The Morphometrical Variance and Abundance Of
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The Pennsylvania State University The Graduate School College of Agricultural Sciences REVIEW OF THE GENUS DIPLOTAXODON WITH A DESCRIPTION OF TWO NEW SPECIES A Thesis in Wildlife and Fisheries Science by Titus Bandulo Phiri © 2013 Titus Bandulo Phiri Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science December 2013 The thesis of Titus Bandulo Phiri was reviewed and approved* by the following: Jay R. Stauffer, Jr. Distinguished Professor of Ichthyology Thesis Adviser Walter Tzilkowski Associate Professor Emeritus of Wildlife Wilson Lazaro Jere Senior Lecturer of Fish Genetics Jeremy Likongwe Associate Professor of Fisheries Sciences Michael G. Massina Head and Professor Department of Ecosystems Science and Management *Signatures are on file in the Graduate School ii ABSTRACT A review of the described species of the genus Diplotaxodon Trewavas 1935, deep water (offshore) fishes endemic to Lake Malaŵi, was conducted and included the description of two new species. Diplotaxodon spp. represent another example of the rapid radiation of the Haplochromine species in Lake Malaŵi. Fishes were collected from Domira Bay and Senga Bay. Morphometric and meristic differences among spp. were analyzed using Sheared Principal Component Analysis (SPCA) for the morphometric data and Principal Component Analysis (PCA) for the meristic data. To date seven species of Diplotaxodon are formally described. The investigations from this study resulted into describing two new species which were known by cheironym. iii TABLE OF CONTENTS LIST OF FIGURES .......................................................................................................................... vi LIST OF TABLES ............................................................................................................................ix AKNOWLEDGEMENTS .................................................................................................................xi CHAPTER 1 ..................................................................................................................................... 1 1.0.0 GENERAL INTRODUCTION ........................................................................................... 1 1.1.0 Study area ...................................................................................................................... 2 1.2.0 Study goals and objectives ........................................................................................................ 5 2.0.0 GENERAL METHODOLOGY ................................................................................................ 7 2.2.0 Collection of Fish Specimen.................................................................................................. 7 2.3.0. Data collection (Meristic and Morphometric) ....................................................................... 7 2.4.0 Meristic data ....................................................................................................................... 10 2.5.0 Data analysis ....................................................................................................................... 14 3.0.0 OVERVIEW OF THE GENUS Diplotaxodon TREWAVAS 1935 ...................................... 14 3.1.0 General Distribution of Diplotaxodon.................................................................................. 15 4.0.0 DESCRIBED SPECIES .......................................................................................................... 15 4.1.0 Diplotaxodon apogon Turner & Stauffer, 1998 ................................................................ 15 4.2.0. Diplotaxodon ecclesi Burgess & Axelrod, 1973 .............................................................. 19 4.3.0 Diplotaxodon macrops Turner & Stauffer, 1998 .............................................................. 20 4.4.0 Diplotaxodon aeneus Turner & Stauffer, 1998 ................................................................. 22 4.5.0 Diplotaxodon limnothrissa Turner, 1994 ........................................................................ 26 4.6.0 Diplotaxodon greenwoodi Stauffer &McKaye, 1986....................................................... 32 4.7.0 Diplotaxodon argenteus Trewavas, 1935 ........................................................................ 38 5.0.0. Overview of undescribed species........................................................................................ 45 CHAPTER 2 ................................................................................................................................... 47 6.0.0 DESCRIPTION OF TWO NEW SPECIES ......................................................................... 47 6.1.0. INTRODUCTION ................................................................................................................. 47 7.0.0 New species of Diplotaxodon .............................................................................................. 50 Diplotaxodon altus n. sp., n. nov (nova species, novem nomen) (Fig. 18)..................................... 50 Diplotaxodon sp. ‘deep’. .............................................................................................................. 50 7.1.0 Results ................................................................................................................................ 52 iv 8.0.0 Diplotaxodon maxillalongus n. sp., n. nov (nova species, novem nomen) (Fig. 18) ............. 67 Diplotaxodon sp. ‘maxillalongus’. ............................................................................................... 67 8.1.0 Results ................................................................................................................................ 70 9.0.0 Discussion .......................................................................................................................... 83 10.0.0 Conclusion ............................................................................................................................ 86 CHAPTER 3 ................................................................................................................................... 88 11.0.0 Dichotomous key for Diplotaxodon ....................................................................................... 88 13.0.0 Remarks............................................................................................................................ 92 LITERATURE CITED .................................................................................................................... 93 v LIST OF FIGURES Figure 1: Maps illustrating location of the study area, Lake Malaŵi showing Domira Bay and Senga Bay in Salima District. Inset is map of Africa showing The Great Lift Valley Lakes and location of Lake Malaŵi. (Source: Google earth and http://www.odci.gov/cia/publications) ..... 4 Figure 2: Diplotaxodon spp. showing landmarks and distances measured as indicated by the numbered landmarks, the measured points not shown in the figure include orbital depth, cheek depth, horizontal and vertical eye diameter. .............................................................................. 8 Figure 3: Illustration of counting of main external meristic sets of variables on dorsal spines, dorsal rays, anal spines, anal rays and lateral line scales in black dots. .............................................. 10 Figure 4: Counting model for rays on pectoral-fin and pelvic-fin as part of the meristic variables. .... 11 Figure 5: Format for counting scales on the cheek, scales appear to form a pattern/ series were considered as indicated by I, II, and III representing 3 series of scales .................................... 12 Figure 6: Part of the gill bar arch from the right hand side indicating how the gill rakers were counted for epibranchial and ceratobranchial. ...................................................................................... 12 Figure 7: Diplotaxodon apogon, holotype from British Museum, described by Turner and Stauffer in (1998). Holotype: BMNH 1996.4.30:21a ripe male of 87.8mm standard length (Source: Turner and Stauffer 1998).................................................................................................................. 17 Figure 8: Plots of the sheared second principle components (morphological data) and the first factor score, first principle component (meristic data) for Diplotaxodon apogon and Diplotaxodon limnothrissa sampled from commercial vessel in Domira Bay, Lake Malawi in 2012 ............. 18 Figure 9: Diplotaxodon ecclesi described from a male holotype: USNM 210696 of 145.7mm standard length, collected off Monkey Bay by Herbert R. Axelrod in 1973, redefined by Turner & Stauffer 1998. Photo by Dr. Herbert R. Axelrod (Source://fishbase.org). ................................ 20 Figure 10: Diplotaxodon macrops, described from a ripe male holotype: BMNH 1996.4.30:1, with 106.7mm in standard length and was collected from 100m depth off Monkey Bay at 14o03′S 34o56′E by Turner in 1991. (Source: //wikipedia.org). ............................................................ 22 Figure 11: Diplotaxodon aeneus, described from a ripe male of 125mm standard length, holotype: BMNH 1996.4.30:16, collected from a depth of 400m northeast of Nkhata Bay by ODA/SADC pelagic Resource Project between 30 November 1993-December 1994 (source: Turner and Stauffer 1998). ......................................................................................................................