 2 4 2 45.5 2 4 3 62.5 2 4 Teeth rows on lower jaw 4 4 87.2 4 5 4 72.7 4 5 4 95.8 4 5

VITA

MATTHEW R. LISY

100 Doral Farms Road • North Branford, CT 06471 • (203) 257-6787 • [email protected]

EDUCATION

Ph.D. Ecology The Pennsylvania State University, University Park, PA 16802, August 1999-anticipated graduation date December 2006

B.S. Biology, Chemistry Minor Baldwin-Wallace College, Berea, OH 44017-2088, August 1994-May 1998

Alternative Route to Certification (ARC) Department of Higher Education, State of Connecticut, ARC II 2004-2005, Initial Educator Certification in Biology 7-12

TEACHING EXPERIENCE

8/04 –Present Science Teacher, Westhill High School - Stamford Public Schools 125 Roxbury Road, Stamford, CT 06902 Courses Taught: Honors Biology, Biology, Marine Biology, Environmental Science, CP Earth and Space Systems, Earth and Space Systems, Physical Science. Also Ecology/Environmental Club advisor; science fair judge. Received AVID (Advancement Via Individual Determination) training in Austin, TX in June, 2005. Awards and Grants: Winner of Spotlight on Teachers Award for the Stamford Public Schools 2006; Received Stamford Chamber of Commerce Creative Classroom Mini- grants for: Mouse Genetics (2005-2006), Modeling an Ecosystem in a Fish Tank (2004- 2005); Received grant for CAPT Club for science (2004-2005).

9/03-12/03; 8/02-12/02; 1/02-5/02; 8/01-12/01; 1/01-5/01; 1/00-5/00 Teaching Assistant, Ichthyology Laboratory (WFS 453) The Pennsylvania State University, School of Forest Resources Ferguson Building, University Park, PA 16802 Duties: Taught lectures on identification of Pennsylvania fish, natural history; construction of dichotomous key; Constructed and graded quizzes, and papers; Maintained collection of specimens; developed syllabus, teaching methods, and lectures.