The Pennsylvania State University

The Graduate School

College of Agricultural Sciences

TAXONOMIC STUDY OF THE GENUS PETROTILAPIA TREWAVAS 1935

(TELEOSTEI: CICHLIDAE) FROM LAKE MALAŴI, AFRICA

A Dissertation in

Wildlife and Fisheries Science

by

Mary Lundeba

© 2009 Mary Lundeba

Submitted in Partial Fulfillment of the Requirements for the Degree of

Doctor of Philosophy

August 2009 The dissertation of Mary Lundeba was reviewed and approved* by the following:

J.R. Stauffer Jr. Distinguished Professor of Ichthyology Dissertation Advisor Chair of Committee Graduate Program Chair

Cecilia Paola Ferreri Associate Professor of Fisheries Management

Walter M. Tzilkowski Associate Professor of Wildlife Science

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

Adrianus Konings Special Signatory Proprietor of Cichlid Press, Texas

* Signatures are on file in the Graduate School.

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ABSTRACT

Lake Malaŵi is the southernmost of the East African Rift Lakes and harbors about 850 cichlid species of which less than 500 have been scientifically described. One of the genera of cichlid fishes is Petrotilapia, which contains the largest mbuna (rock- dwelling cichlids) of Lake Malaŵi. Petrotilapia species are characterized by broad fleshy lips that are densely covered with slender teeth that are visible even when the mouth is closed. The members of the genus Petrotilapia have been divided into three groups namely the P. tridentiger, the P. genalutea, and the P. nigra group. The members of the

P. tridentiger group are mainly found in the wave-washed upper rocky habitats, while those of the P. genalutea group inhabit the sediment-rich and intermediate habitats. The

P. nigra group contains the largest number of species that inhabit the deeper rocky environments; this group prefers sediment-free rocky habitats. The three Petrotilapia groups are also distinguished by female melanin pattern. The pattern in females of the P. tridentiger group consists of indistinct vertical bars. Females of the P. genalutea group are characterized by two rows of spots on the flank, with the lower row consisting of a few large blotches. Females of P. nigra group often have a yellow or golden background color and a pattern consisting of two horizontal rows of dots of about the same size, with the ones of the mid-lateral row slightly larger. A pattern of, sometimes indistinct, vertical bars is superimposed on the two horizontal rows of dots.

Five hundred and sixty newly collected Petrotilapia specimens were evaluated.

New species of Petrotilapia were diagnosed and described by investigating morphological and meristic differences among populations. Petrotilapia populations were compared on the basis of locality, species complex (Petrotilapia tridentiger,

iii Petrotilapia genalutea, or Petrotilapia nigra group), and markings (presence or absence of a black band in the dorsal fin). Differences in morphology were analyzed using sheared principal component analysis (SPCA) of the morphometric data and principal component analysis of the meristic data. Differences among species were illustrated by plotting the second sheared components of the morphometric data against the first principal component of the meristic data 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, a Duncan Multiple Range Test (DMRT)

(p<0.05) was used to determine which clusters differed from each other. If the clusters were not significantly different along either axis, then a multivariate analysis of variance

(MANOVA), in conjunction with a Hottelling-Lawley trace was used (p<0.05).

Five populations of Petrotilapia were found to be different from all other populations as well as from described species and were described as new species. Newly described species include Petrotilapia xanthos from Gallireya Reef, Petrotilapia mumboensis from Mumbo and Thumbi West islands, Petrotilapia pyroscelos from

Mkanila Bay, Chizumulu Island, Petrotilapia flaviventris from Same Bay, Chizumulu

Island, and Petrotilapia palingnathos from Mkanila Bay, Chizumulu Island; this brings the total to 10 described Petrotilapia species. I determined Petrotilapia ‗ruarwe‘ from

Kakusa and Mbowe Island as conspecifics with Petrotilapia microgalana from Nkhata

Bay. There was no morphological difference between Petrotilapia sp. ‗mumbo yellow‘ from Mumbo Island and P. nigra from neighboring Thumbi West Island. I, therefore, considered P. sp. ‗mumbo yellow‘ a geographical variant of P. nigra. There may be five more undescribed species in the genus Petrotilapia.

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TABLE OF CONTENTS LIST OF FIGURES ...... vi

LIST OF TABLES ...... ix ACKNOWLEDGEMENTS ...... x

Chapter 1 ...... 1 Introduction ...... 1 Study goal and objectives………………………………………………………………4 Cichlid phylogeny, speciation, and diversity…………………………………………...5 Stages of evolutionary cichlid radiation………………………………………………..8 Historical review of Petrotilapia ...... 13 Species concepts and criteria used to delimit species of Petrotilapia………………...16

Chapter 2…………………………………………………………………………………20 Materials and Methods ...... 20

Chapter 3…………………………………………………………………………………29 Description of Petrotilapia ...... 29 Generic description ...... 29 Genus Petrotilapia Trewavas, 1935 ...... 29 Petrotilapia tridentiger Trewavas (1935) ...... 31 Petrotilapia genalutea Marsh (1983) ...... 35 Petrotilapia nigra Marsh (1983) ...... 39 Petrotilapia chrysos Stauffer and Van Snik (1996) ...... 43 Petrotilapia microgalana Ruffing, Lambert, and Stauffer (2006) ...... 47

New species of Petrotilapia ...... 52 Petrotilapia xanthos n. sp. (Fig. 12) ...... 52 Petrotilapia mumboensis n. sp. (Fig. 18) ...... 63 Petrotilapia pyroscelos n. sp. (Fig. 21)...... 71 Petrotilapia flaviventris n. sp. (Fig. 24) ...... 79 Petrotilapia palingnathos n. sp. (Fig. 27) ...... 87 Chapter 4 ...... 94 Discussion and conclusion ...... 94

Chapter 5 ...... 96 Taxonomic key to the Species of Petrotilapia: Species Identification………………...... 96 Key to described species of Petrotilapia ...... 97

Literature Cited ...... 102

v LIST OF FIGURES

Figure 1: Illustration of the three stages of evolutionary vertebrate radiation from Lake Malaŵi, Africa ………………………………………………………….………….10

Figure 2: A member of the genus Petrotilapia from Lake Malaŵi, showing visible teeth while mouth is closed, a key diagnostic feature...... 14

Figure 3: Lake Malaŵi: showing some localities of Petrotilapia species ...... 24

Figure 4: Illustration of some of the morphometric and meristic data points ...... 25

Figure 5: Petrotilapia tridentiger male, Boadzulu Island, Lake Malaŵi, Malaŵi...... 32

Figure 6: Petrotilapia genalutea male, Boadzulu Island, Lake Malaŵi, Malaŵi...... 36

Figure 7: Petrotilapia nigra male, Thumbi West Island, Lake Malaŵi ...... 40

Figure 8: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia sp. ‗mumbo yellow‘ from Mumbo Island (N = 39); Petrotilapia nigra from Thumbi West (N = 14 PSU 4801; Petrotilapia nigra from Monkey Bay (Holotype) BMNH 1981.2.2:206……………………………………………………………………..48

Figure 9: Petrotilapia chrysos male, Chinyamwezi Island, Lake Malaŵi,...... 44

Figure 10: Petrotilapia microgalana male, Lion‘s Cove (T‘hoto), Lake Malaŵi ...... 48

Figure 11: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia microgalana from Nkhata Bay (N = 29) PSU 4757; PSU 4756; PSU 4758; Petrotilapia ‗ruarwe‘ from Kakusa (N = 9); Petrotilapia ‗ruarwe‘ from Mbowe Island (N = 7)………………………...51

Figure 12: Petrotilapia xanthos, HOLOTYPE, PSU 4757, adult male, 130.4 mm SL, Gallireya Reef, Lake Malaŵi, Malaŵi ...... 53

Figure 13: Petrotilapia xanthos: male (a) and female (b) at Gallireya Reef, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4757 (c) and female PSU 4758 (d)...... 56

Figure 14: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758; Petrotilapia chrysos (N = 74) PSU 3453; PSU 4810; PSU 4811; PSU 4812; PSU 4813; PSU 4814; PSU 4815; PSU 4816; PSU 4817; PSU 4818; PSU 4819………………………………………………………………………...57

vi Figure 15: Plot of the second sheared principal components (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758; Petrotilapia tridentiger (N = 64) BMNH 1935.6.14.249.263; PSU 3455; PSU 4805; PSU 4806; PSU 4807; PSU 4808: PSU 4809………………………………………………………………………...... 58

Figure 16: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758; Petrotilapia palingnathos (N = 6) PSU 3467; PSU 4766; Petrotilapia flaviventris (N = 18) PSU 4760; PSU 4759………………………...59

Figure 17: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; and Petrotilapia microgalana (N = 29) PSU 3390; PSU 3453; PSU 4755; BC 003; AMNH 238686………………………………………………………...60

Figure 18: Petrotilapia mumboensis HOLOTYPE, PSU 4765, adult male, 106.1 mm SL, from Mumbo Island, Lake Malaŵi, Malaŵi.……...……………………………..62

Figure 19: Petrotilapia mumboensis: male (a) and female (b) at Mumbo Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4747 (c) and female PSU 4764 (d)……………………………………………………………………67

Figure 20: Plot of the second sheared principal components (morphometric data) and the first factor scores (meristic data) of Petrotilapia mumboensis (N = 18) PSU 4757; PSU 4756; PSU 4758; Petrotilapia tridentiger (N = 64) BMNH 1935.6.14.249.263; PSU 3455; PSU 4805; PSU 4806; PSU 4807; PSU 4808: PSU 4809 ...... 68

Figure 21: Petrotilapia pyroscelos HOLOTYPE, PSU 4753, adult male, 107.2 mm SL, from Mkanila Bay, Chizumulu Isalnd, Lake Malaŵi, Malaŵi...... 72

Figure 22: Petrotilapia pyroscelos: male (a) and female (b) at Mkanila Bay, Chizumulu Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4754 (c) and female PSU 4752 (d)…...………………………………………...... 75

Figure 23: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia pyroscelos from Mkanila Bay and Same Bay, Chizumulu Island (N = 28) PSU 4753; PSU 4752; PSU 4754; Petrotilapia genalutea (N = 3) from Mbweca Rocks, PSU 4750; Petrotilapia genalutea (N = 5) from Yofu Bay, PSU 4749...... 77

Figure 24: Petrotilapia flaviventris, HOLOTYPE, PSU 4760, adult male, 110.9 mm SL, from Same Bay, Chizumulu Island, Lake Malaŵi, Malaŵi...... 80

vii Figure 25: Petrotilapia flaviventris: male (a) and female (b) at Same Bay, Chizumulu Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4759 (c) and female PSU 4759 (d)…………………………………………………...... …………83

Figure 26: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia flaviventris (N = 18) PSU 4760 from Same Bay, Chizumulu Island; PSU 4759 from Same Bay, Chizumulu Island, and Petrotilapia microgalana from Nkhata Bay (N = 40) PSU 3390; PSU3453; PSU 4755; BC 003; AMNH 238686……………………………………...... 84

Figure 27: Petrotilapia palingnathos, HOLOTYPE, PSU 4767, adult male, 118.0 mm SL, from Mkanila Bay, Chizumulu Isalnd, Lake Malaŵi, Malaŵi ...... 88

Figure 28: Petrotilapia palingnathos: male (a) and female (b) at Mkanila Bay, Chizumulu Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4766 (c)...... 90

Figure 29: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia palingnathos form Mkanila Bay (N = 6) PSU 4767; PSU 4766; Petrotilapia tridentiger from Nkhata Bay (N = 7) PSU 3455 and Petrotilapia tridentiger from Nkhata Bay, Boadzulu Island, Crocodile Rock, Mazinzi Reef, Mphanga Rocks, Sanga (N = 64) BMNH 1935.6.14.249.263; PSU 3455; PSU 4805; PSU 4806; PSU 4807; PSU 4808: PSU 4809 ...... 92

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LIST OF TABLES

Table 1: Layout of the morphometric measurements and meristic counts ...... 28

Table 2: Morphometric and meristic values of Petrotilapia tridentiger from Lake Malaŵi (N = 64) ...... 34

Table 3: Morphometric and meristic values of Petrotilapia genalutea from Lake Malaŵi (N = 141) ...... 38

Table 4: Morphometric and meristic values of Petrotilapia nigra from Lake Malaŵi (N = 70) ...... 42

Table 5: Morphometric and meristic values of Petrotilapia chrysos from Lake Malaŵi (N = 74) ...... 46

Table 6: Morphometric and meristic values of Petrotilapia microgalana collected at Nkhata Bay, Lake Malaŵi (N = 29) ...... 50

Table 7: Morphometric and meristic values of Petrotilapia xanthos from Gallireya Reef, Lake Malaŵi (N = 40) PSU 4757; PSU 4756; PSU 4758. Ranges include the holotype...... 62

Table 8: Morphometric and meristic values of Petrotilapia mumboensis, from Mumbo Island, Lake Malaŵi (N = 18) PSU 4765; PSU 4764; PSU 4763, and Petrotilapia mumboensis from Thumbi West, Lake Malaŵi, (N = 21) PSU 4748; PSU 4747; PSU 4746. Ranges include the holotype ...... 70

Table 9: Morphometric and meristic values of Petrotilapia pyroscelos from Mkanila Bay, Chizumulu Island, Lake Malaŵi (N = 21) PSU 4753; PSU 474752, and Petrotilapia pyroscelos from Same Bay, Chizumulu Island, Lake Malaŵi, (N = 7) PSU 4754. Ranges include the.holotype ...... 78

Table 10: Morphometric and meristic values of Petrotilapia flaviventris, from Same Bay, Chizumulu Island, Lake Malaŵi (N = 18) PSU 4760; PSU 4759. Ranges include holotype...... 86

Table 11: Morphometric and meristic values of Petrotilapia palingnathos from Mkanila Bay, Chizumulu Island, Lake Malaŵi (N = 6) PSU 4767; PSU 4766. Ranges inclue the holotype ...... 93

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ACKNOWLEDGEMENTS

I extend my sincere gratitude to Dr. Jay R. Stauffer Jr. who has been extremely supportive, understanding, and professional throughout my study. I must recognize his influence, immense patience, and support without which I would not have reached this level of understanding in the field of ichthyology. The work of Dr. Ad Konings must not go unrecognized! Thank you for being a constant source of insight. I am grateful to

Doctors Paola Ferreri, Walter Tzilkowski, and K.C. Kim for their mentorship along the way; you have contributed to my professional growth profoundly and have shaped me into what I have become today. Academic and support staff who have knowingly or unknowingly assisted me in the development of this work, are thanked. My work would not have been complete without Timothy Stecko‘s technical support, thank you. I thank

Dr. Stauffer‘s ichthyology lab members for their support and friendship. I do extend my appreciation to the British Museum of Natural History for providing me with the type specimens of Petrotilapia, without which this work would not have been complete. Dr.

Stauffer‘s crew in Malaŵi, Africa, is recognized for helping in the collection of the specimens of Petrotilapia that I studied.

I am forever indebted to my family and friends for their support, encouragement, patience, and inspiration, especially to Beauty and her husband, Shupa, who assumed the role of parents for my two lovely sons upon the death of my mother; you are a credit to humanity! I acknowledge Terra and Terry for keeping fond memories of mum who has been half way around the world and for such a long time—mummy is coming home!

x Lastly, I have a sense of indebtedness to the Pennsylvania State University,

School of Forest Resources, and to Dr. Jay Stauffer for the financial support without which this work would not have materialized. Special thanks to the Bowers family and

P.E.O. for the International Peace Scholarship.

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DEDICATION

I dedicate my work to my sons, Terra and Terry; what mom has achieved, you can do it, too. This work is also dedicated to the memory of my beloved mum, Ireen who answered God‘s call one month into this program—memories still fresh!

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Chapter 1

Introduction

The African Great Lakes, lakes Victoria, Tanganyika, and Malaŵi, all harbor a large number of endemic species of cichlid fish (Fryer and Iles, 1972; Coulter, 1991).

The fishes in each lake exhibit remarkable diversity in terms of morphology, ecology, and behavior, and this diversity was acquired through independent and explosive adaptive radiation (Danley and Kocher, 2001). Lake Tanganyika, the oldest of the three lakes, has an estimated age of 9-12 million years (Cohen et al., 1993). The lake supports at least 197 endemic cichlids in 49 endemic cichlid genera (Poll, 1986). The cichlid species in Lake

Tanganyika have been classified into 12 tribes (Poll, 1986; Meyer, 1993; Takahashi et al.,

2001), which are relatively old compared to other East African cichlid lineages and phylogenetic evidence suggests that some of the tribes originated at least five million years ago (Nishida, 1991; Sturmbauer and Meyer, 1993). It has been suggested that Lake

Tanganyika is an evolutionary reservoir of multiple ancient lineages (Nishida, 1991;

Sturmbauer and Meyer, 1993; Nishida, 1997). Lake Tanganyika‘s cichlids are morphologically and behaviorally more diverse than cichlids of lakes Victoria and

Malaŵi (Fryer and Iles, 1972), however, the latter two lakes each posses a greater number of cichlid species than Lake Tanganyika (Danley and Kocher, 2001).

Lake Victoria was formed 250,000-750,000 years ago (Johnson et al., 1996) when the tilting of the Tanzanian shield back-pounded west-flowing rivers; thus, it is not a rift lake and therefore shallower than lakes Tanganyika and Malaŵi, which are situated in the cleft of the East African Rift. In spite of its young age, the lake originally had an

1 extremely rich fish fauna, comprising several hundred species of cichlids (Greenwood,

1974; Kaufman et al., 1997). Until the 1970s, the fish fauna of Lake Victoria was dominated by about 500 endemic haplochromine cichlid species, which comprised about

80% of the demersal fish biomass. The cichlids were extremely diverse ecologically. In the first half of the 1980s, the Nile Perch Lates niloticus, an introduced predator, suddenly boomed and cichlids declined dramatically. During the same period, eutrophication increased strongly, probably because of the extensive predation of the Nile

Perch on the open water (plankton-feeding) cichlids, which fed primarily on planktonic algae. With the decline of the Nile Perch population in the 1990s, the cichlids showed some recovery (Witte et al., 2007). More than 300 endemic cichlid species occur in Lake

Victoria (Seehausen, 1996), all of which were thought to be derived from a single common ancestor (Meyer et al., 1990). Recent molecular data, however, suggest that

Lake Victoria was colonized by at least two separate lineages, one representing rock- dwelling cichlids and the other representing all other endemic cichlids from Lake

Victoria. Each of these lineages invaded the lake and rapidly diverged within the last

12,000 years (Nagl et al., 2000).

Lake Malaŵi, the southernmost of the East African Rift lakes, is the ninth largest lake in the world based on surface area and the third largest based on volume, with a surface area of about 31,000 km2 and a maximum depth of 760 m. The lake is 600 km long, 80 km wide at some locations and is situated at an altitude of 472 m above sea level. Its pH varies between 7.8 and 8.5. The difference between these two values is due mainly to the carbon dioxide (CO2) content of the water. In the surf zone gas exchange is optimal, reducing the CO2 content, and thus making the pH higher than in sheltered bays

2 or deeper layers. The conductivity ranges between 200 and 260 microSiemens, which is relatively low in comparison with the other lakes of the East African Rift Valley. The southeasterly wind (mwera) is most prevalent during the dry season from June to August and induces an upwelling of the colder layers in the most southerly parts of the lake, lowering the surface temperature in that region to 20°C. In the rainy season, November to

April, temperatures in sheltered bays may rise to above 30°C. The average surface temperature, however, ranges between 23-28°C. The Shire River at the southern tip is the only outlet from Lake Malaŵi. The shoreline of the lake falls into three main types, namely rocky shores, sandy beaches, and swampy areas with reeds. Most cichlids occur in particular habitats although none is totally restricted to its preferred environment. The lake is also divided into three rocky habitats, the wave-washed upper rocky, the sediment-free rocky, and the deep, sediment- rich rocky habitats (Konings, 2001, 2007).

No lake in the world contains such a diversification and distinctive community of cichlid fishes as Lake Malaŵi (Konings, 2007). The driving mechanism for the speciation events that led to the explosive radiation of the haplochromine cichlids in the Great Lakes of Africa is undiscovered (Stauffer et al., 2005). The two most widely proposed methods are allopatric speciation (Fryer and Iles, 1972) and intrinsic isolating mechanisms

(McKaye and Stauffer, 1986). Sexual selection has long been proposed as a leading mechanism to the diverse cichlid fauna of Lake Malaŵi (Dominey, 1984; McKaye, 1991;

Turner and Burrows, 1995; Deutsch, 1997; Kellogg et al., 2000). The current haplochromine cichlid species of Lake Malaŵi have radiated from a single ancestor within the last million years (Meyer et al., 1990; Albertson et al., 1999; Danley and

Kocher, 2001); thus, these cichlids demonstrate little genetic diversity (Meyer, 1990).

3 Study goal and objectives

It is widely known that the freshwater fishes of Lake Malaŵi provide an estimated

70% of the total animal protein consumed by Malaŵians. This important resource is now being overexploited. For example, Stauffer et al., (1997a; 2006; 2007) observed an increase in the prevalence of schistosomiasis among village residents and expatriate tourists at Lake Malaŵi over the past decades as a likely result of a decrease in snail- eating cichlid fishes. This decrease permitted an increase in the abundance of snails that are intermediate hosts to schistosomes.

It is estimated that in Lake Malaŵi, there may be as many as 850 cichlid species of which less than 500 have been scientifically described. A combination of morphological, genetic, and behavioral data has been used to diagnose the taxonomic status of these fishes (Trewavas, 1935; Marsh, 1983; Bowers and Stauffer, 1993; Stauffer et al., 1997, 2002b).

Over-harvest of cichlids in Lake Malaŵi calls for corrective management strategies. While human population is projected to continue to increase, better management of the lake is needed to ensure sustainability of this important resource.

Therefore, the majority of species should be identified to devise effective management strategies, while such strategies should also account for the maintenance of species diversity. Determination of the specific status of local taxa is critical for the development of management programs for the conservation and utilization of these fishes for food, tourism, disease control, and scientific investigations (Stauffer et al., 1995; Stauffer and

Kocovsky, 2007). Treating many undescribed species as one large group could result in the loss of species diversity, which could adversely affect the ecosystem as demonstrated

4 by Stauffer et al., (1997a) with the over-fishing of Trematocranus placodon coinciding with a significant increase in the rate of infection with urinary schistosomiasis. Stauffer and Kocovsky (2007) pointed out that the management of sustainable fisheries resources depends on an understanding of the taxonomy and systematics of these fishes.

The research discussed in this study contributes to the accurate delimitation of the taxonomic units (species) of the genus Petrotilapia. The goal was to describe additional species of the genus Petrotilapia. My research objectives were to investigate morphological and meristic differences among populations of Petrotilapia, and to diagnose the species of Petrotilapia. In this study, basic data on the biological and geological attributes that make African cichlid species unique and useful systems in which to study evolution, are reviewed. The stages of vertebrate evolutionary radiation, mechanisms that potentially contribute to the spectacular diversification of cichlids are discussed. The genus Petrotilapia is evaluated and new species of the genus are described.

Cichlid phylogeny, speciation, and diversity

The phylogenetic history of the Lake Malaŵi flock suggests a pattern of radiation in three stages (Albertson et al., 1999; Danley and Kocher, 2001). The first step was the occupation of different macrohabitats, followed by a radiation in feeding morphology and finally diversification of some lineages in male breeding colors, presumably under the action of sexual selection (Albertson et al., 2003).

Cichlid fishes represent an outstanding case of explosive radiation and offer extraordinary opportunities to investigate the evolutionary processes that have led to such

5 diversity (Keenlyside, 1991; Stauffer et al., 1995; Kawanabe et al., 1997). Thus, the study of cichlids offers unique opportunities to gain insight into the reasons underlying taxonomic and geographical distributions of species richness and functional diversity

(Turner et al., 2001). Throughout the world, however, these fishes are threatened by over- fishing, introduction of exotics, habitat destruction, and pollution of the environment

(Stauffer et al., 1995).

The cichlid species of the Great Lakes of Africa provide a compelling model system in which to study the process of speciation. In Lake Malaŵi, at least about 850 cichlid species (Konings, 2007) have emerged since the filling of the lake basin. The level of Lake Malaŵi has fluctuated over a few meters in recent decades. Major recessions occurred in the periods before 25,000 years ago and 10,740 ± 130 years ago, with further large falls between 1150 and 1250 A.D., and within the period 1500-1850

(Owen et al., 1990). The most recent of these recessions in Lake Malaŵi is of major interest in evolutionary studies as it places a time limit on the establishment of the unique fish faunas of many rocky islands and shorelines of the lake (Owen et al., 1990). Based on seismic data and estimates of sedimentation rates, Rosendahl (1988) reported the level of the lake, some 25,000 years ago, as being 250-300 m lower than it is now. In Lake

Malaŵi, the severe lake recessions that occurred must have affected the fish fauna and provided countless opportunities for splitting and recombination of fish populations

(Owen et al., 1990). Such dramatic fluctuations in the lake level would undoubtedly have been reflected in the speciation and extinction of the cichlids.

Over 99% of the Lake Malaŵi haplochromine cichlids are endemic (Fryer & Iles,

1972), suggesting that most of this diversification has taken place within the temporal and

6 spatial boundaries set by the lake‘s shores. Moreover, many of these species are endemic to small areas or islands within the lake (Ribbink et al., 1983), indicating that speciation has occurred very recently (after the last low-water level) (Jordan et al., 2007), or perhaps is in progress. Cichlids are one of the most species-rich families of vertebrates. Although native to tropical areas of the world, with the exception of Australia and Southeast Asia, about 70-80% of the cichlid species are found in Africa, with the greatest diversity found in the Great Lakes (Fryer & Iles, 1972). Evolutionists and ecologists are motivated to understand the forces that generate and maintain biological diversity, and attention has focused on the relative roles of natural and sexual selection in vertebrate groups that are considered to be exemplars of evolutionary radiation.

Cichlids have a number of reproductive adaptations associated with their speciation (Stauffer et al., 2006). Several authors (Trewavas, 1983; Barlow, 2000;

Keenleyside, 1991; Kuwamura, 1986) have reviewed reproductive strategies and parental care in cichlid fishes. Breeding tactics have been separated into substrate and mouthbrooders (Trewavas, 1983; Kuwamura, 1986; Barlow, 1991). African cichlids exhibit a spectacular array of different colors and markings which are normally so species-specific that closely-related species may be distinguished from one another by reference to live colors only (Ribbink, 1990). Most rock-dwelling cichlids of lakes

Malaŵi and Tanganyika defend territories and attract females to spawn in either rock crevices, small caves, on the rock surface, or in the water column above rocks. Males of these fishes presumably rely on their brilliant and diverse color patterns to attract females

(Stauffer et al., 2002). Anatomical and physiological evidence strongly suggests that cichlids have color vision. The presence of several spectral classes of cone

7 photoreceptors in cichlids suggest that color vision plays a role in visually guided foraging behavior and mate selection behavior (Jordan et al., 2006).

Stages of evolutionary cichlid radiation

Recent phylogenetic and population genetic evidence (Albertson et al., 1999;

Danley and Kocher, 2001; Streelman and Danley, 2003) shows that seemingly different vertebrate radiations follow similar evolutionary trajectories. Radiation comprises three stages in which divergent selection drives the diversification of specific phenotypes.

First, lineages diverge along the axis of habitat utilization. Next, secondary morphological specializations related to trophic resource acquisition evolve within habitats, and finally, certain groups diversify along the axis of sensory communication

(Streelman and Danley, 2003). A general example of the three stages of diversification is given in Figure 1.

Divergence of habitat utilization plays an important role in the early stages of vertebrate adaptive radiation (Streelman and Danley, 2003). Among temperate freshwater fishes, the evolution of macrohabitat specialists is pervasive (Robinson and Wilson,

1994). Within sticklebacks, Gasterosteus spp., the divergence of benthic and limnetic morphs has occurred independently in multiple postglacial lakes (Schluter and McPhail,

1993; Rundle et al., 2000). Among lakes, independently derived macrohabitat specialists tend to share similar morphological characters such as body shape, gape width, gill raker length and number (Rundle, 2002). A basal split in macrohabitat is also common in tropical lacustrine fish. The radiation of Lake Malaŵi cichlids is characterized by an early divergence between sand and rock-dwelling lineages (Moran et al., 1994; Danley and

8 Kocher, 2001), each containing greater than 200 species (Streelman and Danley, 2003). A large suite of morphological and behavioral characters can be used to distinguish members of each group including body size and shape, dietary preferences, chromatophore patterning, reproductive behavior, and trophic morphology (Streelman and Danley, 2003).

Secondary divergence in morphological traits as an apparent feature of certain vertebrate radiations is exemplified best by East African cichlids that have evolved numerous trophic adaptations for food acquisition within each of the rock and sand habitats. Within the rock-dwelling clade of Lake Malaŵi, secondary morphological divergence gave rise to proto-genera characterized by differences in feeding behavior and the trophic apparatus (Albertson et al., 1999). There is a broad range of behavioral and morphological variation within the Cichlidae, especially in adaptations related to feeding.

In fact, feeding specializations are believed to be the basis for such prolific speciation

(Keenleyside 1991; Nelson, 1994; Stauffer et al., 2002). Trophic adaptations of the pharyngeal apparatus in particularly contributed to cichlid‘s evolutionary success

(Stiassny and Meyer, 1999).

9 a b c

d e f

g

Figure 1: Illustration of three stages of evolutionary vertebrate radiation from Lake Malaŵi, Africa: a, b, c) Divergence of habitat into sandy and rocky habitats, resulting into divergence of lineages along the axis of habitat utilization; d, e, f) divergence in morphological traits, leading to trophic adaptations for food acquistion; g) divergence in sensory communication involving male breeding color, which plays a role in mate choice by females. As regards stages of cichlid evolutionary radiation, morphologically Petrotilapia species have predominantly tricuspid teeth which are long and slender, adapted generally to browse on rocks. Additionally, Petrotilapia are adapted to rocky habitats of Lake Malaŵi. Underwater observations done by Doctors Stauffer and Konings have also revealed that mate choice by females is based on male breeding colors; females are able to recognize their male conspecifics by color.

10 The cichlids of the Great Lakes of Africa have undergone one of the most rapid radiations of any known vertebrate group (Arratia et al., 2004; Liem, 1974; Meyer et al.,

1990). This rapid speciation rate, however, is not correlated with a high genetic diversity

(Meyer et al., 1990). Conversely, the neotropical cichlids, especially the geophagines, have a significantly higher rate of genetic divergence than the African cichlids (Farias et al., 1999; Zardoya et al., 1996). This higher rate of genetic divergence is not expressed in the cichlids of the Great Lakes of Africa, which is the more diversified group (Stauffer et al., 2006), demonstrating that genetic diversity does not translate into species diversity

Although Malaŵi cichlids demonstrate little genetic diversity (Meyer et al.,

1990), these cichlids exhibit great trophic diversity and plasticity, which has led to great species diversity (Stauffer et al., 2004; 2006). The degree of phenotypic plasticity that cichlids exhibit is congruent with the ability of cichlids to take advantage of many habitats and feeding opportunities (Stauffer et al., 2006). Significant overlap of resource use occurs in cichlid communities containing species with multiple pharyngeal morphologies as a result of the high degree of trophic specialization (Greenwood, 1974), although recent work also demonstrates spatial partitioning of resources among rock- dwelling cichlids (Reinthal, 1990; Stauffer and Posner, 2006). In cichlids, mouth structure, dentition, gill raker number and jaw structure vary tremendously, and this variation in structure has been associated to a variety of feeding techniques (Fryer and

Iles, 1972; Kocher, 2004). The oral jaws have developed specializations for acquiring a variety of food items (Fryer & Iles, 1972). Such specializations include paedophagy

(Stauffer and McKaye, 1986), lepidiophagy (Ribbink, 1984), cleaning (Stauffer, 1991), death feigning (Fryer, 1959), and scraping and raking of an algal, diatomaceous and

11 detrital biolayer from the rock surfaces (Fryer, 1959). Phenotypes such as reverse counter shading associated with such bizarre strategies as hunting upside down have also been documented (Stauffer et al., 1999).

Parrotfish (Scarus, Bolbometopon, Cetoscarus, Hipposcarus, Chlorurus) in the marine reef habitat have diverged into lineages that either scrape or excavate algae from coral and rock surfaces (Bellwood and Chaot, 1990; Bellwood, 1994). By contrast, the seagrass (Cryptotomus, Nicholsina, Leptoscarus, Calotomus, Sparisoma) lineage of parrotfish does not exhibit secondary divergence in morphological features; nearly all species are browsers (Bernardi et al., 2000). These differences in trophic utilization are associated with concomitant changes in the cranial-facial skeleton and musculature

(Bellwood and Choat, 1990; Bellwood, 1994). Phylogenies of cichlids and parrotfish suggest that bursts of signaling evolution, usually involving male secondary sexual characteristics, follow bouts of divergence in habitat utilization and trophic morphology

(Streelman et al., 2002; Albertson et al., 1999).

Cichlids of Lake Malaŵi illustrate the rapid evolution of signaling phenotypes.

Lineages have experienced a diversification of male secondary sexual characteristics.

Within the rock-dwelling clade, this phenomenon has been expressed in terms of male nuptial coloration (Deutsch, 1997). Within the sand-dwelling lineage, extended male phenotypes are affected primarily. Sand-dwelling males build intricate species-specific sand bowers that influence reproductive success (Taylor et al., 1998; McKaye et al.,

1990; Kellogg et al., 2000; Stauffer et al., 2005). Among marine parrotfish a tertiary radiation of signaling phenotypes is also observed. Within the reef clade, species that share similar trophic structures differ in male mating coloration (Streelman et al., 2002).

12 This increased variance in color usage and sexual dimorphism is correlated with differences in behavior such as territoriality and a haremic mating system, which is characteristic of two genera (Chlorurus and Scarus) comprising nearly 80% of all parrotfish species (Streelman et al., 2002; Streelman and Danley, 2003).

Historical review of Petrotilapia

The largest mbuna (rock-dwelling cichlids) are found in the genus Petrotilapia

(Ribbink et al., 1983). Members of the genus Petrotilapia are characterized by broad fleshy lips that are densely covered with slender teeth that are visible even when the mouth is closed (Fig. 4). The numerous teeth are used to comb algae for diatoms and loose algal strands. Because Petrotilapia cannot cope with abundant sediment, they usually comb the partially cleared biocover in territories of other species or that on rocky substrates which are exposed to direct sunlight and therefore support a rich algal matrix

(Konings, 2007).

13

Figure 2: A member of the genus Petrotilapia from Lake Malaŵi, showing visible teeth while mouth is closed, a key diagnostic feature. Photo by Ad Konings.

Stauffer and Posner (2006) established that Petrotilapia species feed from slanted and vertical slopes at feeding angles of 84.5-90.2°. Most species of Petrotilapia inhabit the rocky outcroppings where adult males establish territories. Females, juveniles, and non territorial males are found either singularly or in schools throughout the rocky habitat

(Ribbink et al., 1983). The genus Petrotilapia was originally diagnosed as having all tricuspid teeth (Trewavas, 1935) but in subsequent studies, Marsh (1983) noted some distinctly unicuspid teeth in all the specimens he examined.

The members of the genus Petrotilapia have been divided into three groups

(Konings, 2001) namely the P. tridentiger, the P. genalutea, and the P. nigra group. The members of the P. tridentiger group are mainly found in the wave-washed upper rocky

14 habitats, while those of the P. genalutea group inhabit the sediment-rich and intermediate habitats. The P. nigra group is by far the largest, containing more than a dozen species that inhabit the deeper rocky environments. Members of the latter group prefer sediment- free rocky habitats and this isolates them geographically from the neighboring populations (Konings, 2001, 2007). The three Petrotilapia groups can also be identified by basic melanin pattern which is visible in females and juveniles. The pattern in females of the P. tridentiger group consists of vertical bars. These vertical bars are not, however, always evident on a gray-brown to brown-black ground. Females of the P. genalutea group are characterized by two rows of spots on the flank, with the lower row consisting of a few large blotches. The background color is white or very light beige. Females of the

P. nigra group often have a yellow or golden background color and a pattern consisting of two horizontal rows of dots of about the same size with the ones of the mid-lateral row slightly larger. A pattern of vertical bars is superimposed on the two horizontal rows of spots (Konings, 2001), but such bars may not always be distinct.

The five described species in the genus Petrotilapia include P. tridentiger

Trewavas 1935, P. genalutea Marsh 1983, P. nigra Marsh 1983, P. chrysos Stauffer and

Van Snik 1996, and P. microgalana Ruffing et al., 2006. Besides differences in some morphological characters, the most distinctive difference among these species is live coloration. Ribbink et al. (1983) and Konings (2007) suggest that there are probably 16 additional species that are currently undescribed.

15 Species concepts and criteria used to delimit species of Petrotilapia

Delineation of species in Lake Malaŵi is challenging, in part because of the plethora of species concepts that could be applied to its fauna. With the 22 species concepts listed by Mayden (1997), delimitation of species can be a difficult task (Stauffer et al., 2002b). Discussions about definitions of species have dominated the cichlid literature for many years. 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). Though most work before the 1980s assumed the biological species concept of Mayr (1942), subsequent studies considered alternative definitions. While the biological species concept or ‗isolation concept‘ has been called the most useful species concept for the study of speciation (Coyne and Orr, 1998), alternative approaches have been explicitly championed by a number of cichlid researchers.

Historically, the cichlid fishes of Lake Malaŵi have been diagnosed with morphological data. The use of behavioral data as expressed in mate choice based on color patterns or bower shapes have been successfully used to diagnose cichlid species

(Stauffer et al., 2002b). Marsh (1983) used live coloration, particularly of territorial males as a diagnostic character to distinguish three sympatric species of Petrotilapia. 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).

16 Delimitation of species requires a definition of what is meant by the term species.

I view species as being ontological individuals (Wiley 1978; Mayden 1997, 2002; Wiley

2002; Stauffer et al., 2002a), suggesting that species are real; they form a natural group, exist in nature and cease to exist when they go extinct. Species do not just represent an artificial category constructed for human convenience. For example the Lake Malaŵi cichlids that reside in sympatry demonstrate species-specific breeding. These organisms are able to recognize their conspecifics and therefore, exhibit natural groupings, demonstrating that species are indeed natural. I, therefore, adhere to the evolutionary species concept as my theoretical concept when deciding which populations of

Petrotilapia are species. The evolutionary species concept was originally proposed by

Simpson (1957). Wiley (1978): defined the evolutionary species concept as ―…a single lineage of ancestor-descendant populations, which maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate‖. The evolutionary species concept is a species definition that uses as a unifying framework the idea that all species form a lineage, which is separate from all other lineages. The species is therefore a natural entity on an independent evolutionary trajectory. This lineage concept applies to all reproductive modes. The evolutionary species concept therefore encompasses a wide array of possible species, both extant as well as extinct (Stauffer et al., 2002b). The shortcomings of the evolutionary species concept are that it relies on well-resolved phylogenies and it is non-operational (Mayden, 1997; Stauffer and

McKaye, 2001). Stauffer et al., (2002b) reported that in reality many systems, including

Lake Malaŵi, lack a comprehensive and well-supported phylogeny.

17 Practically, I used some criteria associated with other species concepts to serve as surrogates in supporting my hypothesis of an evolutionary species. The first criterion is the reproductive isolation (associated with the biological species concept) to indicate the independence of evolutionary lineages. Reproductive isolation is assessed through observations of mating activities and analysis of gene flow. The second is the morphological and behavioral similarity criteria to aid in species delineation. This criterion is associated with the morphological and phenetic species concepts. The characters I measured are commonly used in body shape morphology, cichlid pharyngeal morphology, and description of new species (Trewavas, 1935; Barel et al., 1977; Marsh,

1983; Stauffer, 1991; Stauffer and Van Snik, 1996).

Because morphological differentiation takes time, I used shape analysis computer programs to aid in the illustration of the minute differences in the recently radiated populations of Petrotilapia. The sheared principal component analysis (SPCA) attempts to maximize small differences. Differences among species or populations are illustrated by plotting one of the sheared components against the first principal components of the meristic data (Stauffer and Hert, 1992). Although these techniques have greatly improved the resolution of slight morphometric differences (Stauffer et al., 1997), taxonomists are stymied because in many cases, distinct species within the Lake Malaŵi cichlid species flock are morphologically similar (Stauffer et al., 2002b). To attempt to diagnose these similarly-shaped populations, I also used color patterns as a diagnostic character to help me delimit species of the genus Petrotilapia. Several studies have recognized the existence of unique color patterns to be a reliable character for distinguishing cichlid species (Barel et al., 1977; Ribbink et al., 1983; Bowers and Stauffer, 1993; Stauffer et

18 al., 1995, 1997). Most species which are now known differ principally by male coloration and melanin pattern, thus pigmentation remains an important tool for describing cichlid fishes of Lake Malaŵi.

Genetic approaches are often problematic because the Lake Malaŵi cichlids are speciating faster than some diagnostic alleles are fixed within a species (Kornfield et al.,

1985; Kornfield and Parker, 1997), thus not always useful in delimiting the species flock of Lake Malaŵi. The use of starch gel electrophoresis of tissue enzymes to aid in the delineation of Lake Malaŵi cichlid species remains inconclusive (Kornfield, 1978) for similar reasons. And also mitochondrial DNA (mtDNA) sequencing remains problematic in delimiting Lake Malaŵi cichlids because of the rapid radiation which may have prevented mitochondrial sorting of lineages, thereby allowing distantly related species to share mtDNA polymorphisms derived from a common ancestor (Moran and Kornfield,

1993). Owing to these constraints, I have delimited new species using morphology, which includes melanin pattern, markings, and shape analysis.

19

Chapter 2

Materials and Methods

Adult fishes of the genus Petrotilapia were collected from 35 sites in Lake

Malaŵi from 1991 to 2008 (Fig. 3) by Dr. Jay R. Stauffer Jr., Dr. Adrianus Konings, and

Stuart M. Grant. Live fishes were collected by chasing them into a monofilament net while SCUBA diving. Upon collection, the fishes were identified and sorted according to site and whether or not the fish populations were described species or undescribed members of the genus; the fishes were then kept separately in individual glass or plastic jars; anaesthetized in clove oil, and fixed in 10 % formalin. Each jar was labeled with information including fish name, collection date, location, collectors, and number of specimens. In addition, detailed field notes were written by the Stauffer (pers. comm.) for each collection which included color notes on live coloration of the specimens.

Determination of whether the fishes were described species or undescribed members of the genus Petrotilapia was done by the above mentioned collectors who identified the fishes underwater by use of color, morphology and observation of mating pairs. At a later stage the fishes were shipped to the Pennsylvania State University for permanent storage and further examination.

The process of measuring morphometric and counting meristic features of individuals in the collections started with a practice sample of 10 fish that was measured repeatedly to ensure precision. Counts and measures from this practice sample were analyzed using a t-test for evaluation of differences in means. Once results were statistically insignificant at alpha 0.05, measurements were taken as precise. After this, I started collecting my data on both described species and undescribed members of the

20 genus. Data were collected from a total of 560 fish that came from different localities

(Fig. 3). I measured 24 morphometric data points and collected 14 meristic counts on each individual fish. Details of counts and measurements are given in Fig. 4 and Table 1.

Counts and measurements followed Stauffer (1991; 1994). The number of fishes examined in each jar varied and all damaged or deformed fish were excluded from the analysis. The morphometric and meristic data points were collected from the left side of the fish body except for gill-raker meristics, which were taken from the right side. Gill- raker meristics require bending and opening of the operculum and making an incision in the gular membrane; thus to prevent damaging the measured side of the fish, gill-raker counts were done on the right side. Standard length (SL) was used throughout and morphometric values are expressed as percent SL for measurements which extend past the operculum of the fish and percent head length (HL) for measurements which do not extend past the operculum. All morphometric characters were measured in millimeters with digital calipers, between consistent landmarks.

Newly collected Petrotilapia were compared with the following type material:

Petrotilapia tridentiger: 1935.6.14.249.263 British Museum of Natural History (BMNH),

Syntype

Petrotilapia tridentiger: BMNH 1935.6.14.2254.256, Paralectotype

Petrotilapia genalutea: BMNH 1981.2.2.221, Holotype

Petrotilapia genalutea: BMNH 1981.2.2.222-226, Paratypes

Petrotilapia nigra: BMNH 1981.2.2:206, Holotype

Petrotilapia chrysos: PSU 3453, Paratypes

21 Petrotilapia microgalana: AMNH (American Museum of Natural History) 238686

Paratypes; PSU 3390, Paratypes, Bunda College (BC) 003, Paratypes

Comparisons of newly collected Petrotilapia were also done with the following non-type material:

Petrotilapia nigra: 3458 Pennsylvania State University (PSU)

Petrotilapia tridentiger: PSU 3455, PSU 4805, , PSU 4806, , PSU 4807, , PSU 4808, ,

PSU 4809.

Petrotilapia genalutea: PSU 3456, PSU 4749, PSU 4750.

Description of nominal species is based upon Lake Malaŵi populations identified by the author.

Meristic data were analyzed using principal component analysis (PCA) in which the correlation matrix was factored. Morphometric data were analyzed using sheared principal component analysis (SPCA), which factors the covariance matrix and restricts size variation to the first principal component (Humphries et al., 1981; Bookstein et al.,

1985; Stauffer et al., 1997b). Comparisons among species were illustrated by plotting the sheared second principal component of the morphometric data against the first principal component of the meristic data to maximize the amount of separation (Stauffer et al.,

1997b). If the minimum polygon clusters resulting from such plots overlapped, I first determined if the clusters were significantly different along one axis, independent of the another; in such cases, a Duncan‘s Multiple Range Test (DMRT) (p<0.05) was used to determine clusters that differed from each other. If the clusters were not significantly different along one axis independent of the other, then a Multivariate analysis of variance

(MANOVA), in conjunction with a Hottelling-Lawley trace was used to determine

22 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).

I also used basic pigmentation pattern as a diagnostic character to distinguish the species of Petrotilapia where necessary. This was based on field notes and underwater observations on pigmentation pattern of both described species and undescribed members of Petrotilapia, which included both male and female fish. Field notes were provided by

Dr. Jay R. Stauffer, Jr. for each collection analyzed (J.R. Stauffer, Penn State University, personal communication). I used pictures of fishes taken in their natural habitat by

Konings (2001; 2007) and the Malaŵi cichlid feeding behavior video (Konings, 2008) to distinguish species of Petrotilapia based on color pattern.

23 TANZ ANI A

Chitande Isl. Mphanga Roc ks Manda Hara

Gallireya Reef Kakusa Ruarwe Mbowe Island Taiwanee Reef Sanga Lion’s Cove Londo Nkhata Bay Chizumulu Kande Island Island Lake Cobwé Likoma Island Malaŵi MALAŴI Mbweca

Lumessi Mbenji Islands MO ÇAMBIQUE Namalenje Island Nakantenga Island Senga Bay Chinyankwazi Island Mumbo Island Chinyamwezi Island Kadango Reef Thumbi West Island Monkey Bay Mazinzi Reef

Kanchedza Island Boadzulu Isl. Crocodile Rocks

Figure 3: Lake Malaŵi: showing some localities of Petrotilapia species including Luwino Reef and Katale Island, south of Mphanga Rocks; Yofu Bay, east of Chizumulu Island. New species described are from locations in blue. Map of Lake Malaŵi courtesy Ad Konings.

24

Snout to Dorsal Fin Base Vertical Lengt Dorsal Eye h Diameter Standard Length

Horizontal Eye DiameterHead Least Length PeduncleCaudal Snout to Pelvic Fin Depth Dorsal Spines Dorsal Rays

Lateral Line Scale s Pectoral Rays Cheek Scales Anal Upper Tooth Rays Row Lower Tooth Rows Pelvic Anal s Rays Spines

Figure 4: Illustration of some of the morphometric and meristic data points. The measurements that were expressed as percent standard length (SL) were SNDOR,

SNPEL, DFBL, ADAA, ADPA, PDAA, PDPA, PDVC, PADC, ADP2, PDP2, CPL, and

LCPL; while SNL, POHL, HED, VED, HD, PRE, CD, and LJL were expressed as percent head length (HL).

A detailed description of morphometric measurements and meristic counts (Table 1) is given below:

. SL: from snout tip to origin caudal fin

. HL: from snout tip to posterior-most point of operculum

. SNL: from rostral tip upper jaw to anterior orbit margin

. POHL: from posterior orbit margin to posterior-most point of operculum

. HED: horizontal distance between orbit margins

. VED: vertical distance between orbit margins

25 . PRE: length intersection of preorbital bone with line continuing radius of orbit

and parallel to snout profile

. CD: from ventral orbit margin to ventral-most point along ventral borderline of

adductor muscle

. LJL: from rostral tip of dentary to caudal tip of retro-articular process

. HD: perpendicular line from hyoid symphysis to top of head

. BD: from origin dorsal fin to origin pelvic fin

. SNDOR: from snout tip to origin dorsal fin

. SNPEL: from snout tip to origin pelvic fin

. DFBL: from origin to end of dorsal fin

. ADAA: from origin dorsal fin to origin anal fin

. ADPA: from origin dorsal fin to posterior end anal fin

. PDAA: from posterior end dorsal fin to origin anal fin

. PDPA: from posterior end dorsal fin to posterior end anal fin

. PDVC: from posterior end dorsal fin to ventral origin caudal fin

. PADC: from posterior end anal fin to dorsal origin caudal fin

. ADP2: from origin dorsal fin to origin pelvic fin

. PDP2: from posterior end dorsal fin to origin pelvic fin

. CPL: distance between vertical line of posterior end anal fin to that of origin

caudal fin

. LCPD: least vertical depth of caudal peduncle

. Dspines: number of spines of dorsal fin

. Drays: number of soft rays of dorsal fin

26 . Aspines: number of spines of anal fin

. Arays: number of rays of anal fin; if the last ray branches but has same origin, it

was counted as one ray

. P2rays: number of rays of pelvic fin

. P1rays: number of rays of pectoral fin

. LLS: number of scales in upper lateral line plus those of lower lateral line that lie

caudal to last upper lateral line scale

. PSPLL: pored lateral line scales past the hypurals

. CS: number of cheek scales in vertical line from posterior orbit margin

. GRLOW: number of gill rakers on lower arm of anterior-most gill-arch

(ceratobranchial) not including gill raker in angle that separates upper and lower

arm

. GRUP: number of gill rakers on upper arm of anterior-most gill-arch

(epibranchial) not including gill raker in angle that separates upper and lower arm

. TORLLJ: number of teeth in outer row of left lower jaw counting from symphysis

of dentary to end of outer row including developing teeth

. TRU: number of tooth rows in upper jaw

. TRL: number of tooth rows in lower jaw

27 Table 1: Morphometric measurements and meristic counts that were taken on each individual fish Morphometric measurements Meristic counts Standard Length (SL) Doral fin Spines (DSPINES) Head Length (HL) Dorsal fin Rays (DRAYS) Snout Length (SL) Anal fin Spines (ANAL SPINES) Post-Orbital Head Length (POHL) Anal fin Rays (ANAL RAYS) Horizontal Eye Diameter (HED) Pelvic fin Rays (P2RAYS) Vertical Eye Diameter (VED) Pectoral fin Rays (P1RAYS) Pre-Orbital depth (PRE) Lateral Line Scales (LLS) Cheek Depth (CD) Pored Scales Past Lateral Line (PSPLL) Lower Jaw Length (LJL) Cheek Scales (CS) Head Depth (HD) Gill Rakers on ceratobranchial (GRLOW) Body Depth (BD) Gill Rakers on epibranchial (GRUP) Snout to Dorsal fin origin (SNDOR) Teeth Outer Row of Left Lower Jaw (TORLLJ) Snout to Pelvic fin origin (SNPEL) Tooth Rows on Upper jaw (TRU) Dorsal Fin Base Length (DFBL) Tooth Rows on Lower jaw (TRL) Anterior Dorsal fin to Anterior Anal fin (ADAA) Anterior Dorsal fin to Posterior Anal fin (ADPA) Posterior Dorsal fin to Anterior Anal fin (PDAA) Posterior Dorsal fin to Posterior Anal fin (PDPA) Posterior Dorsal fin to Ventral origin Caudal Fin (PDVC) Posterior Anal fin to Dorsal Caudal fin origin (PADC) Anterior Dorsal fin to Pelvic fin origin (ADP2) Posterior Dorsal fin to Pelvic fin origin (PDP2) Caudal Peduncle Length (CPL) Least Caudal Peduncle Depth (LCPD)

28 Chapter 3

Description of Petrotilapia

Generic description

Genus Petrotilapia Trewavas, 1935

Type Species.—Petrotilapia tridentiger Trewavas 1935.

Annual Magazine of Natural History 16:65-118.

Type locality.—Monkey Bay, Lake Malaŵi.

Diagnosis.—The presence of tricuspid teeth in the outer rows of both the lower and upper jaws, and the visibility of some of the inner tooth rows when the jaws are closed distinguishes Petrotilapia from all other mbuna genera. Labeotropheus is the only other mbuna genus with tricuspid teeth in the outer row but upper and lower jaw perfectly fit when the mouth is closed and inner rows of teeth are then not visible. Cynotilapia,

Labidochromis, and Gephyrochromis all have unicuspid teeth in the anterior portion of the outer rows of both jaws. Metriaclima is distinguished by the presence of bicuspid teeth in the anterior portion of the outer row of both the upper and lower jaws. Tropheops is distinguished from Petrotilapia by the presence of one row of bicuspid outer teeth followed by a series of tricuspid inner teeth; while Cyathochromis is distinguished because its teeth consist of slender shafts with compressed, bicuspid crowns.

Description.—Petrotilapia comprises the largest mbuna (rock-dwelling cichlids) of Lake Malaŵi, which are characterized by tiny scales on the breast. The maximum recorded SL in this study is 13.0 cm for P. xanthos.holotype from Gallireya Reef. Some members of the genus such as P. ‗likoma barred‘ and P. palingnathos can attain a

29 maximum total length of 18 and 19 cm respectively (Konings, 2007). The genus

Petrotilapia has many rows of predominantly tricuspid teeth in both lower and upper jaws. Three groups of Petrotilapia have been recognized based on habitat and melanin pattern as described above.

Distribution. —Petrotilapia is distributed throughout the whole lake in rocky habitats.

Etymology. —The name Petrotilapia is from petros (Greek), meaning rock, and

Tilapia, a genus of African cichlids, referring to the feeding habit of members of the genus that generally browse on rocks.

30

Petrotilapia tridentiger Trewavas (1935)

Annual Magazine of Natural History 16:65-118.

Type Specimens

BMNH 1935.6.14.249.263 Syntype BMNH 1935.6.14.2254.256, Paralectotype Non-type material Collectors: Konings, Stauffer, and Grant

PSU 3455, Nkhata Bay; March 1995 PSU 4806 Boadzulu Island ; January 2003

PSU 4805 Crocodile Rock; February 2003

PSU 4808 Mazinzi Reef; January 2003

PSU4809 Mphanga Rocks; January 2007

PSU4807 Sanga; February 2005

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed places this species in Petrotilapia. In live and preserved specimens the dorsal fin has no submarginal band (Marsh, 1993). Absence of the dark submarginal band in the dorsal fin

(Fig. 5) distinguishes it from P. microgalana, P. genalutea, P. nigra, P. chrysos, and from the types of P. mumboensis, and P. pyroscelos which have this band. Adult males of

P. tridentiger are light blue with dark blue bars which distinguishes them from P. chrysos, which are dark blue with black bars and from the types of P. xanthos which are yellow. Adult males of P. tridentiger are also distinct from types of P. flaviventris, and P. palingnathos. Males of P. flaviventris are yellow on ventral and mid sides with scales

31 outlined in blue. Males of P. palingnathos are dark gray with orange markings and scales outlined in blue. Females of P. tridentiger differ from those of P. xanthos, P. chrysos, P. palingnathos, and P. flaviventris. Petrotilapia tridentiger females are brown which distinguishes them from those of P. xanthos which are light brown and from P. chrysos which are golden yellow; females of P. flaviventris are yellow brown and P. palingnathos are orange brown.

Morphometric and meristic data for P. tridentiger non-types are shown in Table 2.

Figure 5: Petrotilapia tridentiger male, Boadzulu Island, Lake Malaŵi, Malaŵi. Photo by A. Konings.

Description.—This species is one of the largest mbuna in Lake Malaŵi, attaining

14.1 cm SL, from Nkhata Bay, and 13.7 cm, from Monkey Bay (Marsh, 1983). The maximum SL of the specimens I examined is 11.0 cm with a maximum body depth of

37.8 % SL. Males from Boadzulu are light blue with dark blue bars and 2 yellow orange ocelli on anal fin. Cheek, gular, breast, and belly purple blue. Dorsal fin with white margin and without black submarginal band; pelvic fin with white leading edge. Females brown with 8 dark brown vertical bars and without horizontal pigmentation elements.

Cheek, gular, breast, and belly orange brown

32 Distribution.—Petrotilapia tridentiger is found at every rocky shore on the western and northeastern side of the lake. It is, however, absent from all islands and from the eastern coast between Cobwe and Makanjila point (Konings, 2007). Specimens examined in this study were collected at Mphanga Rocks, Mazinzi Reef, Sanga,

Boadzulu, Crocodile Rock, Otter Island, and Mbowe Island, all localities in Malaŵi.

Etymology.—The name tridentiger from tri (Latin), meaning three and dentiger from dentis (Latin), meaning tooth to denote the predominantly three-crowned teeth of this species.

33 Table 2: Morphometric and meristic values of Petrotilapia tridentiger types and non- types from Lake Malaŵi, (N = 64).

Variable Mean SD Range Standard length (mm) 110.4 17.8 75.4-163.3 Head length (mm) 36.0 5.6 25.7-53.4 Percent standard length Body depth 37.8 1.5 34.4-41.1 Snout to dorsal-fin origin 34.1 1.2 31.7-36.2 Snout to pelvic-fin origin 41.0 1.9 34.7-44.7 Dorsal-fin base length 61.0 1.6 56.3-64.3 Anterior dorsal to anterior anal 54.4 1.2 52.1-57.6 Anterior dorsal to posterior anal 64.8 1.4 61.6-67.7 Posterior dorsal to anterior anal 31.9 1.4 28.2-34.8 Posterior dorsal to posterior anal 16.3 0.8 13.9-18.5 Posterior dorsal to ventral caudal 18.0 0.9 16.3-20.3 Posterior anal to dorsal caudal 19.0 0.8 16.9-21.8 Anterior dorsal to pelvic fin origin 40.1 1.4 36.7-43.1 Posterior dorsal to pelvic fin origin 57.5 1.7 53.6-61.0 Caudal peduncle depth 13.3 1.1 10.8-15.4 Least caudal peduncle depth 13.7 0.7 12.3-15.4 Percent head length Snout length 40.1 2.0 33.8-44.6 Postorbital head length 38.0 1.4 34.7-40.5 Horizontal eye diameter 28.5 2.2 23.6-33.6 Vertical eye diameter 29.0 2.2 25.2-34.9 Head depth 91.7 5.2 78.2-105.0 Preorbital depth 27.4 2.1 23.2-32.2 Cheek depth 27.0 2.1 22.8-32.0 Lower jaw length 35.0 2.0 31.0-40.9 Counts Mode %Frequency Range Dorsal-fin spines 17 62.5 16-19 Dorsal-fin rays 9 65.6 8-10 Anal-fin spines 3 100 _3_ Anal-fin rays 8 89.1 7-9 Pelvic-fin rays 5 100 _5_ Pectoral-fin rays 14 67.2 10-15 Lateral line scales 30 68.8 29-31 Pored scales past lateral line 2 75.0 0-2 Scale rows on cheek 4 82.8 3-5 Gillrakers on first ceratobranchial 10 60.9 8-13 Gillrakers on first epibranchial 3 82.8 2-3 Teeth on outer row of left lower jaw 14 15.6 10-25 Teeth rows on upper jaw 15 18.8 10-26 Teeth rows on lower jaw 16 18.8 11-24

34 Petrotilapia genalutea Marsh (1983)

J.L.B. Institute of Ichthyology 48:1-14. South Africa.

Type Specimens

Petrotilapia genalutea: BMNH 1981.2.2.221, Holotype

Petrotilapia genalutea: BMNH 1981.2.2.222-226, Paratypes

Non-Type Specimens

Collectors: Stauffer, Konings, and Grant

Specimens examined were collected from 1995-2006. Most specimens prior to the year 2000 were collected by Stauffer and Grant; collections after 2000 were done by

Konings, Stauffer and Grant. Specimens examined include: PSU 4820 (Kanchedza

Island), PSU 4821(Kanchedza Island), PSU 4822 (Namalenje Island), PSU 4823 (Harbor

Island) , PSU 4824 (Mbenji Island) , PSU 4825 (Harbor Island), PSU 4826 (Mazinzi

Reef) , PSU 4827 (Harbor Island) , PSU 4828 (Namalenje Island) , PSU 4829 (Harbor

Island) , PSU 4830 (Mazinzi Reef), PSU 4831 (Mazinzi Reef), PSU 4832 (Mazinzi

Reef), PSU 4833 (Mazinzi Reef), PSU 4834 (Mazinzi Reef), PSU 4835 (Mazinzi Reef),

PSU 4836 (Mazinzi Reef).

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed, places this species in Petrotilapia. In live and preserved specimens, the dorsal fin has a submarginal band. A dark submarginal band in the spinous dorsal fin of both male and female (Fig. 6) distinguishes it from P. tridentiger, P. xanthos, P. flaviventris, and P. palingnathos which lack such a band. Males of P. genalutea differ from those of P. nigra, P. microgalana, P. mumboensis, and P. pyroscelos. Males of P. genalutea are dull

35 gray to blue which distinguishes them from P. nigra and P. chrysos which are predominantly blue-black, and from those of P. microgalana which are bright blue.

Petrotilapia genalutea is further distinguished from males of P. pyroscelos which are blue with bronze highlights and from those of P. mumboensis which have an overall blue color. Females of P. genalutea are off-white to pale yellow brown which distinguishes them from those of P. nigra which are pale brown, from females of P. microgalana which are golden yellow, and from those of P. pyroscelos which are brown with faint blue and yellow highlights.

Morphometric and meristic data for P. genalutea are given in Table 3.

Figure 6: Petrotilapia genalutea male, Boadzulu Island, Lake Malaŵi, Malaŵi. Photo by A. Konings.

Description.—Examined specimens with maximum SL of 10.4 cm; body depth

36.0 % SL. Petrotilapia genalutea shows considerable geographical variation in male coloration Male population from Boadzulu dull gray blue; 5 black vertical bars; yellow cheek and, black gular. Dorsal fin with broad black submarginal band; margin whitish-

36 purple; soft dorsal and caudal rays with yellow leading edge. Anal fin black with 3 yellow ocelli and purple leading edge. Females from Thumbi West off-white to pale yellow brown with horizontal band of large blotches mid-laterally. Cheek, gular, breast, and belly whitish brown. Anal fin with black leading edge; anal soft ray black with 2 orange ocelli.

Distribution.—Petrotilapia genalutea is the most widespread species and occurs lake-wide; specimens examined were collected at Yofu Bay, Mazinzi Reef, and Mitande

Rocks, and at Boadzulu, Mbenji, Harbour, Namalenje, and Kanchedza islands (all locations in Malaŵi, and at Mbweca Rocks and Londo in Mozambique.

Etymology.—The name genalutea from gena (Latin), meaning cheek, and lutea

(Latin) meaning orange in reference to the orange cheeks of adult males.

37 Table 3: Morphometric and meristic values of Petrotilapia genalutea Lake Malaŵi, Africa (N = 141).

Variable Mean SD Range Standard length (mm) 103.5 10.5 73.1-124.3 Head length (mm) 33.2 3.3 22.7-39 Percent standard length Body depth 36.0 1.4 32.1-39.1 Snout to dorsal-fin origin 32.9 1.0 30.5-35.6 Snout to pelvic-fin origin 40.0 1.3 36.9-44.7 Dorsal-fin base length 61.2 1.5 56.5-64.0 Anterior dorsal to anterior anal 54.3 1.5 50.2-57.3 Anterior dorsal to posterior anal 64.8 1.5 60.4-68.1 Posterior dorsal to anterior anal 32.3 1.2 28.5-34.7 Posterior dorsal to posterior anal 16.4 0.8 14.6-18.3 Posterior dorsal to ventral caudal 17.9 0.7 16.4-20.1 Posterior anal to dorsal caudal 19.2 0.7 17.6-20.9 Anterior dorsal to pelvic fin origin 38.6 1.3 35.6-41.3 Posterior dorsal to pelvic fin origin 57.6 1.7 53.0-62.6 Caudal peduncle depth 13.4 0.9 11.0-15.5 Least caudal peduncle depth 13.5 0.5 11.8-14.5 Percent head length Snout length 37.3 2.0 32.2-41.9 Postorbital head length 37.4 1.1 34.8-40.3 Horizontal eye diameter 30.6 1.5 27.5-36.9 Vertical eye diameter 31.2 1.5 28.4-37.3 Head depth 87.2 4.2 76.1-97.4 Preorbital depth 25.9 1.7 20.0-31.4 Cheek depth 25.1 1.7 19.8-29.1 Lower jaw length 35.0 1.6 29.6-40.6 Counts Mode %Frequency Range Dorsal-fin spines 18 78.0 16-19 Dorsal-fin rays 9 79.4 8-10 Anal-fin spines 3 100.0 _3_ Anal-fin rays 8 88.7 6-9 Pelvic-fin rays 5 100.0 _5_ Pectoral-fin rays 14 52.5 12-15 Lateral line scales 30 49.7 29-32 Pored scales past lateral line 2 76.6 0-2 Scale rows on cheek 4 64.5 3-5 Gillrakers on first ceratobranchial 10 47.5 9-12 Gillrakers on first epibranchial 3 81.6 2-4 Teeth on outer row of left lower jaw 14 16.3 10-21 Teeth rows on upper jaw 13 18.4 8-20 Teeth rows on lower jaw 14 19.9 10-21

38 Petrotilapia nigra Marsh (1983)

J.L.B. Institute of Ichthyology 48:1-14.

Type Locality—Monkey Bay, Lake Malaŵi.

Type Specimens

BMNH 1981.2.2:206, Holotype Non-type material Collectors: Konings (PSU 4801), Stauffer, Turner

PSU 3458, Harbor Island; March 1996 PSU 4801 Thumbi West Island; February 2003

PSU 4802 Harbor Island; February 1995

PSU 4803 Mazinzi Reef; January 1991

PSU 4804 Harbor Island; March 1995

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed places this species in Petrotilapia. The dark submarginal band in the spinous part of the dorsal fin of both male and female (Fig. 7) distinguishes it from P. tridentiger, P. xanthos, P. flaviventris, and P. palingnathos which lack such a band. Males of P. nigra are predominantly blue-black which distinguishes them from those of P. genalutea which are dull gray to blue, from those of P. microgalana which are bright blue, from those of P. pyroscelos which are blue with bronze highlights, and from those of P. mumboensis which have an overall blue color. Males of P. nigra are also distinct from those of P. chrysos. Territorial males of P. chrysos are .blue with 7-9 black bars; while non-territorial males are gold with blue highlights. Females of P. nigra are pale brown and are

39 distinguished from those of P. genalutea which are off-white to pale yellow brown, and from those of P. microgalana and P. chrysos which are golden yellow; they are further distinguished from those of P. mumboensis which are gray brown to light brown and from females of P. pyroscelos which are brown with faint blue and yellow highlights.

Morphometric and meristic data for P. nigra are given in Table 4.

Figure 7: Petrotilapia nigra male, Thumbi West Island, Lake Malaŵi, Malaŵi. Photo by A. Konings.

Description.—Examined specimens with mean SL 9.5 mm; body depth 36.8 %

SL. Males predominantly blue-black; dorsal fin with broad, black submarginal band, white margin, and yellow lappets, soft dorsal rays with 2 orange ocelli. Cheek, gular, and pelvic fin black. Females pale brown with two horizontal bands of small spots.

Etymology.—The name nigra from the Latin meaning black to note the dominant dark color of territorial males.

Distribution.—Petrotilapia nigra occurs in the south of the lake around the

Nankumba Peninsula and its associated islands (Konings, 2007). Petrotilapia nigra at

Mumbo Island was previously known as P. sp. ‗mumbo yellow‘. Petrotilapia sp. ‗mumbo yellow‘ belongs to P. nigra group. I compare P. sp. ‗mumbo yellow‘ to the neighboring

40 population of P. nigra from Thumbi West. A comparison of P. sp. ‗mumbo yellow‘ to P. nigra did not revealed a distinct clustering of P. sp. ‗mumbo yellow‘ when the first principal component of the meristic data is plotted against the sheared second principal component of the morphometric data (Fig. 8), suggesting that P. sp. ‗mumbo yellow‘ could be a geographical variant of P. nigra.

0.10 P. 'mumbo yellow' P. nigra

P. nigra

) a

t 0.05

a

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i

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e

m o

h 0.00

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2

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_ -0.05

D

R

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-0.10

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

Figure 8: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia sp. ‗mumbo yellow‘ from Mumbo Island

(N = 39); Petrotilapia nigra from Thumbi West Island (N= 14) PSU 4801; Petrotilapia nigra from Monkey Bay (Holotype) BMNH 1981.2.2:206.

41 Table 4: Morphometric and meristic values of Petrotilapia nigra from Lake Malaŵi, Africa (N = 70).

Variable Mean SD Range Standard length (mm) 94.5 11.8 63.0-116 Head length (mm) 30.2 3.6 20.4-36.6 Percent standard length Body depth 36.8 1.4 33.4-41.0 Snout to dorsal-fin origin 32.6 1.2 30.2-34.9 Snout to pelvic-fin origin 39.5 1.4 35.6-43.7 Dorsal-fin base length 61.3 1.2 58.5-64.0 Anterior dorsal to anterior anal 54.4 1.1 51.7-57.2 Anterior dorsal to posterior anal 64.9 1.1 62.6-67.4 Posterior dorsal to anterior anal 31.9 1.2 29.4-34.9 Posterior dorsal to posterior anal 16.6 1.9 14.8-31.1 Posterior dorsal to ventral caudal 17.4 0.7 15.5-18.9 Posterior anal to dorsal caudal 18.7 0.6 17.2-20.0 Anterior dorsal to pelvic fin origin 38.7 1.1 36.7-42.0 Posterior dorsal to pelvic fin origin 59.6 1.4 55.0-62.2 Caudal peduncle depth 12.8 0.9 10.8-14.8 Least caudal peduncle depth 13.5 0.5 12.2-15.0 Percent head length Snout length 37.2 2.4 30.6-41.8 Postorbital head length 38.2 1.4 34.5-41.2 Horizontal eye diameter 29.9 1.6 26.4-33.8 Vertical eye diameter 30.5 1.6 26.2-34.1 Head depth 86.8 4.1 77.3-97.8 Preorbital depth 26.5 2.3 21.7-34.7 Cheek depth 25.5 1.9 20.2-29.8 Lower jaw length 33.9 2.4 24.7-38.2 Counts Mode %Frequency Range Dorsal-fin spines 18 71.4 17-19 Dorsal-fin rays 9 68.6 8-9 Anal-fin spines 3 100.0 3__ Anal-fin rays 8 81.4 7-9 Pelvic-fin rays 5 100.0 _5_ Pectoral-fin rays 14 54.3 12-15 Lateral line scales 31 52.9 29-32 Pored scales past lateral line 2 75.71 0-2 Scale rows on cheek 3 54.3 3-4 Gillrakers on first ceratobranchial 10 51.4 9-14 Gillrakers on first epibranchial 3 77.4 2-4 Teeth on outer row of left lower jaw 15 18.6 10-21 Teeth rows on upper jaw 12 17.1 10-18 Teeth rows on lower jaw 13 22.9 10-19

42 Petrotilapia chrysos Stauffer and Van Snik (1996)

Copeia 695-702

Type Specimens

PSU 3453 Paratypes Non-type material

Collectors: Stauffer, Bowers, and Grant

PSU 4810, Chinyamwezi Island; March 1991 PSU 4811 Chinyankwazi Island; May 1988

PSU 4812 Chinyamwezi Island; March 1991

PSU 4813 Chnyankwazi Island; March 1991

PSU4814 Chinyamwezi Island, May 1991

PSU4815 Chnyankwazi Island; May 1991

PSU4816 Chnyankwazi Island; May 1991

PSU4817 Chinyamwezi Island, February 1988

PSU4818 Chinyamwezi Island, June 1991

PSU4819 Chinyankwazi Island; May 1988

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed places this species in Petrotilapia. The presence of a dark submarginal band in the dorsal fin

(Fig. 8) distinguishes it from P. xanthos, P. tridentiger, P. palingnathos, and P. flaviventris which lack such a band. Males of P. chrysos are predominantly blue-black which distinguishes them from those of P. genalutea which are dull gray to blue, from

43 those of P. microgalana which are bright blue, from those of P. pyroscelos which are blue with bronze highlights, and from those of P. mumboensis which have an overall blue color. Territorial males of P. chrysos are .blue with 7-9 black bars; while non-territorial males are gold with blue highlights Females of P. chrysos, which lack a black submarginal band in the dorsal, are golden yellow which distinguishes them from those of P. tridentiger which are brown, from those of P. xanthos which are light brown, from those of P. flaviventris which are yellow brown, and from those of P. palingnathos which are orange brown.

Morphometric and meristic data for P. chrysos are given in Table 5.

Figure 9: Petrotilapia chrysos territorial male, Chinyamwezi Island, Lake Malaŵi, Malaŵi. Photo by A. Konings.

Description.—Examined specimens with mean SL 9.7 cm; body depth 37.4 %

SL. Adult males dark blue with black vertical bars; males from Chinyamwezi dark blue with black bars; blue gular and cheek. Females golden brown overall.

44 Distribution.—Petrotilapia chrysos is the only representative of the genus at the two islets, Chinyamwezi and Chinyankwazi, in the southeastern arm of the lake (Ribbink et al., 1983).

Etymology.—The name chrysos, from the Greek, meaning gold in reference of the gold coloration of females.

45 Table 5: Morphometric and meristic values of Petrotilapia chrysos Lake Malaŵi, Africa (N = 74).

Variable Mean SD Range Standard length (mm) 96.9 13.0 60.9-123.8 Head length (mm) 31.7 4.3 19.6-38.6 Percent standard length Body depth 37.4 1.5 33.2-41.8 Snout to dorsal-fin origin 33.4 1.1 30.7-35.8 Snout to pelvic-fin origin 40.4 1.8 34.1-44.5 Dorsal-fin base length 61.6 1.3 58.7-64.1 Anterior dorsal to anterior anal 55.2 1.4 52.5-61.2 Anterior dorsal to posterior anal 65.8 1.3 62.9-69.7 Posterior dorsal to anterior anal 31.8 1.2 29.1-35.7 Posterior dorsal to posterior anal 16.9 0.9 14.5-19.2 Posterior dorsal to ventral caudal 17.5 0.6 16.1-19.3 Posterior anal to dorsal caudal 18.6 0.7 17.2-20.3 Anterior dorsal to pelvic fin origin 39.3 1.2 36.5-43.4 Posterior dorsal to pelvic fin origin 58.5 1.7 54.4-61.8 Caudal peduncle depth 12.1 0.9 10.4-14.5 Least caudal peduncle depth 13.6 0.6 12.3-16.2 Percent head length Snout length 36.9 2.0 32.7-41.6 Postorbital head length 39.1 1.0 36.7-41.1 Horizontal eye diameter 29.7 1.6 25.6-34.4 Vertical eye diameter 30.4 1.6 26.2-34.7 Head depth 89.8 4.5 79.1-101.3 Preorbital depth 26.2 1.7 22.5-30.6 Cheek depth 24.8 2.0 20.1-29.3 Lower jaw length 33.2 2.4 24.6-36.9 Counts Mode %Frequency Range Dorsal-fin spines 18 79.7 17-19 Dorsal-fin rays 9 78.4 8-10 Anal-fin spines 3 100 _3_ Anal-fin rays 8 94.6 7-9 Pelvic-fin rays 5 100 _5_ Pectoral-fin rays 14 82.4 12-14 Lateral line scales 30 62.2 28-30 Pored scales past lateral line 2 78.4 0-2 Scale rows on cheek 3 68.9 3-4 Gillrakers on first ceratobranchial 10 82.4 10-11 Gillrakers on first epibranchial 3 60.8 2-3 Teeth on outer row of left lower jaw 13 20.3 10-21 Teeth rows on upper jaw 13 25.7 8-18 Teeth rows on lower jaw 13 20.3 8-21

46 Petrotilapia microgalana Ruffing, Lambert, and Stauffer (2006)

Biological Society of Washington 119:534-539.

Type Locality.—Nkhata Bay

Collectors: Stauffer and Grant; 1995, 2001

Type Specimens.—PSU 3390, Paratypes; AMNH 238686, Paratypes; BC 003, Paratypes

Non-type material.—PSU 4755, Nkhata Bay, February 2001.

Collectors: Stauffer and Grant

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed places this species in Petrotilapia. The dark submarginal band in the spinous part of the dorsal fin of both male and female (Fig. 9) distinguishes it from P. tridentiger, P. xanthos, P. flaviventris, and P. palingnathos which lack such a band. Adult males of P. microgalana are bright blue which distinguishes them from those of P. genalutea which are dull gray to blue and from those of P. nigra and P. chrysos which are predominantly blue-black; males P. microgalana are distinguished from those of P. mumboensis which have an overall blue color and from those of P. pyroscelos which are blue with bronze highlights.

Females of P. microgalana are golden yellow and lack a black submarginal band in the dorsal fin. They are distinguished from those of P. tridentiger which are brown, from those of P. xanthos which are light brown, from those of P. flaviventris which are yellow brown, and from those of P. palingnathos which are orange brown. Females of P. microgalana are distinguished from those of P. chrysos which are golden yellow and without a horizontal pigmentation pattern.

47 Morphometric and meristic data for P. nigra are given in Table 6.

Figure 10 Petrotilapia microgalana male, Lion‘s Cove (T‘hoto), Lake Malaŵi, Malaŵi.

Photo by A. Konings.

Description.— Examined specimens with mean SL 9.5 cm; body depth 35.3 %

SL. Lateral body coloration of territorial males from Nkhata Bay blue with 5-7 faint black vertical bars. Head blue with one broad black interorbital bar; light blue cheek and yellow gular. Caudal fin with blue membrane, gray rays, and orange tips. Anal fin with two orange ocelli. Females light brown to golden yellow; 5-7 faint brown vertical bars, and horizontal pigmentation pattern.

Distribution. —Petrotilapia microgalana was originally described from Nkhata

Bay, Malaŵi (Fig. 3), but this study has recognized that populations north of Nkhata Bay, at Mbowe Island and Kakusa, previously known as P. ‗ruarwe‘ (Ribbink et al. 1983;

Konings, 2007), are likely conspecific (Fig. 11).

48 Etymology.—The name micro, from the Greek, meaning small, and galanos which, according to Stauffer & Kellog (2002), means blue; in reference to the blue color of territorial males.

49 Table 6: Morphometric and meristic values of Petrotilapia microgalana collected at Nkhata Bay, Lake Malaŵi, Africa, (N = 29) PSU 3390: PSU 3453; PSU 4755; BC 003; AMNH 238686.

Variable Mean SD Range Standard length (mm) 95.0 7.7 78.9-111.9 Head length (mm) 30.6 2.8 25.1-36.7 Percent standard length Body depth 35.3 1.2 32.7-36.9 Snout to dorsal-fin origin 33.7 0.9 31.8-35.5 Snout to pelvic-fin origin 39.9 1.6 35.7-45.7 Dorsal-fin base length 62.2 1.2 59.9-64.3 Anterior dorsal to anterior anal 54.6 1.1 53.0-57.3 Anterior dorsal to posterior anal 65.6 1.2 63.3-67.5 Posterior dorsal to anterior anal 32.2 1.1 30.0-34.5 Posterior dorsal to posterior anal 16.3 0.7 14.8-17.8 Posterior dorsal to ventral caudal 17.8 0.6 16.6-19.1 Posterior anal to dorsal caudal 18.9 0.6 17.7-20.2 Anterior dorsal to pelvic fin origin 37.8 1.3 34.9-40.9 Posterior dorsal to pelvic fin origin 56.8 1.3 53.8-58.9 Caudal peduncle depth 13.1 1.0 11.0-15.0 Least caudal peduncle depth 13.6 0.4 12.7-14.3 Percent head length Snout length 38.1 1.6 33.5-40.5 Postorbital head length 38.1 1.2 35.9-40.3 Horizontal eye diameter 30.2 1.1 27.7-32.8 Vertical eye diameter 31.0 1.2 26.9-32.5 Head depth 88.9 3.2 82.1-96.2 Preorbital depth 26.2 1.4 23.9-28.5 Cheek depth 26.1 1.5 23.6-28.9 Lower jaw length 36.1 1.1 34.4-38.0 Counts Mode %Frequency Range Dorsal-fin spines 18 82.8 17-19 Dorsal-fin rays 9 82.8 9-10 Anal-fin spines 3 100.0 __ Anal-fin rays 8 93.1 7-8 Pelvic-fin rays 5 100.0 __ Pectoral-fin rays 14 62.1 13-14 Lateral line scales 31 68.1 30-31 Pored scales past lateral line 2 68.1 0-2 Scale rows on cheek 4 86.2 3-5 Gillrakers on first ceratobranchial 10 58.6 3-10 Gillrakers on first epibranchial 3 89.7 2-3 Teeth on outer row of left lower jaw 12 27.6 8-17 Teeth rows on upper jaw 11 24.1 7-13 Teeth rows on lower jaw 12 20.7 8-17

50

P. microgalana Nkhata Bay 0.05 Kakusa

) Mbowe Island

a

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o 0.00

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2

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-0.10 -3 -2 -1 0 1 2 PC1 (meristic data)

Figure 11: Plot of the second sheared principal components (morphometric data) and the first factor scores (meristic data) of Petrotilapia microgalana from Nkhata Bay (N = 29) PSU 4757; PSU 4756; PSU 4758; red dot:from Kakusa (N = 9); green dot: from Mbowe Island (N = 7).

51 New species of Petrotilapia

Petrotilapia xanthos n. sp. (Fig. 12) Petrotilapia sp. ‗hara‘ Konings, 2007.

HOLOTYPE.—PSU 4757, adult male, 130.4 mm, S 10°0 29.998' E 34° 14.126'

Gallireya Reef, Lake Malaŵi, Africa; Konings and Stauffer, collectors; January 2007

(Fig. 11).

PARATYPES.—PSU 4756, 24, same collection data as for holotype; PSU 4758,

16, Gallireya Reef, Lake Malaŵi, Africa; June 2008; (74.68-124.45 mm).

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed clearly places this species in the genus Petrotilapia. Absence of a dark submarginal band in the dorsal fin (Fig. 12) distinguishes it from P. microgalana, P. genalutea, P. nigra, P. chrysos, P. mumboensis, and P. pyroscelos which have such a band. Adult males of P. xanthos are yellow, which distinguishes them from P. tridentiger, which are light blue with dark blue bars. Adult males of P. xanthos are also distinct from those of P. flaviventris, and P. palingnathos. Males of P. flaviventris are yellow on ventral and mid sides with scales outlined in blue; the dorsal one-third is blue to gray with yellow and orange highlights; cheeks and gular are yellow to orange. Males of P. palingnathos s are dark gray with orange markings and scales outlined in blue; cheeks are orange with a blue gular. Females of P. xanthos are light brown, which distinguishes them from those of P. tridentiger, which are brown, from P. chrysos and P. microgalana, which are golden yellow, from those of P. flaviventris, which are yellow brown with interrupted black stripes and fading to dark brown dorsally, and from P. palingnathos, which are

52 orange brown. Females of P. xanthos have a melanin pattern that is different from those of P. chrysos, P. microgalana, and P. flaviventris. Females of P. xanthos have both horizontal and vertical melanin pattern, which distinguishes them from those of P. chrysos which lack both horizontal and vertical melanin pattern and from P. microgalana and P. flaviventris which have no vertical melanin pattern.

Caudal peduncle depth is larger in P. xanthos (mean 14.7 % SL; range 13.1-17.0

% SL) than in P. chrysos (mean 12.1 % SL; range 10.4-14.5 % SL), P. tridentiger (mean

13.3 % SL; range 10.8-15.4 % SL), P. flaviventris (mean 12.4 % HL; range 11.2-13.7 %

HL) and in P. microgalana (mean 13.1 % HL; range 11.0-15.0 % HL), but similar to that in P. palingnathos (mean 15.0% HL; range 13.8-17.6 % HL).

Morphometric and meristic data for P. xanthos are shown in Table 7.

Figure 12: Petrotilapia xanthos, HOLOTYPE, PSU 4757, adult male, 130.4 mm SL,

Gallireya Reef, Lake Malaŵi, Malaŵi.

53 Description.—Petrotilapia xanthos is a moderate elongate species; holotype with body depth of 35.6 % SL, smaller than holotypes of P. mumboensis (42.0 % SL), P. flaviventris (38.5 % SL) and P. palingnathos (38.0 % SL), but larger than P. pyroscelos

(34.5 % SL). Greatest body depth of P. xanthos holotype at about base of eighth dorsal spine. Dorsal body profile gradually curving to caudal peduncle; Dorsal head profile concave between snout tip and interorbital, making 45–60° angle with body axis, then rounding to dorsal fin origin; horizontal and vertical eye diameter (29.8 % HL and 30.1

% HL) larger than depth preorbital (25.7% HL). Snout long with isognathous jaws and thickened lips; teeth on lower jaw in 10–17 rows with outer row and inner rows tricuspid.

Dorsal fin with XVII–XVIIII (mode XVIII) spines and 6–10 (mode 9) soft rays. Anal fin with III spines and 7–8 (mode 8) soft rays. First 3-5 dorsal spines gradually increasing in

1 length posteriorly with first spine about /2 length of eighth spine; last 13 spines slightly increasing in length posteriorly with last spine longest; third or fourth dorsal ray longest, past base of caudal fin in males and about to base caudal in females. Anal spines progressively increasing in length posteriorly; third or fourth ray longest, past base of caudal fin, usually shorter than dorsal in males but longer than dorsal in females. Caudal fin subtruncate to emarginate. Pelvic fin not reaching anal fin in females; reaching second to third anal spine in males. Pectoral fin rounded, paddle-shaped, short, reaching vertical through base of 10th or 11th dorsal spine. Flank scales large, ctenoid; abrupt difference to small scales on breast and belly; cheek with 2–5 rows of scales. Lateral line with 29-31 pored scales.

54 Live coloration.—Live males with gold heads and black opercle spot (Fig.13).

Lateral coloration gold with 6-7 faint gray bars. Breast gold with white belly, gold dorsal fin; caudal fin with gold rays and blue membranes. Anal fin blue to gray with 5 yellow ocelli. Anterior pelvic fin dark gold to clear posteriorly. Pelvic fins gold rays and clear membranes. Females with brown heads; white gular, and purple highlights on cheek; black opercle spot with green highlights. Lateral coloration brown with 6 dark brown bars, and brown breast and white belly. Dorsal spines and rays clear, membranes clear with brown spot between spines and black spots between rays. Proximal half of caudal fin brown, distal half light gray and caudal fin membrane with light gray circles.

Proximal three-quarters of anal fin dark brown, distal half clear, anal fin with orange tips and one faint orange ocelli. Pelvic and pectoral fins clear.

Preserved pattern.—Coloration in preserved males consists of yellow to light brown heads, black opercle, and yellow to brown gular; lateral coloration yellow with 7-8 light brown vertical bars and yellow breast and belly. The dorsal, caudal, anal, and pectoral fins and membranes faint yellow. Pelvic fin yellow with yellow to clear membranes. Preserved females with dark to light brown heads, black opercle, and yellow gular. Lateral coloration light brown to yellow with two horizontal dark stripes and 5-7

55 dark bars; breast and belly yellow to whitish (Fig. 13).

a b

c d

Figure 13: Petrotilapia xanthos: male (a) and female (b) at Gallireya Reef, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4757 (c) and female PSU 4758 (d). Distribution.—Petrotilapia xanthos is only known from the type locality,

Gallireya Reef, Malaŵi (Fig. 3).

Etymology.—The name xanthos, from the Greek, meaning yellow referring to the yellow breeding color of males.

Discussion.—Petrotilapia xanthos is probably a member of the P. nigra group and I have compared it to other members of that group (P. chrysos, P. microgalana, and

P. flaviventris) and to other Petrotilapia that lack a submarginal band in the dorsal fin (P. tridentiger and P. palingnathos). Comparison of P. xanthos with P. chrysos, (a member

56 of the P. nigra group), and P. tridentiger, P. flaviventris, and P. palingnathos, males of which lack a dark submarginal band in the dorsal fin, using the principal component analysis revealed distinct clustering of P. xanthos when the first principal component of the meristic data is plotted against the sheared second principal component of the morphometric data (Figs. 14, 15, 16). A comparison of P. xanthos to P. microgalana, which is a neighboring member of the P. nigra group revealed a distinct clustering of P. xanthos when the first principal component of the meristic data is plotted against the sheared second principal component of the morphometric data (Fig. 17).

0.15 P. xanthos P. chrysos

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Figure 14: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758; Petrotilapia chrysos (N = 74) PSU 3453; PSU 4810; PSU 4811; PSU 4812; PSU 4813; PSU 4814; PSU 4815; PSU 4816; PSU 4817; PSU 4818; PSU 4819.

57 P. xanthos 0.2

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Figure 15: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758; Petrotilapia tridentiger (N = 48) BMNH 1935.6.14.249.263; PSU 3455; PSU 4805; PSU 4806; PSU 4807; PSU 4808; PSU 4809.

58 0.15 P. xanthos P. palingnathos P. flaviventris

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Figure 16: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758; Petrotilapia palingnathos (N = 6) PSU 4767; PSU 4766; Petrotilapia flaviventris, (N = 18) PSU 4760; PSU 4759.

59 P. xanthos P. microgalana

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Figure 17: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia xanthos (N = 40) PSU 4757; PSU 4756; PSU 4758 and Petrotilapia microgalana (N = 29) PSU 3390; PSU 3453; PSU 4755; BC 003; AMNH 238686.

In the comparison with P. chrysos, variables with the highest loadings on the sheared second principal component were postorbital head length (-0.23), snout length (-

0.26), and caudal peduncle depth (0.70); while those with the highest loadings on the principal components of the meristic data were teeth on outer left lower jaw (0.46), teeth rows on upper jaw (0.50, and teeth rows on lower jaw (0.51). In the comparison with P. tridentiger, variables with the highest loadings on the sheared second principal component were cheek depth (-0.34), snout length (-0.39), and caudal peduncle length

(0.46); while those with the highest loadings on the principal components of the meristic

60 data were dorsal spines (-0.37), teeth rows on upper jaw (0.50), and teeth rows on lower jaw (0.52). When compared to P. flaviventris and P. palingnathos, variables with the highest loadings were snout length (-0.32), cheek depth (-0.35), and caudal peduncle depth (0.45); while those with the highest loadings on the principal components of the meristic data were teeth on outer left lower jaw (0.43) teeth rows on upper jaw (0.49), and teeth rows on lower jaw (0.52). When P. xanthos was compared to P. microgalana, variables with highest loadings were lower jaw length (0.34), cheek depth (0.43), and caudal peduncle depth (0.47); while those with the highest loadings on the principal components of the meristic data were teeth on outer left lower jaw (0.53) teeth rows on upper jaw (0.53), and teeth rows on lower jaw (0.52).

61 Table7: Morphometric and meristic values of Petrotilapia xanthos, from Gallireya Reef, Malaŵi, (N = 40) PSU 4757; PSU 4756; PSU 4758. Rages include the holotype.

Variable Holotype Mean SD Range Standard length (mm) 130.4 104.2 15.2 74.7-130.4 Head length (mm) 39.0 32.0 4.3 23.6-39.0 Percent standard length Body depth 35.6 35.1 1.5 32.3-39.2 Snout to dorsal-fin origin 31.7 31.7 1.0 29.9-33.8 Snout to pelvic-fin origin 39.8 39.1 1.2 37.1-42.0 Dorsal-fin base length 63.1 60.9 1.3 58.8-63.9 Anterior dorsal to anterior anal 55.0 54.1 1.5 50.5-57.0 Anterior dorsal to posterior anal 64.8 64.2 1.1 62.0-67.2 Posterior dorsal to anterior anal 32.1 32.5 1.1 30.5-34.3 Posterior dorsal to posterior anal 16.5 16.4 0.7 15.3-17.9 Posterior dorsal to ventral caudal 17.8 18.3 0.6 17.1-20.2 Posterior anal to dorsal caudal 18.8 19.4 0.5 18.6-20.9 Anterior dorsal to pelvic fin origin 40.2 38.6 1.6 34.4-42.1 Posterior dorsal to pelvic fin origin 60.9 59.7 1.5 55.7-62.7 Caudal peduncle depth 14.6 14.7 0.8 13.1-17.0 Least caudal peduncle depth 13.5 13.3 0.4 12.7-14.3 Percent head length Snout length 37.8 35.8 1.5 33.4-38.6 Postorbital head length 38.8 37.6 0.9 35.5-38.9 Horizontal eye diameter 29.8 32.6 1.4 29.8-35.5 Vertical eye diameter 30.1 33.0 1.3 30.1-36.9 Head depth 92.5 86.7 5.0 77.0-96.3 Preorbital depth 26.8 25.7 1.4 22.6-28.3 Cheek depth 25.9 24.6 2.1 15.8-28.8 Lower jaw length 34.0 33.7 1.6 29.1-36.7 Counts Mode %Frequency Range Dorsal-fin spines 18 18 72.5 17-19 Dorsal-fin rays 9 9 80 6-10 Anal-fin spines 3 3 100 _3_ Anal-fin rays 9 8 70 7-9 Pelvic-fin rays 5 5 100 _5_ Pectoral-fin rays 14 14 67.5 12-14 Lateral line scales 30 31 50 29-31 Pored scales past lateral line 1 2 62.5 0-3 Scale rows on cheek 4 4 72.5 2-5 Gillrakers on first ceratobranchial 10 11 50 10-12 Gillrakers on first epibranchial 3 3 95 2-3 Teeth on outer row of left lower jaw 19 14 20 10-19 Teeth rows on upper jaw 15 13 30 9-16 Teeth rows on lower jaw 16 14 30 10-17

62 Petrotilapia mumboensis n. sp. (Fig. 18)

Petrotilapia ‗mumbo blue‘ Ribbink et al. 1983.

HOLOTYPE.—PSU 4765, adult male, 106.1 mm; S 13o 59.530' E 034° 45.397'

Mumbo Island, Lake Malaŵi, Africa; Konings, Stauffer, and Grant, collectors; April

2003 (Fig. 18).

PARATYPES.—PSU 4764, 8, Mumbo Island, Lake Malaŵi, Malaŵi, April 2003;

PSU 4763, 9, Mumbo Island, Lake Malaŵi, Malaŵi, February 2003, (78.92-105.69 mm);

PSU 4748, 1, Thumbi West, Lake Malaŵi, Malaŵi; 0-4 m, February 2003 (127.16 mm);

PSU 4747, 16, Thumbi West, Lake Malaŵi, Malaŵi, January 2000; PSU 4746, 4,

Thumbi West, Lake Malaŵi, Malaŵi, February 2003; (80.2-121.2 mm).

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed clearly places this species in the genus Petrotilapia. Its presence in the upper rocky habitat makes it a member of the P. tridentiger group. The dark submarginal band in the spinous part of the dorsal fin of both male and female (Fig. 18) distinguishes it from P. tridentiger, P. xanthos P. flaviventris, and P. palingnathos which lack such a band. Males of P. mumboensis differ from those of P. genalutea, P. nigra, P. chrysos, P. microgalana and P. pyroscelos by its light blue overall color with 8 dark-blue vertical bars, light-blue cheeks, and a light blue to gray gular region. Males of P. genalutea are dull gray-blue with 5-7 black vertical bars, have an orange-yellow cheek, and a black gular. Males of P. nigra and P. chrysos are predominantly blue-black with 7-10 gray/brown bars, have a dark-blue cheek, and a black gular. Males of P. microgalana are bright blue with 5-7

63 faint black vertical bars, and have a light-blue cheek and a yellow gular while those of P. pyroscelos are blue with bronze highlights and have a red pelvic fin. Females of P. mumboensis are gray-brown to light brown with a conspicuous black submarginal band in the dorsal fin and are distinguished from those of P. genalutea, P. nigra, P. chrysos, and

P. microgalana by the lack of a horizontal pigmentation pattern and are distinguished from females of P. pyroscelos which are brown with faint blue and yellow highlights.

Morphometric and meristic data for P. mumboensis are shown in Table 8.

Figure 18: Petrotilapia mumboensis HOLOTYPE, PSU 4765, adult male, 106.1 mm SL, from Mumbo Island, Lake Malaŵi, Malaŵi.

Description.—Petrotilapia mumboensis holotype has the deepest body in comparison to other Petrotilapia type species; BD (42.0 % SL), P. xanthos (35. 6 % SL),

P. pyroscelos (34.5 % SL), P. flaviventris (38.5 % SL), and P. palingnathos (38.0 % SL).

Petrotilapia mumboensis has a mean BD of 39.2% SL. Greatest body depth at about base of ninth spine. Dorsal body profile gradually curving to caudal peduncle. Dorsal head profile concave between snout tip and interorbital, making 45–55° angle with body axis,

64 then rounding to dorsal fin origin; horizontal and vertical eye diameter (26.7 % HL and

27.5 % HL) larger than depth preorbital (mean 23.7 % HL). Snout with isognathous jaws and thickened lips; teeth on lower jaw in 10–17 rows with both outer row and inner rows tricuspid, except for a few unicuspid teeth at the posterior end of the rows.

Dorsal fin with XVII–XVIII (mode XVII) spines and 9–10 (mode 9) soft rays.

Anal fin with III spines and 7–9 (mode 8) soft rays. First 3-6 dorsal spines gradually

1 increasing in length posteriorly with first spine less than /2 length of ninth spine; last 13 spines slightly increasing in length posteriorly with seventeenth spine longest; third or fourth ray longest, past base of caudal fin. Anal spines progressively increasing in length posteriorly; fourth ray longest, in some males past base of caudal fin while in others reaching to about base of caudal fin, slightly shorter than dorsal in males. Caudal fin subtruncate to emarginate. Pelvic fin not reaching anal fin in some females; reaching first to second anal spine in males. Pectoral fin rounded, paddle-shaped, short, reaching vertical through base of 10th dorsal spine. Flank scales large, ctenoid; cheek with 3–5 rows of scales. Lateral line with 29-31 pored scales.

Live coloration.—Males with dark blue/gray heads and two light blue interorbital bars, opercle and pre-opercle with light blue highlights and a black opercle spot; gular light/blue gray. Lateral coloration blue/green with 6 black bars; breast and belly white with black/gray markings. Caudal peduncle dark blue/gray. Dorsal fin with broad black submarginal band. Proximal one-quarter of the dorsal fin light blue, lappets light blue with orange tips. Caudal fin with gray rays, orange tips, and light blue membrane. Anal fin blue/gray with a black spinous portion which extend as abroad black marginal bar over rayed portion; 2-6 dark orange ocelli on anal fin. Pelvic fin light blue with white

65 leading edge; first two rays of fin and membrane black, rest of the membrane clear.

Pectoral fin with gray rays and clear membrane. Females with brown head and gray gular. Lateral coloration brown with light-brown to white belly and dark gray breast.

Anal fin black with 1-4 small orange ocelli. Pelvic, pectoral, caudal, and dorsal fins as in males.

Preserved pattern.—Coloration in preserved males consists of dark-brown head, brown opercle, and brown gular. Lateral coloration dark brown to gray with 7-8 faint dark vertical bars, white to gray belly, and dark brown breast. Dorsal fin light brown to dark, with broad submarginal black band that extends into rays; dorsal fin membrane light brown. Caudal fin brown to gray with light-brown membrane. Anal fin gray with black marginal band that extends to first ray; spines black with dark-brown rays and light-brown membrane. Pelvic fin light brown with black spine; light-brown membrane and black marginal band that extends to first ray. Pectoral fin light brown with clear membrane. Females with dark-brown head, brown opercle, and yellow to light-brown gular. Lateral coloration brown with 8-9 vertical bars, yellow to light brown breast, and light brown belly. Dorsal fin light brown with broad submarginal band and light-brown membrane. Caudal fin brown with light-brown membrane. Anal fin light brown to gray, light-brown membrane, and black spines with submarginal band that extends to anal rays.

Pelvic fin gray to white; black spine with first three rays dark and last two rays white to gray. Pectoral fin dark to gray with clear membrane. Figure 19 below shows live and preserved specimens of P. mumboensis.

66 a b

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Figure 19: Petrotilapia mumboensis: male (a) and female (b) at Mumbo Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4747 (c) and female PSU 4764 (d).

Distribution.—Petrotilapia mumboensis occurs at Mumbo and Thumbi West islands, Malaŵi. Ribbink et al. (1983) report this species‘ presence at Mbenji Island as well.

Etymology.—The name mumboensis is in reference to Mumbo Island, Malaŵi where this species is very common and from where the holotype was collected.

Discussion.—Petrotilapia mumboensis is a member of the P. tridentiger group but is easily distinguished from P. tridentiger by the black submarginal band in the dorsal.

Petrotilapia mumboensis is sympatric with P. genalutea and P. nigra. The form of the latter species at Mumbo Island was previously known as P. sp. ‗mumbo yellow‘, but as I

67 demonstrated on page 41, I was unable to find any morphological difference between the

Mumbo population and those of the neighboring rocky areas, i.e. Thumbi West Island. A plot of the sheared second principal component of the morphometric data versus the first principal component of the meristic data show that the minimum polygon cluster formed by P. mumboensis does not overlap with the cluster formed by P. tridentiger (Fig. 20).

0.15 P. mumboensis P. tridentiger

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Figure 20: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia mumboensis (N = 39) PSU 4765; PSU 4764; PSU 4763; PSU 4748: PSU 4747: PSU 4746 and Petrotilapia tridentiger (N = 64) BMNH 1935.6.14.249.263; PSU 3455; PSU 4805; PSU 4806; PSU 4807; PSU 4808; PSU 4809.

Variables with the highest loadings on the sheared second principal component were preorbital head length (-0.61), caudal peduncle depth (-0.49), and least caudal

68 peduncle length (0.26); while those with the highest loadings on the principal components of the meristic data were teeth rows on upper jaw (0.45), lateral line scales

(0.45), and teeth rows on lower jaw (0.55).

69 Table 8: Morphometric and meristic values of Petrotilapia mumboensis, from Mumbo Island, Lake Malaŵi, Africa, (N = 18) PSU 4765; PSU 4764; PSU 4763, and Petrotilapia mumboensis from Thumbi West, Lake Malaŵi, Africa (N = 21) PSU 4748; PSU 4747; PSU 4746. Ranges include the holotype.

Variable Holotype Mean SD Range Standard length (mm) 106.1 100.9 12.5 78.9-127.2 Head length (mm) 36.9 34.0 3.6 27.6-40.0 Percent standard length Body depth 42.0 39.2 1.4 36.3-42.5 Snout to dorsal-fin origin 35.9 35.0 1.1 32.5-36.9 Snout to pelvic-fin origin 40.7 40.1 1.9 33.6-44.3 Dorsal-fin base length 63.6 61.8 1.5 59.5-64.9 Anterior dorsal to anterior anal 58.0 54.9 4.2 30.3-58.4 Anterior dorsal to posterior anal 67.9 66.2 1.4 64.1-69.4 Posterior dorsal to anterior anal 34.9 32.9 1.3 30.3-35.1 Posterior dorsal to posterior anal 17.0 16.9 0.7 15.4-18.3 Posterior dorsal to ventral caudal 18.0 18.2 0.6 17.0-19.4 Posterior anal to dorsal caudal 18.7 18.9 0.6 17.4-20.0 Anterior dorsal to pelvic fin origin 44.1 41.1 1.3 38.3-44.1 Posterior dorsal to pelvic fin origin 61.6 59.4 1.4 57.1-62.7 Caudal peduncle depth 12.2 12.3 0.9 10.0-14.1 Least caudal peduncle depth 15.5 14.6 0.6 13.6-15.7 Percent head length Snout length 37.0 38.7 2.4 32.3-42.8 Postorbital head length 39.5 39.9 1.3 36.5-43.0 Horizontal eye diameter 26.7 27.3 1.6 23.4-30.2 Vertical eye diameter 27.5 28.1 1.6 24.7-31.5 Head depth 94.0 92.2 6.0 80.8-111.7 Preorbital depth 23.7 24.7 2.6 20.0-28.5 Cheek depth 27.4 27.2 2.3 22.4-33.0 Lower jaw length 37.2 34.9 1.7 30.6-37.4 Counts Mode %Frequency Range Dorsal-fin spines 18 17 84.6 16-18 Dorsal-fin rays 8 9 79.5 8-10 Anal-fin spines 3 3 100 _3_ Anal-fin rays 8 8 89.7 7-9 Pelvic-fin rays 5 5 100 _5_ Pectoral-fin rays 14 14 82.1 13-15 Lateral line scales 29 29 64.1 29-31 Pored scales past lateral line 1 2 82.1 1-2 Scale rows on cheek 4 4 74.4 3-5 Gillrakers on first ceratobranchial 10 10 71.8 9-12 Gillrakers on first epibranchial 3 3 69.2 2-3 Teeth on outer row of left lower jaw 19 16 18.0 10-27 Teeth rows on upper jaw 21 13 23.1 9-21 Teeth rows on lower jaw 9 14 18.0 9-21

70

Petrotilapia pyroscelos n. sp. (Fig. 21)

Petrotilapia sp. ‗orange pelvic‘ Ribbink et al. 1983.

HOLOTYPE.—PSU 4753, adult male 107.2 mm; S 12o 00.566' E 34° 36.882'

Mkanila Bay, Chizumulu Island, Malaŵi, Africa (Fig. 19); Grant, Konings, Stauffer, collectors; January 2003 (Fig. 21).

PARATYPES.—PSU 4752, 21, (71.9-103.56 mm) same data as for holotype;

PSU 4754, 7, Same Bay, Chizumulu Island, Malaŵi; Grant, Konings, Stauffer, collectors;

February 2006, (77.23-100.79 mm).

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed clearly places this species in the genus Petrotilapia. The dark submarginal band in the spinous part of the dorsal fin of both male and female (Fig. 21) distinguishes it from P. tridentiger, P. xanthos, P. flaviventris, and P. palingnathos which lack such a band.

Males of P. pyroscelos differ from those of P. genalutea, P. nigra, P. chrysos, P. microgalana and P. mumboensis by its blue and bronze ground color with 7-8 gray vertical bars, whitish to gray gular and purple cheek. Males of P. genalutea are dull gray- blue with 5-7 black vertical bars, have an orange-yellow cheek, and a black gular. Males of P. nigra and P. chrysos are predominantly blue-black with 7-10 gray/brown bars, have a dark-blue cheek, and a black gular. Males of P. microgalana are bright blue with 5-7 faint black vertical bars, and have a light-blue cheek and a yellow gular while those of P. mumboensis are blue with 8 dark-blue vertical bars, light-blue cheeks, and a light-blue to gray gular region. Females of P. pyroscelos are brown with faint blue and yellow

71 highlights with a conspicuous black submarginal band in the dorsal fin. Females of P. pyroscelos are distinguished from those of P. genalutea, P. nigra, P. chrysos, and P. microgalana by the lack of horizontal stripes or spots in the melanin pattern and are distinguished from females of P. mumboensis which are gray-brown to light brown.

Morphometric and meristic data for P. pyroscelos are shown in Table 9.

Figure 21: Petrotilapia pyroscelos HOLOTYPE, PSU 4753, adult male, 107.2 mm SL, from Mkanila Bay, Chizumulu Island, Lake Malaŵi, Malaŵi.

Description.—Petrotilapia pyroscelos holotype has the smallest body depth (34.5

% SL) compared P. mumboensis, P. xanthos, P. flaviventris, and P. palingnathos; see values above.on page 64. Greatest body depth at about base of tenth dorsal spine. Dorsal body profile gradually curving to caudal peduncle; Dorsal head profile concave between snout tip and interorbital, making 30–50° angle with body axis, then rounding to dorsal fin origin; horizontal and vertical eye diameter (31.5 and 32.6 % HL) larger than depth preorbital 28.2 % HL). Snout with isognathous jaws and somewhat thickened lips; teeth on lower jaw in 10–18 rows with both outer row and inner rows tricuspid, except for posterior end rows which are unicuspid.

72 Dorsal fin with XVII–XVIIII (mode XVIII) spines and 8–10 (mode 9) soft rays.

Anal fin with III spines and 7–8 (mode 8) soft rays. First 4-5 dorsal spines gradually

1 increasing in length posteriorly with first spine less than /2 length of tenth spine; last 14 spines slightly increasing in length posteriorly with last spine longest; soft dorsal with subacuminate tip, fourth or fifth ray longest, past base of caudal fin in males, about reaching base of caudal fin in females. Anal spines progressively increasing in length posteriorly; fourth or fifth ray longest, reaching to about base of caudal fin, slightly shorter than dorsal in males. Caudal fin subtruncate to emarginate. Pelvic fin not or about reaching anal fin in females; reaching first to second anal spine in males, and in some males, reaching first soft ray. Pectoral fin rounded, paddle-shaped, short, reaching vertical through base of 10th or 11th dorsal spine. Flank scales large, ctenoid; cheek with

3–5 rows of scales. Lateral line with 29-33 pored scales.

Live coloration.—Males with dark gray interorbital, no interobital bar; cheek, preorbital, opercle, and pre-opercle purple with light blue highlights and a dark gray opercle spot; gular whitish to gray. Lateral coloration blue with bronze highlights and 5 dark gray to blue vertical bars; breast and belly whitish to gray. Dorsal fin with broad black submarginal band and light blue lappets. Proximal one-third to half of dorsal fin blue, and membrane between ray blue. Caudal fin with gray rays and blue membrane.

Anal spines and membrane black, rays black to gray and with clear membrane and 2 yellow ocelli. Pelvic fin orange with white leading edge. Pectoral fin withdark blue rays and clear membrane. Females with brown heads with blue and green highlights on cheek, black opercle spot and white gular. and gray gular. Lateral coloration brown with faint blue and yellow highlights; scales outlined in blue; pale yellow breast with white belly.

73 Anal fin with gray rays, clear membrane and 3 yellow ocelli. Dorsal fin with brown spines and rays; black submarginal band, yellow and light blue lappets with brown tips.

Caudal fin with gray rays and clear membrane. Pelvic fin with white to light blue leading edge. Pectoral fin with brown rays and clear membrane.

Preserved pattern.—Males consists of dark brown head, black opercle, brown cheek, and yellowish to light brown gular. Lateral coloration dark brown; faint dark vertical bars, white belly, and brown breast. Dorsal fin with a black submarginal band, gray membrane, and gray to brown spines and rays. Caudal fin with dark to gray rays and gray membrane. Anal spines and membrane black, rays black to gray with clear membrane. Pelvic spine and first three rays gray with a clear membrane, posterior rays and membrane clear. Pectoral fin with gray rays and clear membrane. Preserved females with brown head, black opercle, brown cheek, and a yellowish to brown gular. Lateral coloration brown with blotches and faint vertical bars; breast yellow, belly white to yellow. Dorsal fin with black submarginal band, spines and membrane gray; rays with brown spots. Caudal fin with brown to gray rays and clear membrane. Anal fin with gray to brown spines with brown to clear membrane. Pelvic spines and first two rays dark with dark to clear membrane. Pectoral fin with gray rays and clear membrane. Figure 22 below shows live and preserved coloration of P. pyroscelos.

74 a b

c d

Figure 22: Petrotilapia pyroscelos: male (a) and female ( b) at Mkanila Bay, Chizumulu Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4754 (c) and female PSU 4752 (d)

Distribution.—Petrotilapia pyroscelos is only known from the type locality,

Chizumulu Island, Malaŵi (Fig. 3).

Etymology.—The name pyroscelos is from pyros (Greek), meaning fire and skelos

(Greek), meaning leg in reference to the pelvic fins of this species that have the color of fire.

Discussion.—Petrotilapia pyroscelos is probably a member of the P. genalutea group and is restricted to Chizumulu Island where it is commonly encountered in the shallow wave-washed rocky areas (Konings, 2007). Petrotilapia genalutea is absent from

Chizumulu Island. Territorial male Petrotilapia pyroscelos are easily distinguished from those of all other species in Petrotilapia by its orange pelvic (Fig. 20). Comparison with

P. genalutea from nearby populations, Mbweca Rocks and Yofu Bay, indicated separation of the two species. The plot of the sheared second principal component of the

75 morphometric data versus the first principal component of the meristic data show that the minimum polygon cluster formed by P. pyroscelos does not overlap with the clusters formed by P. genalutea (Fig. 23). Snout length is about the same in P. pyroscelos (mean

36.3% HL; range 33.5-39.9% HL) and P. genalutea from Mbweca Rocks (mean 36.2%

HL; range 34.6-37.6% HL) but smaller in P. genalutea from Yofu Bay (mean 33.0 %

HL; range 32.2-34.4 % HL). Both preorbital depth and lower jaw length are larger in P. pyroscelos (mean 25.3% HL; range 22.2-28.6 % HL; mean 35.7; range 29.2-38.6 % HL) than in P. genalutea from Mbweca Rocks (mean 24.8 % HL; range 23.8-25.7 % HL; mean 34.8 % HL; range 33.6-35.7 % HL), and in P. genalutea from Yofu Bay (mean

23.8 % HL; range 23.1-24.6 % HL; mean 34.0 % HL; range 32.5-35.6 % HL).

76 P. pyroscelos 0.10 P. genalutea

P. genalutea

)

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Figure 23: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia pyroscelos from Mkanila Bay and Same Bay, Chizumulu Island (N = 28) PSU 4753; PSU 4752; PSU 4754; Petrotilapia genalutea (N = 3) from Mbweca Rocks, PSU 4750; Petrotilapia genalutea (N = 5) from Yofu Bay, PSU 4749.

Variables with the highest loadings on the sheared second principal component were snout length (0.35), lower jaw length (0.38), and preorbital head length (0.60); while those with the highest loadings on the principal components of the meristic data were teeth on outer left lower jaw (0.38), teeth rows on upper jaw (0.55), and teeth rows on lower jaw (0.58).

77 Table 9: Morphometric and meristic values of Petrotilapia pyroscelos, from Mkanila Bay, Chizumulu Island, Malaŵi, (N = 21) PSU 4753; PSU 4752, and Petrotilapia pyroscelos from Same Bay, Chizumulu Island, Lake Malaŵi, Africa (N = 7) PSU 4754. Ranges include the holotype.

Variable Holotype Mean SD Range Standard length (mm) 107.2 92.3 8.8 71.9-106.1 Head length (mm) 33.1 29.0 2.6 23.2-33.5 Percent standard length Body depth 34.5 35.0 1.1 32.1-37.6 Snout to dorsal-fin origin 32.4 33.7 1.4 31.0-36.1. Snout to pelvic-fin origin 40.6 39.2 1.1 37.2-41.3 Dorsal-fin base length 59.6 60.6 1.4 57.9-62.8 Anterior dorsal to anterior anal 54.1 53.4 1.1 50.5-55.4 Anterior dorsal to posterior anal 63.9 63.5 1.3 60.7-65.6 Posterior dorsal to anterior anal 30.8 30.0 0.9 27.9-31.7 Posterior dorsal to posterior anal 16.3 15.2 0.6 14.1-16.5 Posterior dorsal to ventral caudal 18.1 17.8 0.7 15.9-19.3 Posterior anal to dorsal caudal 19.2 19.0 0.8 17.1-20.5 Anterior dorsal to pelvic fin origin 37.4 37.5 1.2 34.1-39.9 Posterior dorsal to pelvic fin origin 56.0 57.0 1.5 53.6-59.1 Caudal peduncle depth 15.4 14.2 1.1 11.6-16.3 Least caudal peduncle depth 13.2 12.8 0.4 11.9-16.3 Percent head length Snout length 38.6 36.3 1.7 33.5-39.9 Postorbital head length 37.4 35.9 1.0 33.6-37.7 Horizontal eye diameter 31.5 32.3 1.5 28.9-35.9 Vertical eye diameter 32.6 33.7 1.0 31.1-35.9 Head depth 85.3 84.5 5.0 74.0-93.3 Preorbital depth 28.2 25.3 1.5 22.2-28.6 Cheek depth 26.6 23.8 1.5 20.6-26.6 Lower jaw length 36.8 35.7 2.2 29.2-38.6 Counts Mode %Frequency Range Dorsal-fin spines 18 18 64.3 17-19 Dorsal-fin rays 9 9 71.4 8-10 Anal-fin spines 3 3 100 _3_ Anal-fin rays 7 8 67.9 7-8 Pelvic-fin rays 5 5 100 _5_ Pectoral-fin rays 14 14 57.1 12-15 Lateral line scales 30 31 46.4 29-33 Pored scales past lateral line 2 2 64.3 1-2 Scale rows on cheek 4 4 78.6 3-5 Gillrakers on first ceratobranchial 11 10 53.6 8-11 Gillrakers on first epibranchial 3 3 89.3 2-3 Teeth on outer row of left lower jaw 13 14 25.0 9-20 Teeth rows on upper jaw 14 12 28.6 10-18 Teeth rows on lower jaw 15 13 28.6 10-18

78

Petrotilapia flaviventris n. sp. (Fig. 24)

Petrotilapia sp. ‗yellow ventral‘ Ribbink et al. 1983.

HOLOTYPE.—PSU 4760, adult male 110.9 mm; Same Bay, Chizumulu Island,

Lake Malaŵi, Africa; Grant, Konings, Stauffer, collectors; February 2006 (Fig. 24).

PARATYPES.—PSU 4759, 17, (68.3-110.89 mm) same data as for holotype.

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed clearly places this species in the genus Petrotilapia. Absence of a dark submarginal band in the dorsal fin (Fig.24) distinguishes it from male P. microgalana, P. genalutea, P. nigra, P. chrysos, P. mumboensis, and P. pyroscelos which have such a band. Males of P. flaviventris are distinguished from those of P. tridentiger, P. xanthos, and P. palingnathos. Adult males of P. flaviventris are yellow on ventral and mid sides with scales outlined in blue; the dorsal one-third is blue to gray with yellow and orange highlights; cheeks and gular are yellow to orange. Males of P. tridentiger are light blue with dark blue bars; while those of P. xanthos are yellow. Males of P. palingnathos are dark gray with orange markings and scales outlined in blue; cheeks are orange with a blue gular. Females of P. flaviventris are yellow brown with interrupted black stripes and with a background coloration fading to dark brown dorsally, which distinguishes them from those of P. tridentiger which are brown and from P. chrysos and P. microgalana which are golden yellow. Females of P. flaviventris are further distinguished from those of P. xanthos which are light brown, and from P. palingnathos which are orange brown.

Morphometric and meristic data for P. flaviventris are shown in Table 10.

79

Figure 24: Petrotilapia flaviventris, HOLOTYPE, PSU 4760, adult male, 110.9 mm SL, from Same Bay, Chizumulu Island, Lake Malaŵi, Malaŵi.

Description.—Petrotilapia flaviventris has a moderate deep body in comparison to other type species (see values on page 64); BD 38.5 % SL; mean 37.6 % SL. Greatest body depth at about base of eighth to ninth dorsal spine. Dorsal body profile gradually curving to caudal peduncle. Dorsal head profile concave between snout tip and interorbital, making 40–50° angle with body axis, then rounding to dorsal fin origin; horizontal and vertical eye diameter (29.7 and 30.7 % HL) larger than depth preorbital

(26.9 % HL) Snout with isognathous jaws and somewhat thickened lips; teeth on lower jaw in 10–20 rows with both outer row and inner rows tricuspid, except for posterior end rows which are unicuspid. Dorsal fin with XVII–XVIIII (mode XVIII) spines and 8–9

(mode 9) soft rays. Anal fin with III spines and 7–8 (mode 8) soft rays. First 3–5 dorsal

1 spines gradually increasing in length posteriorly with first spine less than /2 length of eighth and ninth spine; last 13 spines slightly increasing in length posteriorly with last

80 spine longest; soft dorsal with rounded or subacuminate tip, second, third ray longest, not or about reaching base of caudal fin; in some specimens past base of caudal fin. Anal spines progressively increasing in length posteriorly; third or fourth ray longest, reaching or past base of caudal fin, at about the same as dorsal in males. Caudal fin subtruncate to emarginate. Pelvic fin not reaching anal fin in females; not reaching or reaching first anal spine in males. Pectoral fin rounded, paddle-shaped, short, reaching vertical through base of 13th or 14th dorsal spine. Flank scales large, ctenoid; cheek with 3–4 rows of scales.

Lateral line with 30-31 pored scales.

Live coloration.—Males with yellow cheek, ventral preopercle, opercle and gular, preopercle, and opercle with blue highlights; interorbital dark blue (Fig. 25). Lateral coloration blue and yellow; ventral one-third yellow, mid one-third yellow with scales outlined in blue and yellow highlights; breast and belly yellow. Dorsal fin yellow and blue with blue membrane. Caudal fin with blue rays and yellow membrane. Anal fin spines and membrane black, anterior rays black, posterior rays blue with a single ocelli anal fin membrane blue and black. Pelvic fin yellow with a blue leading edge. Pectoral fin with yellow rays and clear membrane. gold with 6-7 faint gray bars. Females with dark brown heads, black opercle with blue green highlights and a gray gular. Lateral coloration yellow brown with interrupted black stripes and fading to dark brown dorsally.

Breast and belly light gray. Dorsal spines brown, proximal two-thirds of membrane brown, distal one-third light blue; gray brown lappets, and membrane of rays with brown spots. Caudal fin with brown rays and blue gray membrane. Proximal two-thirds light brown, distal one-third clear with no ocelli. Pelvic fin with first two rays with yellow

81 brown membrane, remaining membrane clear. Pectoral fin with brown rays and clear membrane.

Preserved coloration.—Males with brown head, light brown cheek, black opercle, and light brown to yellow gular. Lateral coloration of ventral and mid side light brown to yellowish and scales outlined in brown with 6 faint brown vertical bars; Lateral coloration dorsally dark brown with dark spots; belly and breast light brown to white.

Dorsal fin light brown to white with clear spines, rays and membrane; dorsal fin without black submarginal band. Caudal fin and rays light brown with clear membrane. Anal fin light brown with dark rays and spine and dark leading edge. Pelvic fin light brown with clear spine, rays and membrane. Pectoral fin with light brown rays and clear membrane.

Females with dark brown heads, black opercle, brown cheek and brown to light brown gular. Lateral coloration brown with black stripes past hypural plate and forming blotches on some places; belly yellow to whitish, breast brown to yellow. Dorsal fin light brown with clear membrane and no submarginal black band; brown spots on rays. Caudal fin with gray rays, clear membrane and black blotch. Anal fin with brown rays, spines and membrane clear. Pelvic fin with brown spine and anterior rays; posterior rays and membrane clear (Fig. 25).

82 a b

c d

Figure 25: Petrotilapia flaviventris: male (a) and female (b) at Same Bay, Chizumulu Island, Malaŵi (photos by A. Konings); preserved coloration of male PSU 4759 (c) and female PSU 4759 (d).

Distribution.—Petrotilapia flaviventris is found at Chizumulu Island. Konings

(2007) reports the presence of P. flaviventris between Mbweca, Mozambique and Undu

Point, Tanzania.

Etymology.—The name flaviventris, from flavus (Latin), meaning yellow and ventriculus (Latin), meaning belly, referring to the yellow color on the ventral side of this species.

Discussion.—Petrotilapia flaviventris is a member of the P. nigra group and mature males are easily distinguished from those of P. tridentiger, P. xanthos, and P.

83 palingnathos, which also lack a dark submarginal band in the dorsal fin, by its pigmentation pattern. Comparison of P. flaviventris with P. xanthos, P. palingnathos, and

P. microgalana, which occur in the same locality and nearby areas, using the principal component analysis revealed distinct clustering of P. flaviventris from the other three species when the first principal component of the meristic data are plotted against the sheared second principal component of the morphometric data (Figs. 16 above, 26).

0.100 P. flaviventris P. microgalana

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Figure 26: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia flaviventris (N = 18) PSU 4760; PSU 4759, and Petrotilapia microgalana (N = 40) PSU 3390; PSU3453; PSU 4755; BC 003; AMNH 238686.

84 Variables with the highest loadings on the sheared second principal component were least caudal peduncle depth (-0.32), anterior dorsal to pelvic fin origin (0.36), and posterior dorsal to posterior anal (-0.40); while those with the highest loadings on the principal components of the meristic data were teeth on outer left lower jaw (0.49), teeth rows on lower jaw (0.51), and teeth rows on upper jaw (0.53).

85 Table 10: Morphometric and meristic values of Petrotilapia flaviventris, from Same Bay, Chizumulu Island, Lake Malaŵi, Africa , (N = 18) PSU 4760; PSU 4759. Ranges include the holotype.

Variable Holotype Mean SD Range Standard length (mm) 110.9 94.1 14.2 68.0-111.2 Head length (mm) 36.2 30.5 4.5 21.6-36.2 Percent standard length Body depth 38.5 37.6 0.8 36.3-39.0 Snout to dorsal-fin origin 33.7 33.3 0.8 32.1-34.9 Snout to pelvic-fin origin 42.1 40.9 1.2 37.4-42.9 Dorsal-fin base length 63.5 61.2 1.5 57.9-63.5 Anterior dorsal to anterior anal 55.8 55.5 1.1 52.7-57.0 Anterior dorsal to posterior anal 66.2 64.9 1.3 61.6-66.4 Posterior dorsal to anterior anal 32.5 31.2 0.7 29.9-32.7 Posterior dorsal to posterior anal 16.4 15.7 0.6 13.7-16.5 Posterior dorsal to ventral caudal 17.5 17.1 0.4 16.1-17.8 Posterior anal to dorsal caudal 18.3 18.3 0.5 17.4-19.3 Anterior dorsal to pelvic fin origin 41.5 41.1 0.9 38.5-41.6 Posterior dorsal to pelvic fin origin 59.7 40.4 1.6 56.7-62.1 Caudal peduncle depth 12.7 12.3 0.6 11.4-13.7 Least caudal peduncle depth 13.7 12.9 0.3 12.4-13.7 Percent head length Snout length 38.6 37.2 1.8 32.6-39.9 Postorbital head length 39.0 37.6 1.1 35.5-40.1 Horizontal eye diameter 29.7 30.5 1.7 28.0-34.7 Vertical eye diameter 30.7 31.3 1.5 28.6-33.7 Head depth 93.2 90.6 1.0 86.9-94.8 Preorbital depth 26.9 26.2 2.4 24.2-28.6 Cheek depth 26.2 25.6 1.7 22.4-29.9 Lower jaw length 34.3 34.7 1.0 33.0-36.4 Counts Mode %Frequency Range Dorsal-fin spines 18 18 66.7 17-19 Dorsal-fin rays 9 9 77.8 8-9 Anal-fin spines 3 3 100 _3_ Anal-fin rays 8 8 88.9 7-8 Pelvic-fin rays 5 5 100 _5_ Pectoral-fin rays 14 14 61.1 13-14 Lateral line scales 30 30 61.1 30-31 Pored scales past lateral line 1 2 77.9 1-2 Scale rows on cheek 4 4 88.9 3-4 Gillrakers on first ceratobranchial 11 10 83.3 10-11 Gillrakers on first epibranchial 2 3 72.2 2-3 Teeth on outer row of left lower jaw 14 13 27.8 10-18 Teeth rows on upper jaw 14 14 27.8 10-17 Teeth rows on lower jaw 16 13 22.2 11-18

86

Petrotilapia palingnathos n. sp. (Fig. 27)

Petrotilapia ‗retrognathous‘ Ribbink et al. 1983.

HOLOTYPE.—PSU 4767, adult male 118.0 mm; S 12o 00.566' E 34° 36.882'

Same Bay, Chizumulu Island, Lake Malaŵi, Africa; Stauffer and Konings collectors;

February 2006 (Fig. 27).

PARATYPES.—PSU 4766, 5, (96.29-117.72 mm) same data as for holotype.

Diagnosis.—The presence of predominantly triscupid teeth in both outer and inner rows of the upper and lower jaws that are visible when the mouth is closed clearly places this species in the genus Petrotilapia. Absence of a dark submarginal band in the dorsal fin (Fig. 27) distinguishes it from P. microgalana, P. genalutea, P. nigra, P. chrysos, P. mumboensis, and P. pyroscelos which have such a band. Adult males of P. palingnathos are dark gray with orange markings and scales outlined in blue with orange cheeks and a blue gular, which distinguishes them from P. tridentiger, which are light blue with dark blue bars. Adult males of P. palingnathos are also distinct from those of P. xanthos and P. flaviventris. Males of P. xanthos are yellow while those of P. flaviventris are yellow on ventral and mid sides with scales outlined in blue; the dorsal one-third is blue to gray with yellow and orange highlights; cheeks and gular are yellow to orange.

Females of P. palingnathos differ from those of P. tridentiger, P. chrysos, P. microgalana, P. xanthos, and P. flaviventris by the orange brown color, which distinguishes them from those of P. tridentiger which are brown and from P. chrysos and

P. microgalana which are golden yellow. Females of P. palingnathos are further

87 distinguished from those of P. xanthos which are light brown, and from P. flaviventris, which are yellow-brown with interrupted black stripes and fading to dark brown dorsally.

Both vertical eye diameter and horizontal eye diameter are smaller in P. palingnathos (mean 26.2 % HL; range 25.6-27.7 %; mean 24.9 % HL; range 23.6-26.2) than in P. tridentiger (mean 29.0 % HL; range 25.2-34.9 % HL; mean 28.5 % HL; range

23.6-33.6 % HL) respectively. Caudal peduncle depth is larger in P. palingnathos (mean

15.0 % SL; range 13.8-17.6 % SL) than in P. tridentiger (mean 13.3 % SL; range 10.8-

15.4 % SL).

Morphometric and meristic data for P. palingnathos are shown in Table 11.

Figure 27: Petrotilapia palingnathos, HOLOTYPE, PSU 4767, adult male, 118.0 mm SL, from Same Bay, Chizumulu Island, Lake Malaŵi, Malaŵi.

Description.—Petrotilapia palingnathos is a deep body species (BD 38.0 % SL), similar to P. flaviventris (38.8 % SL), but smaller than P. mumboensis (42.0 % SL)

Greatest body depth at about base of ninth dorsal spine. Dorsal body profile gradually curving to caudal peduncle. Dorsal head profile concave between snout tip and interorbital, making 50–60° angle with body axis, then rounding to dorsal fin origin; horizontal and vertical eye diameter (24.8 and 25.6 % HL) similar to depth preorbital

88 (24.9). Snout with isognathous jaws and thickened lips; teeth on lower jaw in 18–26 rows with both outer row and inner rows tricuspid, except for posterior end rows which are unicuspid.

Dorsal fin with XVII–XVIII (mode XVIII) spines and 8–10 (mode 9) soft rays.

Anal fin with III spines and 7–8 (mode 8) soft rays. First 2–4 dorsal spines gradually

1 increasing in length posteriorly with first spine less than /2 length of ninth spine; last 13 spines slightly increasing in length posteriorly with last spine longest; soft dorsal with rounded or subacuminate tip, fourth ray longest, reaching past base of caudal fin in males. Anal spines progressively increasing in length posteriorly; third or fourth ray longest, reaching about base of caudal fin, slightly shorter than dorsal in males. Caudal fin subtruncate to emarginate. Pelvic fin reaching second anal spine to first ray in males.

Pectoral fin rounded or subacuminate, paddle-shaped, short, reaching vertical through base of 12th or 13th dorsal spine. Flank scales large, ctenoid; cheek with 4 rows of scales.

Lateral line with 30-31 pored scales;

Live coloration.—Males with orange cheek and preorbital, preopercle and opercle orange with blue highlights; gular blue; interorbital dark gray with light blue bar. Lateral coloration dark gray with scales outlined in blue, and with some orange markings. Dorsal spines dark blue gray; blue membrane with orange markings. Caudal fin with blue rays and orange membrane. Anal fin with faint orange cast and two yellow ocelli. Pelvic fin with yellow to clear membrane. Pectoral fins with brown rays and clear membrane (Fig.

28). Live coloration for female P. palingnathos was not recorded in the field; the following description of females is based on the photo taken from Konings (2007).

89 Females with an overall orange brown color with 8 dark brown vertical bars; dark brown opercle; gular, breast and belly yellow to orange. Dorsal fin without black submarginal band; spines and rays brown with orange lappets and clear membrane. Pelvic fin brown with blue leading edge and clear membrane. Pectoral fin brown with clear membrane

(Fig. 28).

Preserved pattern.—Males with brown heads, black opercle, and light brown gular. Lateral coloration dark brown and without vertical bars; belly and breast brown.

Dorsal fin without black submarginal band; spines rays, and membrane gray. Caudal fin with dark brown rays and brown membrane. Anal fin light brown with two light brown ocelli; membrane light brown. Pelvic fin with light brown spine and rays; membrane clear. Pectoral fin with light brown rays and clear membrane (Fig. 28). Preserved coloration of females is not recorded because there were no female preserved specimens.

a b

c

Figure 28: Petrotilapia palingnathos: male (a) and female (b) at Chizumulu Island (photos by A. Konings); preserved coloration of male PSU 4766 (c).

90 Distribution.—Petrotilapia palingnathos occurs at Chizumulu Island. Konings

(2007) reports this species‘ presence at Taiwanee Reef as well. It is normally found in the shallow but not wave-washed habitat. The presence of P. palingnathos at Taiwanee Reef may indicate that it is an ―old‖ species which inhabited the lake when the water was at a much lower level several tens of thousands of years ago (Konings, 2007).

Etymology.—The name palingnathos, from the Greek, meaning backward jaw, referring to the shorter lower jaw.

Discussion.—Petrotilapia palingnathos is member of the P. tridentiger group and easily identified by the downward-opening mouth, a character which is unique to this species of Petrotilapia (Konings, 2001; 2007). Comparison of P. palingnathos with P. xanthos, P. flaviventris, and P. tridentiger, males of which also lack a dark submarginal band in the dorsal fin, using the principal component analysis revealed distinct clustering of P. palingnathos from the three species when the first principal component of the meristic data are plotted against the sheared second principal components of the morphometric data (Figs. 16, 29).

91 0.15 P. palingnathos P. tridentiger P. tridentiger

) 0.10

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Figure 29: Plot of the second sheared principal component (morphometric data) and the first factor scores (meristic data) of Petrotilapia palingnathos (N = 6) PSU 4767; PSU 4766; Petrotilapia tridentiger Nkhata Bay (N = 7) PSU 3455, and Petrotilapia tridentiger (N = 48) BMNH 1935.6.14.249.263; PSU 3455; PSU 4805; PSU 4806; PSU 4807; PSU 4808; PSU 4809.

Variables with the highest loadings on the sheared second principal component were vertical eye diameter (0.39), horizontal eye diameter (0.52), and caudal peduncle depth (-0.62); while those with the highest loadings on the principal components of the meristic data were teeth on outer left lower jaw (0.39), teeth rows on upper jaw (0.43), and teeth rows on lower jaw (0.52).

92 Table 11: Morphometric and meristic values of Petrotilapia palingnathos, from Mkanila Bay, Chizumulu Island, Lake Malaŵi, Africa , (N = 6) PSU 4767; PSU 4766. Ranges the include holotype.

Variable Holotype Mean SD Range Standard length (mm) 118.0 111.5 8.3 96.3-118.0 Head length (mm) 37.7 36.3 2.3 32.4-38.4 Percent standard length Body depth 38.0 38.6 0.3 38.0-38.9 Snout to dorsal-fin origin 34.1 35.1 0.8 34.1-36.2 Snout to pelvic-fin origin 41.0 40.5 0.9 38.8-41.2 Dorsal-fin base length 61.5 61.3 1.2 58.9-62.5 Anterior dorsal to anterior anal 56.3 55.6 0.3 55.1-56.3 Anterior dorsal to posterior anal 65.4 65.1 0.9 63.5-66.4 Posterior dorsal to anterior anal 30.6 30.4 0.4 29.4-30.7 Posterior dorsal to posterior anal 16.0 16.2 0.5 15.3-16.7 Posterior dorsal to ventral caudal 18.5 18.1 0.5 17.5-18.7 Posterior anal to dorsal caudal 19.4 19.3 0.2 18.8-19.6 Anterior dorsal to pelvic fin origin 39.0 39.8 1.0 38.5-41.2 Posterior dorsal to pelvic fin origin 55.9 57.0 1.0 55.9-58.5 Caudal peduncle depth 17.6 15.0 1.4 13.8-17.6 Least caudal peduncle depth 13.7 13.9 0.2 13.7-14.1 Percent head length Snout length 40.5 41.9 1.4 40.5-44.5 Postorbital head length 34.6 36.5 1.4 34.6-38.8 Horizontal eye diameter 24.8 24.9 0.9 23.6-26.2 Vertical eye diameter 25.6 26.2 0.8 25.6-27.7 Head depth 92.6 91.0 2.4 88.5-94.8 Preorbital depth 24.9 27.6 2.3 24.9-30.8 Cheek depth 25.8 26.6 0.9 25.2-27.6 Lower jaw length 35.2 34.0 1.3 31.5-35.2 Counts Mode %Frequency Range Dorsal-fin spines 18 18 66.7 17-18 Dorsal-fin rays 9 9 50.0 8-10 Anal-fin spines 3 3 100 _3_ Anal-fin rays 8 8 66.7 7-8 Pelvic-fin rays 5 5 100 _5_ Pectoral-fin rays 14 14 66.7 14-15 Lateral line scales 31 31 66.7 30-31 Pored scales past lateral line 2 1 50.0 1-2 Scale rows on cheek 4 4 100 -4- Gillrakers on first ceratobranchial 10 10 83.3 9-10 Gillrakers on first epibranchial 2 2 100 -2- Teeth on outer row of left lower jaw 25 20 33.3 19-25 Teeth rows on upper jaw 21 17 33.3 16-21 Teeth rows on lower jaw 26 26 33.3 18-26

93

Chapter 4

Discussion and conclusion

Petrotilapia was established by Trewavas (1935) as having all tricuspid teeth that are visible when the mouth is closed. Subsequently, Marsh (1983) modified the generic diagnosis to include cichlids with predominantly triscupid teeth because all the species he examined had some distinctly unicuspid teeth. I observed some unicuspid tooth rows on the innermost rows of both lower and uppers jaws in all the specimens I examined.

I worked on several populations and species of Petrotilapia. The genus

Petrotilapia is a widely distributed group of rock-dwelling cichlids of Lake Malaŵi. Prior to my work, the genus Petrotilapia consisted of five species namely P. tridentiger, P. genalutea, P. nigra, P. chrysos, and P. microgalana. Petrotilapia genalutea is the most wide spread species of the genus. Comparison of P. ‗ruarwe‘ from Kakusa and Mbowe

Island with P. microgalana using principal component analysis revealed that these two populations of Petrotilapia north of Nkhata Bay are conspecific with P. microgalana when the first principal component of the merisitc data is plotted against the sheared second principal component of the morphometric data. Not only did these populations overlap but also females of both P. microgalana and P. ‗ruarwe‘ have the same pigmentation pattern consisting of two horizontal rows of dots of about the same size with the mid-lateral row seemingly larger. Additionally, males of both species have a black submarginal band in the dorsal fin. The three populations are also from neighboring sites. I, therefore, consider P. ‗ruarwe‘ from Kakusa and Mbowe Island P. microgalana.

I compared P. sp.‗mumbo yellow‘ from Mumbo Island with P. nigra from

Thumbi West Island. Konings (2007) reports females of P. sp. ‗mumbo yellow‘ as having

94 the characteristic pattern of the P. nigra group (yellow or golden background color and a pattern consisting of two horizontal rows of dots) and the strong yellow color of the males as being uncommon and found only in members of the P. nigra group in the northern half of the lake. Although the strong yellow color of the males of P. sp. ‗mumbo yellow‘ is not common among populations in the south of the lake, I found no morphological difference between the two neighboring populations of P. nigra and P. sp.‗mumbo yellow‘ (Fig. 8). Petrotilapia sp. ‗mumbo yellow‘ is indistinguishable from

P. nigra. I, therefore, consider P. sp. ‗mumbo yellow‘ a geographical variant of P. nigra.

I recognized five new species of Petrotilapia namely P. xanthos from Gallireya

Reef, P. mumboensis from Mumbo and Thumbi West islands, P. pyroscelos, P. flaviventris, and P. palingnathos all from Chizumulu Island. These species were distinguished on the basis of color pattern and markings, species group, and morphometric and meristic differences. The above mentioned new species were compared with neighboring populations and were distinguishable from such other populations.

The genus Petrotilapia still contains species that are known but not scientifically described yet. Some of the species are P. ‗black flank‘ (P. nigra group), P. ‗chitande‘

(only species that belongs to the P. genalutea group), P. ‗fuscous‘ (P. nigra group), P. sp.

‗likoma variable‘ (P. nigra group), P. ‗yellow chin‘ (P. tridentiger group), and P. sp.

‗likoma barred‘ (P. tridentiger group). Petrotilapia sp. ‗likoma barred‘ is the largest mbuna known attaining a maximum total length of 19 cm (Konings, 2007). It is evident from the above data that there remains more species in the genus Petrotilapia to be described.

95 Chapter 5

Taxonomic key to the Species of Petrotilapia: Species Identification

The vent is the opening between anus and anal fin where the fish will excrete either eggs or sperm; the vent is more rounded and broader in females than in males where it is pointed and thinner. Photo by Gary Kratochvil, 1997.

96 Key to described species of Petrotilapia

1a. Absence of a black submarginal band in the dorsal fin of males……….2

1b. Presence of a black submarginal band in the dorsal fin of males……….5

2a. Live adult males light blue with dark blue vertical bars; purple blue cheek; whitish blue gular; females dark brown with dark vertical bars……….Petrotilapia tridentiger

2b. Live adult males not light blue; vertical bars not dark blue; females not dark brown and without vertical bars……….3

3a. Adult males blue and yellow; ventral one-third yellow, mid one-third yellow with scales outlined in blue and yellow highlights; cheeks and gular yellow to orange; females yellow brown with interrupted black stripes and fading to dark brown dorsally………..Petrotilapia flaviventris

3b. Adult males not blue and yellow; ventral one-third and mid one-third not yellow; cheeks and gular not yellow to orange; females not yellow brown and without interrupted black stripes…………..4

97 4a. Adult males dark gray with orange markings and scales outlined in blue; orange cheeks; blue gular; females orange brown with 8 dark brown vertical bars and without horizontal pigmentation; both males and females with downward-opening mouth……….Petrotilapia palingnathos

4b. Live adult males entirely yellow with 6-7 faint vertical bars; females light brown with 6 dark brown vertical bars and with horizontal pigmentation……….Petrotilapia xanthos

5a. Adult males with vertical bars; females with horizontal pigmentation pattern…….6

5b. Adult males with vertical bars; females without horizontal pigmentation pattern….8

98 6a. Adult males dull gray-blue with 5-7 black vertical bars; orange-yellow cheek, and a black gular; females off-white to pale yellow-brown……….Petrotilapia genalutea

6b. Adult males not dull gray to blue; vertical bars 7 or more; cheek not orange-yellow; females not off-white to pale yellow-brown……….7

7a. Adult males predominantly blue-black with 7-10 gray brown vertical bars; dark-blue cheek, and a black gular; females pale brown……….Petrotilapia nigra

99 7b. Adult males bright blue with 5-7 faint black vertical bars; light-blue cheek and a yellow gular; females golden yellow and without a black submarginal band……….Petrotilapia microgalana

8a. Adult males light blue overall color; 8 dark-blue vertical bars; light blue cheek; light blue to gray gular; two light blue interorbital bars; females gray brown to light brown……Petrotilapia mumboensis

Interorbital bars

8b. Adult males not light blue overall color; vertical bars not dark-blue and less than 8; cheek not light blue; gular not light blue to gray; no interorbital bars; females not gray brown to light brown………….9

100 9a. Adult males blue with bronze highlights and orange pelvic; 5 gray vertical bars; purple cheek; gular whitish to gray; females brown with faint blue and yellow highlights………….Petrotilapia pyroscelos

9b. Live adult males dark blue with 7-9 black vertical bars; non-territorial males gold with blue highlights and 7-9 black vertical bars; females golden brown and without a black submarginal band……Petrotilapia chrysos

Non-territorial male

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109 VITA

MARY LUNDEBA

P. O. Box 350103 Chilanga, Lusaka, Zambia. (+260 966 533 148)

[email protected]

EDUCATION

Ph.D. Wildlife and Fisheries Science The Pennsylvania State University, University Park, PA 16802; August 2006-August 2009

Master of Science, Aquaculture and Fisheries Science University of Malaŵi, Bunda College of Agriculture, Lilongwe, Malawi, August 2003- November 2005

Bachelor of Science, Agriculture, Fisheries Major University of Malaŵi, Bunda College of Agriculture, Lilongwe, Malaŵi, October 1995- April 2000

TEACHING EXPERIENCE August 2007-May 2009 Teaching Assistant, Ichthyology Laboratory Course (WFS 301) Duties: Taught lectures on vertebrate biology, set exams, quizzes and did the grading. The Pennsylvania State University, School of Forest Resources, University Park, PA 16802

January 2005-May 2006 Teaching Assistant, University of Malaŵi, Bunda College of Agriculture Course Taught: Integrated Aquaculture Farming Systems

2000-2002: U.S. Peace Corps Technical Trainer, National Aquacultural Development Center, Kitwe, Zambia (Rural Aquaculture Development Project) Causes Taught: Fish Biology, Fish Ecology, Fish Nutrition, Water Quality Management, and Rural Development courses

Scholarship Awards Icelandic International Development Agency (ICEIDA) 1995-2000 Japanese International Cooperation Agency (JICA) 2001 NFS/NIH Joint Program, Ecology of Infectious Diseases 2003-2006 Elizabeth Bowers Zambia Education Fund (EBZEF) 2003 The Pennsylvania State University, School of Forest Resources 2006-2009 P.E.O. International Peace Scholarship 2007-2008