PROCEEDINGS OF THE ACADEMY OF NATURAL SCIENCES OF PHLADELPHIA 148: 189-230. 31 OCTOBER 1997

A revision of the blue-blackPseudotropheus zebra (Teleostei: Cichlidae) complex from LakeMalaw'i, Africa, with a description of a new genus and ten new species

JAY R. STAUFFER,JR.,1 NANCY J. BOWERS,2KAREN A. KELLOGG1AND KENNETH R. MCKAYE3 'School of ForestResources, Penn State University, University Park, PA 16802, USA 2EnvironmentalSciences and Resources,Portland State University, Portland, OR 97207, USA 3AppalachianLaboratory, University of MarylandSystem, Frostburg, MD 21532, USA

ABSTRACT - A new genus, Metriaclima,is described for members of the Pseudotropheuszebra complex from Lake Malawi. The presence of bicuspid teeth in the anterior portion of the outer row of both the upper and lower jaws distinguishes Metriaclima from many of the previously described genera of rock-dwelling cichlids that inhabit Lake Malaw'i, including Cyathochromis,Cynotilapia, Gephyrochromis,Labidochromis, and Petrotilapia. The absence of two horizontal stripes along its flanks, distinguishes it from Melanochromis. The isognathous jaws of Metriaclimadelimits it from Genyochromis,which is characterizedby having the lower jaw extend in front of the upper jaw. The mouth of Metriaclima is terminal, while that of Labeotropheusis inferior. Within the genus Pseudotropheusas it is now recognized, species of Metriaclima are unique because they have a moderately sloped ethmo-vomerine block and a swollen rostral tip. Ten previously undescribed species that have a slight variation from the characteristic blue/black barring are described. The new species are recognized primarily by their distinctive adult coloring in conjunction with the discontinuity of morphological differences throughout their range. GENERA ET SPECIES NOVAE: Metriadima, M. melabranchion,M. chrysomallos,M. phaeos,M. cyneusmarginatus,M. benetos,M. pyrsonotos,M. sandaracinos,M. emmiltos, M. mbenjii, M. thapsinogen.

INTRODUCTION genera within the lake. Oliver & Loiselle (1972) later added Iodotropheusto this group. The majori- In the lakes of East Africa, fishes of the family ty of the species within this group are all strongly Cichlidae have undergone an extraordinarily rapid lithophilous and form complex communities and extensive radiation. Within Lake Malaw'ithere throughout the lake along the rocky shorelines and are over 450 described species of fish, and there rocky islands. Endemism within this group is may be as many as 1500 species which inhabit the high, and many species are restricted to a single, lake, all of which have arisen in the last 2 million isolated rock group (Fryer and Iles 1972, Ribbink years (Bowers et al. 1994). The cichlids within et al. 1983, Stauffer 1988). Although the mbuna Lake Malawi are representedby two major groups, cannot be distinguished from other genera of which are regardedas tribes by some investigators: cichlids on the basis of synapomorphies, they can the tilapiines, of which there are from five to seven be recognized by a suite of characters, some of species and the haplochromines, of which there are which may be shared by other deep-water and some 450 described species (Konings 1990). Within sand-dwelling haplochromine taxa. These charac- the haplochromines, two major and several minor ters include large number of small scales in the lineages exist; the major lineages being the nape and chest region, an abrupt transition from rock-dwelling cichlids, commonly called mbuna, large flank scales to small chest scales, reduction of and the pelagic and sand-dwelling cichlids, al- the left ovary, coloration, and habitat preferences though recent molecular evidence suggests splitting (Fryer 1959). The possession of true ocelli has also the second major lineage into several minor ones been suggested to be a distinguishing characteristic (Moran et al. 1994). of mbuna (Oliver 1984). Recently, Moran et al. Trewavas (1935) recognized the nine genera of (1994) identified three additional genera, Aulono- mbuna (Cyathochromis,Cynotilapia, Genyochromis, cara, Alticorpus, and Lethrinops, which although Gephyrochromis, Labeotropheus, Labidocbromis, lacking many of mbuna characters, shared an Melanochromis,Petrotilapia, and Pseudotropheus)to affinity with the mbuna on the basis of RFLP be more closely related to each other than to other analysis of mtDNA.

189 190 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE

350 E

NkhataBay

Chizimulu Island %U 0 MbenjeIsland Likoma Island

Nknotakota Namalenge IWand Mbenje Island NamalenjeIsland

onkey Bay

MaleriIslands

Domwe Island MumboIsland S \ Thumbi West \ Island *Thumbi East Island .Sangwe HIN /Mazinzi Reef *Mpanda Sland Kanjedza Island Boadzulu Island

\ Crocodile\ Rocks

MAP. Locations of sites mentioned in the text. PSEUDOTROPHEUSZEBRA COMPLEX 191

Of the aforementionedmbuna genera, Pseudo- used throughout. Except for gill raker meristics, tropheusis the most diverseand certainly polyphy- all counts and measurementswere made on the left letic. The type species of the genus is Pseudo- side of the fish. Morphometric values are ex- tropheuswilliamsi (Gunther), and species complexes pressed as percent SL or percent head length (HL). recognizedwithin the genus include the: P. wil- Differences in body shape were analyzed using liamsi complex, Pseudotropheustropheops Regan sheared principal component analysis (SPCA) of complex,Pseudotropheus elongatus Fryer complex, the morphometric data (Humphries et al. 1981, Pseudotropheusaggressive complex, and the Pseudo- Bookstein et al. 1985). Pectoral-fin length and tropheuszebra (Boulenger) complex. Additionally, pelvic-fin length were not included in the analysis, there is a miscellaneousgroup within Pseudo- because these measurements are dependent on the tropheusthat shows no clear affinitywith any of reproductive stage of the fish. Meristic data were the othercomplexes and is itselfcertainly polyphy- analyzed using principal component analysis letic (Ribbinket al. 1983). Finally,the placement (PCA). The correlation matrix was factored in all of Pseudotropheusfainzilberi Staeck in Pseudo- principal component analyses of meristic data, tropheusis problematic. Althoughits body color- while the covariance matrix was factored in the ation superficiallyresembles P. zebra,it differs, calculation of all sheared principal components of because of its peculiar mouth shape and teeth the morphometric data. This analysis ordinates position, which more closely resemblesthat of factors independently of a main linear ordination Petrotilapiaspecies (Staeck 1976, fig. 12, p. 489). (Reyment et al. 1984). Differences among species Currently,the P. zebracomplex consists of 13 were illustrated by plotting either the sheared described species including:P. zebra, P. elegans second or third principal components of the Trewavas,P. livingstonii(Boulenger), P. heteropictus morphometric data against the first principal Staeck,P. lombardoiBurgess, P. auroraBurgess, P. components (labeledas factor score) of the meristic IanisticolaBurgess, P. greshakeiMeyer and Foerster, data (Stauffer and Hert 1992). If the mean multi- P. barlowiMcKaye and Stauffer,P. xanstomachus variate scores of the clusters were significantly Staufferand Boltz, P. pursusStauffer, P. callainos different along one axis, independent of the other Stauffer and Hert, P. hajomaylandiMeyer and axis, a Duncan's multiple range test (p < 0.05) was Schartl,and P. estberaeKonings. In addition,there used to determine which clusters differed from are some 20 speciesthat are currentlyrecognized each other. If, in fact, the clusters were not by aquaristsfor which thereare no formaldescrip- significantly different along one axis independent tions (Ribbinket al. 1983,Konings 1990). As with of the others, then a MANOVA, in conjunction the majorityof the mbuna,most of these species with a Hotelling-Lawley trace, was used to deter- live over a rocky substrate;however, there are mine whether the mean multivariate scores of severalforms in this complex representedby P. clusters formed by the minimum polygons of the elegans,P. Zivingstonii,P. lanisticola,and P. pursus, PCA scores were significantly different (p<0.05). that have invadedthe sandhabitat and are associat- ed with snail shells either as juveniles,adults, or both life stages. The purposeof this paperis to TAXONOMIC ANALYSES describea new genusfor membersof the P. zebra complexand to describeten new speciesthat have [NOTE: Tables 1-11, containing morphometric and the characteristicblue/black (BB) barring found in meristic values for type and referred material, are P. zebra. Future papers will describe the grouped at the end of this paper.] sand-dwellingforms and the formsof this complex that do not conformto the BB color pattern. Metriaclima Stauffer, Bowers, Kellogg and McKaye, n. gen. METHODS AND MATERIALS TypeSpecies.-Metriaclima zebra Boulenger. Diagnosis.-This genus comprises a series of Adult fishes were collected throughoutLake rock-dwelling species endemic to Lake Mala'wi, Malawiby chasingthem into a monofilamentnet Africa. The presence of many small scales in the (7 m x 1 m x 1.5 cm) while SCUBA diving. nape region and the reduction of the left ovary Externalcounts andmeasurements follow Barelet aligns this genus to the other rock-dwelling genera al. (1977) and Stauffer(1991). The number of in Lake Malaw'i. The presence of bicuspid teeth in scalesin the lateral-lineseries do not includescales the anterior portion of the outer row of both the in the overlappingportion of the lower lateralline; upper and lower jaws distinguishes Metriaclima pored scaleslocated posterior to the hypuralplate from many of the previously described genera of were recordedseparately. Standard length (SL)is mbuna. Cyatbochromisis distinguished because its 192 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

Fig. 1. Skull of Metriadima zebra.

Fig. 2. Skullof Pseudotropheusdlongatus. teeth consist of slender shafts with compressed The mouth of Metriaclimais terminal, while that crowns. Cynotilapia,Gephyrocbromis, and Labido- of Labeotropheusis inferior. The absence of two chromisall have unicuspidteeth in the anterior horizontgal stripes along its flanks, delimits Metri- portion of the outer rows of both the upper and aclima from Melanochromis(Bowers and Stauffer lower jaws. Petrotilapiapossesses tricuspid teeth in 1997). Within the genus Pseudotropheusas it is the outer rows of both the lower and upperjaws now recognized, those species being placed in and is further distinguishedby visibility of the Metriaclimaare unique because they have a moder- teeth when the jaws are closed. The isognathus ately sloped ethmo-vomerine block and a swollen jaws of Metriaclimadistinguishes it from Genyo- rostral tip (Fig. 1), compared to P. elongatus (Fig. chromis,which is characterizedby having the 2) and P. tropheops(Fig. 3) complexes. Pseudotro- lower jaw extendingin front of the upper jaw. pheus williamsi complex differs in that when the PSEUDOTROPHEUSZEBRA COMPLEX 193

Fig. 3. Skull of Pseudotropheustropheops. mouth is closedthe lower jaw is parallelto a line triangularin outline. Scalesalong side ctenoid; from the tip of the snoutto the hypuralplate (Fig. 29-33 lateral-line scales. First gill arch with 9-14 4), while the lower jaw of Metriaclimaspecies is rakers on the ceratobranchial, 1-3 on epibranchial, angledapproximately 450 relativeto this line (Fig. and 1 between the epibranchialand ceratobranchial 5). Meyer and Foerster (1984) suggestedthat (Table 1). certainmembers of this group be includedin the Breeding males have a blue ground coloration subgenus Maylandia;however, although Maylandia laterally with 6-8 black vertical bars, very faint or was proposed as a subgenusno descriptionor no bars on posterior one-third. Although extreme- diagnosisaccompanied this suggestion.Therefore, ly rare, a few males in some populations (e.g. Maylandia,in effect, is a nomen nudumand thus Songwe Hill) exhibit a blue/black or orange/black is an invalidname. blotched coloration. Belly anterior to the pelvic Etymology.-Metriaclima, from the Greek, fins (P) black fading to light blue posteriorly; head meaningmoderately sloped to indicatethe moder- dark blue to black with a single light interorbital ately sloped head of membersof this genus. bar; gular dark gray in most populations, but yellow in some northern populations. Dorsal fin Metriaclimazebra Boulenger uniformly blue gray in most populations, with tips Diagnosis.-The moderatelysloped head, swol- of posterior rays orange in some populations and len rostral tip of the neurocranium,isognathus a continuous black horizontal stripe in some jaws, and the presenceof bicuspidteeth in the northern populations; caudalfin blue/gray; anal fin outer rows of the jaws place this speciesin Metri- blue gray with 1-5 yellow ocelli; in some popula- aclima. The absenceof any interruptedpigment in tions the proximal two-thirds of the anal fin is the pale blue dorsalfin in combinationwith black black, while the distal one-third is light gray; P2 barson a blue groundcolor distinguishthis form light blue leading edge, spine and first two rays from all other membersof the genus. black, posterior rays clear; P1 clear. Females Description.-Jawsisognathous; teeth on jawsin generally with brown cast and 1-4 faint ocelli on 2-5 rows;majority of teeth in outerrows bicuspid, anal fin. The females alternatively may be dull with a few posterior unicuspidteeth; those in blue or exhibit an orange/black blotched color- inner rows tricuspid;4-14 teeth in outer rows of ation. This blotched coloration is observed more the left lower jaw. Dorsal fin with 16-19spines frequently in females than males and appearsto be and 7-10 rays. Pectoralfins (P1)with 12-15rays; completely absent in some populations. anal fin with 3 spines except in some specimens Distribution.-Metriaclima zebra is widely from MazinziReef and Mitande Rocks, which had distributed throughout the lake. Differences in 4; analfin with 6-8 rays. Lowerpharyngeal bone meristics and morphometrics among populations 194 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

Fig. 4. X-ray of Pseudotropheuswdliamsi.

Fig. 5. X-ray of Metriadima zebra. can be attributedto clinal variation,as there is a [HL), a greater distance between the anterior great deal of overlap in the minimum polygon insertionof the dorsalfin and the anteriorinser- clustersthat are formedwhen the first principal tion of the anal fin (57.9 % HL vs. 47.0 - 57.3 % componentsof the meristicdata are plotted against HL), and more teeth rows on the lower jaw (5 vs. the shearedsecond principalcomponents of the 2-4) when compared to all other populations of M. morphometricdata (Figs. 6a, 6b, & 6c). Thus, zebra (Table 1). The lack of a more specific type thesepopulations are consideredto be conspecific, locality precluded collecting topotypes. Because even though McKayeet al. (1984)reported allelic there is no overlap when the first principal compo- frequencydifferences of the same magnitudeas nents of the meristic data are plotted against the observedbetween sympatric heterospecific popula- sheared second principal components of the mor- tions of Metriaclima. phometric data (Fig. 6c) with M. zebra and any of Discussion.-The type locality of Metriaclima the populations of M. zebra, we were first tempted zebrais Lake Nyassa and the originaldescription to restrict the name M. zebra to the single speci- was basedon a singlespecimen (BMNH 91.12.17.7) men housed in the Natural History Museum cataloged into the Natural History Museum (London) and propose a new name for the other (London). The HOLOTYPE of M. zebra has a populations. Ultimately, we chose to preserve the longer snout length (40 % HL vs. 30.7 - 39.3 % name M. zebra, because of its wide usage in both PSEUDOTROPHEUSZEBRA COMPLEX 195

0.15

* ThuambiEast Island - SongweHill A Mazinzi Reef

-0.1-\0 x x BoadzuluIsland 0~~~~~~~~~~~~~~ x CrocodileRocks -~~~~~ *~~~ MitandeRocks _ +-l zebra holotype -0.05-0.15 -|lll- - --

x x

x ~~~~~~~~~~~+ -0.1I

-0.15 - i -3 -2 -1 0 1 2 3

Factor I

Fig. 6a. Plot of the sheared second principal cmponents (morphometric data) and the first factor score (meristic data) for Metriaclimazebra populations from the uthern end of Lake Malaxwi.

0.1I

0.05

*NkicataBay as *~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~o5kn N. ofNklisataBay cli -0.05 ~~~~~~~~~~~~~~~~~~~~~~~~AChilunba 12 ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~xM zebra holotype

-0.1I

-0.15 x

-0.2 I I I -I -2 -1.5 -l -0.5 0 0.5 1 1.5 2 2.5

Factor1

Fig. 6b. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriaecli'mazebrae populations from the northern end of Lake Mala4iWi. 196 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE

0.15

0.1 A A

*0.1 A A A PA 0.05 \ A A

A A#A A*

/ A I:$* M. zebra holotype

-0.15- -3 -2 0 1 A 2 3 Factor A Fig. 6c. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriaclimazebra populationsfrom the northernand southernend of LakeMalaw'i. the scientific and popular literature and regardthe Metriaclima heteropictus (Staeck) holotype as being atypical. Diagnosis.-The moderately sloped head, swoi- Etymology.-The name zebra, from the Greek, len rostral tip of the neurocranium, isognathus meaning black-barredto note the black bars on the jaws, and the presence of bicuspid teeth in the lateral side. outer rows of the jaws place this species in Metri- SpecimensExamined.-BMNH 91.12.17.7, Lake aclima. The high number of teeth rows on the Malawi;PSU 3036, PSU 3037, 3; USNM 341908, 3; upper jaw (6) and the lower jaw (5-6) distinguishes ANSP 176182,3; MFU (Malaw'iFisheries Unit) 11, 3; this form from all other member of the genus. (63.7 - 88.4 mm), MazinziReef, Lake Malaw'i,Malawi Description.-Jaws isognathous; teeth on jaws in Africa, 7-10 m, March, 1991. PSU 3038, 10; USNM 5-6 rows; majority of teeth in outer rows bicuspid, 3; ANSP 176183,3; MFU 12,3; (56.1- 91.9mm), 341909, with a few unicuspid teeth; those in inner rows CrocodileRocks, LakeMalaw'i, Malaw'i, Africa, 2-10 m, May, 1991. PSU 3039, 9; USNM 341910, 5; ANSP tricuspid; 14-16 teeth in outer row of left lower 176184,5; MFU 13, 5; (61.2- 89.6mm), Boadzulu Island, jaw. Dorsal fin with 18 spines and 9-10 rays. Lake Mala4wi,Malawii, Africa, 1-10 m, January,1991. Pectoral fin with 13 rays; anal fin with 3 spines PSU 3040, 8; USNM 341911,4; ASNP 176185,4; MFU and 9-10 rays. Scales along side ctenoid with 32 14, 4; (55.5 - 80.0 mm), Thumbi East Island,3-7 m, lateral-linescales. First gill arch with 12 rakers on February,1991. PSU 3041, 6; USNM 341912,2; ANSP the ceratobranchialand 2 on the epibranchial, and 176186,3; MFU 15, 2; (56.2- 80.0 mm), SongweHill, 1 between the epibranchial and ceratobranchial LakeMalawi, Malawii, Africa, 3-5 m, January,1991. PSU (Table 2). 3042, 8; USNM 341913,4; ANSP 176187,4; MFU 16, 4; Distribution.-Staeck (1980) lists the type - mm), MitandeRocks (ThumbiWest Island), (57.7 90.6 locality as Thumbi West Island, Lake Malaw'i; LakeMalawii, Malawii, Africa; 3-5 m, January,1991. PSU 3043,7; USNM 341914,3; ANSP 176188,3; MFU 17,3; however, no forms which resemble the paratypes (57.0 - 84.2 mm), Nkhata Bay, Lake Malawii,Malaw'i, have ever been collected at this locality. The Africa,2-4 m, May, 1991. PSU 3044,6; USNM 341915, name, Metriaclimabeteropictus, has historically been 3; ANSP 176189,3; MFU 18, 3; (60.2- 83.6 mm), 5 km applied to a large species from Thumbi West Island north of NkhataBay, LakeMalawi, Malaw4i, Africa, 2-6 (Konings 1995); thus, the locality of this form is m, June, 1991. PSU 3045, 7; USNM 341916,3; ANSP currently unknown. 176190,3; MFU 19, 3; (61.2- 84.6mm), Chitande Rocks, MaterialExamined.-SMF (Natur-Museum und LakeMalaw4i, Mala'w4i, Africa, 2-6m, July, 1991. Forschungs-Institut Senckenberg) 15179, SMF 15180, PARATYPES. PSEUDOTROPHEUSZEBRA COMPLEX 197

Fig.7. Metriadima melabranchion,HOLOTYPE, PSU 3049, adultmale, 84.8 mm SL.

Fig. 7. Metriaclimamelabranchion, HOLOTYPE, PSU 3049, adultmale, 84.8 mm SL.

Metriaclima melabranchion Stauffer, Bowers, Kellogg and McKaye, n. sp. (Fig. 7) Pseudotrophuszebra, Ribbink et al. 1983:158(in part); Konings(1990) (in part). HOLOTYPE.-PSU 3049, adult male, 85.7 mm, Mumbo Island, Lake Malawi, Malawi, Africa, 1-5 m, May, 1991 (Figs. 7 & 8). PARATYPES.-PSU3050, 10; USNM 341921, 3; ANSP 176191, 3; MFU 24, 3; (76.8 - 91.5 mm), data as for holotype. PSU 3051, 11; USNM 341922, 3; ANSP 176192, 3; MFU 25, 3; (78.2 - 105.4 mm); Zimbawe Rocks, Lake Mala'wi,Africa, 1-3 m, May, 1991; PSU 3046, 11; USNM 341917, 3; ASNP 176193, 3; MFU 20, 3; (64.2 - 81.5 mm); Mitande Rocks, Cape MaClear, Lake Malawi, Malawi, Africa; 11-17 m; March, 1988. PSU 3047, Fig. 8. Lower pharyngeal bone of the holotype of 11; USNM 341918, 3; ASNP 176194, 3; MFU 21, Metriadima melabranchion,PSU 3049. 3; (55.3 - 88.6 mm); Domwe Island, Cape MaClear, holotype and 12-14 in paratypes; anal fin with 3 Lake Malawi, Malawi, Africa; 1-3 m, May, 1991. spines, 8 rays in holotype and 7-9 in paratypes. PSU 3048, 11; USNM 341919, 3; ASNP 176195, 3; Lower pharyngeal bone triangular in outline (Fig. MFU 22, 3; (64.3 - 86.6 mm); Namalenge Island, 8). Scales along side holotype with 32 Lake Malawi, Malaw'ri,Africa, 3-9 m, July, 1991. ctenoid; lateral-line scales and 31-34 in paratypes. First gill Diagnosis.-The moderately sloped head, swol- arch with 11-14 rakers on the ceratobranchial, 2-5 len rostral tip of the neurocranium, isognathous on epibranchial, and 1 between the epibranchial jaws, and the presence of bicuspid teeth in the and ceratobranchial 3). outer rows of the jaws place this species in Metri- (Table Breeding males have a dark blue ground color- aclima. The extension of the lateral vertical bars ation laterally with 6-10 vertical bars; breast with into the dorsal fin discriminates M. melabranchion a reddish cast on some specimens and tends to be from M. zebra. more pronounced in the population from Description.-Jaws isognathous (Fig. 7); teeth on Zimbawe Rocks; head dark blue to black with 1-2 jaws in 2-5 rows; majority of teeth in outer rows blue interorbital bars, gular blue/black. Dorsal fin bicuspid, with a few posterior unicuspid teeth; blue with black vertical bars from side extending those in inner rows tricuspid; 9 teeth in outer row into dorsal fin; some individuals have orange spots of left lower jaw of holotype, 3-12 in paratypes. in the posterior three membranes. Caudal fin with Dorsal fin with 18 spines in the holotype and black rays and blue/gray membranes; anal fin 16-19 in paratypes; dorsal-fin rays 8 in holotype gray/black with 3-5 yellow ocelli; P2 black with and 8-10 in paratypes. Pectoral fin with 12 rays in 198 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE

0.15

+

0.1

0.05 . MitandeRocks M*A Ao zebra- holotype A MumboIsland A /;xc : ) x NamalengeIsland A 1 t I o- A * o/ * \ 3 xDomwe Island

\ "' \,1 * x x o ZimbaweRocks \^< - \! X X + L+M.heteropictus -paratypes

-0.05 \ x X

-0.1.I

-0.15 -I I I - I -3 -2 1 0 1 2 3 4

Fig. 9. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriaclima melabranchion populations and Metriaclima heteropictus PARATYPES. white leading edge; P1 clear. Females brown/green Metriaclima chrysomallos Stauffer, Bowers, laterally with blue/green highlights and 5-7 vertical Kellogg and McKaye, n. sp. (Fig. 10) bars; head dorsally dark brown with 1-2 green HOLOTYPE.-PSU 3052, adult male, 84.8 mm, interorbital bars; gular blue/white with scattered Mumbo Island, Lake Malaw'i,Malaw'i, Africa, 1-5 micromelanophores; cheek and opercular light m, May, 1991 (Figs. 10 & 11). blue. Dorsal fin blue/gray to brown; caudal fin PARATYPES.-PSU 3053, 7; USNM 34192, 4; brown with yellow tips in some individuals; anal ANSP 176196, 4; MFU 23, 4; (61.0 - 89.6 mm); fin gray to brown with 0-4 yellow ocelli; P2 data as for holotype. blue/white leading edge, first two rays and mem- Diagnosis.-The moderately sloped head, swol- branes black with posterior ones clear; P1 clear. len rostral tip of the neurocranium, isognathus Distribution.-Metriaclima melabranchionoccurs jaws, and the presence of bicuspid teeth in the at Mitande Rocks (Thumbi West Island), Domwe outer rows of the jaws place this species in Metri- Island, Mumbo Island, Zimbawe Rocks, and aclima. The continuation of the vertical bars on Namalenje Island. The minimum polygon clusters the side of fish into the dorsal fin separates this formed when the first principal components of the species from M. zebra. The gold highlights on the meristic data are plotted againstthe shearedsecond light blue ground coloration distinguishes this principal components of the morphometric data species from M. melabranchion. for the populations of M. melabranchionappear to Description.-Jaws isognathous (Fig. 10); teeth form a clinal grade (Fig. 9). The fact that M. on jaws in 2-3 rows; majority of teeth in outer melabranchion is sympatric with M. zebra at rows bicuspid, with a few posterior unicuspid Mitande Rocks supports the conclusion that these teeth; those in inner rows tricuspid; 13 teeth in two taxa are heterospecific. outer row of left lower jaw of holotype, 6-13 in Etymology.-The name malabranchionfrom the paratypes. Dorsal fin with 17 spines im the holo- Greek, meaning black fin to note the extension of type and 17-18 in paratypes; dorsal-fin rays 9 in the lateral black bars onto the dorsal fin. holotype and 8-10 in paratypes. Pectoral fin with 13 rays; anal fin with 3 spines and 6-8 rays. Lower pharyngeal bone triangular in outline (Fig. 11). Scales along side ctenoid; holotype with 32 lateral PSEUDOTROPHEUSZEBRA COMPLEX 199

Fig. 10. Metriadima chrysomallos,HOLOTYPE, PSU 3052, adult male, 85.7 mm SL.

Distribution.-Metriaclima cbrysomallos is en- demic to Mumbo Island. Etymology.-The name chrysomallos,from the Greek, meaning gold-fleeced to note the gold highlights on the lateral side in both sexes.

Metriadima phaeos Stauffer, Bowers, Kellogg and McKaye, n. sp. (Fig. 12) HOLOTYPE.-PSU 3054, adult male, 85.1 mm, Cobue, Lake Malaw'i,Mozambique, Africa, 1-5 m, March, 1996 (Figs. 12 & 13). PARATYPES.-PSU3055, 5; USNM 341923, 1; ANSP 176197, 1; MFU 26, 1; (68.3 - 86.5 mm), data as for holotype. Diagnosis.-The moderately sloped head, swol- Fig. 11. Lower pharyngeal bone of the type specimen of len rostral tip of the neurocranium, isognathous Metriadiima PSU 3052. chbysomat1os, jaws, and the presence of bicuspid teeth in the scales and 31-33 in paratypes. First gill arch with outer rows of the jaws place this species in Metri- 11-14 rakers on ceratobranchial, 2-3 on epibranch- aclima. Metriaclimaphaeos superficially resembles ial, and 1 between the epibranchial and cerato- M. zebra, but the dusky submarginal band in the dorsal fin of M. phaeos distinguishes it from all branchial (Table 4). Breeding males have a light blue ground color- other BB members of the genus. The minimum ation laterally with gold highlights and 6 gray polygons formed by plotting the first principal vertical bars; ventral portion fades to pale components of the meristic data against the sheared second of blue/green; head gray with 2 blue interorbital bars, principal components the mor- green opercle, blue cheek, and gray gular. Dorsal phometric data (Fig. 14) illustrate that the mini- formed M. fin pale blue with vertical bars extending into it; mum polygons by phaeos do not over- with those formed posterior 4 membranes are rust/orange distally. lap by any of the populations Caudal fin light blue/green; anal fin blue with 3-5 of M. zebra. yellow ocelli; P2 with light blue leading edge, spine Description.-Jaw isognathous (Fig. 12);teeth on and first 2 membranes black, remainder clear; P1 jaws in 3-4 rows; majority of teeth in outer rows clear. Females have a pale green/beige ground bicuspid, with a few posterior unicuspid teeth; in coloration with gold highlights and 6 gray vertical those inner rows tricuspid; 12 teeth in outer row of left lower of in bars; head gray with 2 blue interorbital bars, green jaw holotype, 11-13 para- Dorsal fin with 17 in opercle and cheek, and white gular. types. spines the holotype and 17-18 in paratypes; dorsal-fin rays 9. Pectoral 200 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

Fig.12. Metriadimaphacos, HOLOTYPE, PSU 3054, adultmale, 85.1 mm SL.

Fig. 12. Metriaclimapbaeos, HOLOlYPE,PSU 3054, adult male, 85.1 mm SL. with trailing yellow edge; anal fin light blue/purplebecoming dusky then white in posteri- or distal portion with 3 yellow ocelli; P2 blue/white leadingedge with distinct, bold sub- marginalblack band, remainderhyaline; P2 dear. Femalesdark blue/gray with 6-8 dark gray bars; ventralone-fifth of flankyellow/brown; head dark brown with faint blue/brown interorbitalbar; cheek and operculumyellow/brown; dorsal fin yellow/brown highlightson a light blue ground; very faintthin duskysubmarginal band with white lappetsand yellow/brown tips of serialelements; rayedportion with yellow/brownspots; caudal fin with faint light blue membranes and yel- low/brown rays;anal fin blue/graywith transpar- ent edge and 3 faint yellow/brown ocelli; P2 with white leading edge and black submarginalband with yellow/brown membranes;P1 clear. Fig. 13. Lower pharyngeal bone of the type specirnen of Distribution.-Metriaclimaphaeos was collected Metriadima phaeos, PSU 3054. only at Cobue,Mozambique, but may havea more extensive distribution along the Mozambique fin with 14 rays in holotype and 14-15 in para- coastline. types; anal fin with 3-4 spines and 8 rays. Lower Etymology.-Thename phaeos, from the Greek, pharyngeal bone triangular in outline (Fig. 13). meaningdusk to note the duskysubmarginal band Scales along side ctenoid; holotype with 31 later- presentin the dorsalfin of both sexes. al-line scales and 31-32 in paratypes. First gill arch with 11-13 rakers on the ceratobranchial, 2-4 on Metriaclimacyneusmarginatus Stauffer, Bowers, the epibranchial, and 1 between the epibranchial Kelloggand McKaye,n. sp. (Fig. 15) and ceratobranchial (Table 5). HOLoTYPE.-PSU 3056, adultmale, 73.9 mm, Breeding males dark blue ground coloration Nkhota khota, LakeMalaw'i, Malaw'i, Africa, 1-3 laterally with 6-8 vertical bars; belly white; head m, February, 1995 (Figs. 15 &16). dark blue with iridescent blue interorbital bar; PARATYPES.-PSU3057, 11; USNM 341924, 3; fin gular gray/white; dorsal light blue proximally, ANSP 176198, 3; MFU 27, 3; (51.9 - 80.4 mm), becoming lighter distally; thin dusky submarginal dataas for holotype. band and white lappets; rayed portion of the Diagnosis.-The moderatelysloped head, swol- dorsal fin is yellow/white in distal one-third; len rostral tip of the neurocranium,isognathus caudal fin deep blue proximally, light blue distally jaws, and the presenceof bicuspid teeth in the PSEUDO TROPHEUS ZEBRA COMPLEX 201

0.2

0.15

l 0.1 YU

o zebra - Northern 0.050.05 U ~~~~~~~~~~~~~~~~~~~~~~~~~M /C~ .AI zebra - Southern 0 % ir ~~~~~~~~~~~~~~~~AAl. zebra holotvpe Wo *0 */C .* X M phaeos no 0 ii *4 I^ e > | xXM cyneaumaorinatus 0 ~ ~ ~ ~

-0.05 U U

A

x -0.15 -3 -2 -1 0 1 2 3 FactorI

Fig. 14. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metrtaclimazebra populations from the northern and southern end of Lake Malaw'i,as well as, Metriaclimaphaeos and Metriaclimacyneusmarginatus.

Fig. 15. Metriaclimacyneusmarginatus, HOLOTYPE, PSU 3056, adult male, 73.9 mm SL. outer rows of the jaws place this species in Metri- Description.-Jaws isognathous (Fig 15);teeth on aclima. The blue marginal band and brown jaws in 2-3 rows; majority of teeth in outer rows lappets distinguishes it from other members of bicuspid, with a few posterior unicuspid teeth; Metriaclima. The minimum polygons formed by those in inner rows tricuspid; 12 teeth in outer plotting the first principal components of the row of left lower jaw of holotype, 8-12 in para- meristic data against the sheared second principal types. Dorsal fin with 17 spines in holotype, 17-18 components of the morphometric data (Fig. 14) in paratypes; dorsal-fin rays 9 in holotype, 8-10 in illustrate that the minimum polygon formed by M. paratypes. Pectoral fin with 13 rays in holotype, cyneusmarginatus is significantly different with 11-15 in paratypes; anal fin with 3 spines; anal-fin those formed by M. phaeos. rays 8 in holotype, 7-8 in paratypes. Lower phar- 202 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

Distribution.-Metriaclima cyneusmarginatuswas collected only at Nkhota khota. Etymology.-The name cyneusmarginatus,from the Greek, meaning blue margin to note the blue marginal band in the dorsal fin.

Metriadima benetos Stauffer,Bowers, Kellogg and McKaye, n. sp. (Fig. 17) Pseudotropheuszebra 'mazinzi', Ribbink et al. 1983:160; Pseudotropheuszebra Konings 1990: 442 (in part). HOLOTYPE.-PSU 3058, adult male, 81.4 mm, Mazinzi Reef, Lake Malawi, Malawi, Africa, 10-15 m, January, 1991 (Figs. 17 & 18). PARATYPES.-PSU 3059, 9; USNM 349125, 3; ANSP 176199, 3; MFU 28, 3; (59.7 - 81.4 mm), data as for holotype. Diagnosis.-The moderately sloped head, swol- len rostral tip of the neurocranium, isognathus Fig. 16. Lower pharyngeal bone of the type specimen of jaws, and the presence of bicuspid teeth in the Metriadima cyneusmarginatus,PSU 3056. outer rows of the jaws place this species in Metri- yngeal bone triangular in outline (Fig. 16). Scales aclimna. The pale blue ground color with the along side ctenoid; holotype with 32 lateral-line absence of distinct black vertical bars distinguishes scales and 31-33 in paratypes. First gill arch with M. benetos (Fig. 19a) from all other members of 10-12 rakers on the ceratobranchial, 2-3 on epi- this genus, with the exception of Metriaclima branchial, and 1 between the epibranchial and callainos (Fig. 19b). Metriaclimabenetos females are ceratobranchial (Table 6). brown with green highlights, while M. callainos Breeding males have a blue ground coloration females are either light blue or white (Fig. 19c). In laterally with 5-6 gray vertical bars and a white general, M. benetos have fewer dorsal-fin spines belly. Dorsal fin blue/gray with a blue marginal (mode - 17) than M. callainos (mode - 18) (Table 7). band and brown lappets; caudal-fin rays gray, The first principal components of the meristic data membranes blue, with brown tips; anal fin blue were plotted against the sheared second principal with 2 yellow ocelli; P2 with light blue leading components of the morphometric data. There was edge, spine and first 2 membranes black, posterior no overlap in the minimum polygon clusters rays and membranes clear; P1 clear. Females formed by M. benetos with that formed by M. similarly colored but not as intense and they lack callainos (Fig. 20). Variables that had the highest ocelli on the anal fin. loadings on the sheared second principal compo-

Fig. 17. Metriadima benetos, HOLOTYPE, PSU 3058, adult male, 81.4 mm SL. PSEUDOTROPHEUSZEBRA COMPLEX 203

Fig. 19a. Metriaclima benetosfrom Mazinzi Reef.

Fig. 18. Lower pharyngeal bone of the type specimen of Metriadima benetos, PSU 3058. nents were horizontal eye diameter (-0.439), caudal peduncle length (0.409), and preorbital depth (0.313); while those with the highest loadings on i : the principal components of the meristic data were number of dorsal spines (0.227), number of P1 rays Fig. 19b. Metriadima callainos male from Mitande (0.215), and number of teeth on the left side of the Rocks. lower jaw (0.216). Description.-Jaws isognathous (Fig. 17); teeth on jaws in 3-4 rows; majority of teeth in outer rows bicuspid, with a few posterior unicuspid teeth; those in inner rows tricuspid; 9 teeth in outer row of left lower jaw of holotype, 9-13 in paratypes. Dorsal fin with 16 spines in holotype - , -~~~~~~~~~~~~*; and 16-18 in paratypes; dorsal-fin rays 9 in holo- type, 8-10 in paratypes. Pectoral fin with 13 rays in holotype and 12-14 in paratypes; anal fin with 3 spines; anal fin with 7 rays in holotype and 6-8 in paratypes. Lower pharyngeal bone triangularin Fig. 19c. Metriadiima callainos female, white morph, outline (Fig. 18). Scales along side ctenoid; holo- from Mitande Rocks. type with 29 lateral-line scales and 29-32 in para- types. First gill arch with 9-12 on the cerato- tween spine and first ray, remainderof the mem- branchial, 3-4 on epibranchial, and 1 between the braneslight blue with 2 yellow ocelli; P2 with yellow raysand epibranchial and ceratobranchial (Table 7). light transparentblue membranes; Breeding males have a light blue ground color- P1yellow raysand dear membranes.Females have ation laterally that is darker dorsally and fades to a brown ground colorationwith green and blue white ventrally. For the most part, no vertical highlightsand 6-8 faint verticalbars; head brown bars are visible, but on occasion, faint bars are with bluehighlights below eye andon cheek,gular noticeable immediately after preservation. Head white with blue cast. Dorsal fin uniformly or- blue/gray, cheek light blue, white gular. Some ange/brown;caudal fin orange/brown;anal fin larger males in the wild have yellow gulars. transparentbrown with no ocelli;P2 brown with Although rarely seen in wild specimens, alpha blackleading edge; P1 clear. males held in aquaria will sometimes develop a Distribution.-Metriaclimabenetos is endemicto bright yellow gular. Dorsal fin proximally blue Mazinzi Reef in the southeasternarm of Lake gray with green highlights and white marginal Malawi. namebenetos, from the Greek, band; caudal fin proximally blue/gray with white Etymology.-The membranes with a yellow cast on distal two-thirds; meaningblue to note the light blue groundcolor- anal fin with white membrane with yellow cast be- ation of breedingmales. 204 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE

0.12 ; 0.17

0.08 t . v

0.06

c 0.04

0.020.0 *AMLcallanos mM benetos 0

-0.02

-0.04

-0.06-

-0.08

-0.1 I -2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 Factor I

Fig. 20. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriaclima benetos and Metriaclimacallainos.

Metriaclima callainos (Stauffer and Hert) Hert 1992); teeth on jaws in 3-4 rows; majority of Diagnosis.-The moderately sloped head, swol- teeth in outer rows bicuspid, with a few posterior len rostral tip of the neurocranium, isognathous unicuspid teeth; those in inner rows tricuspid; jaws, and the presence of bicuspid teeth in the 13-17 teeth in outer row of left lower jaw. Dorsal outer rows of the jaws place this species in Metri- fin with 17-18 spines and 8-9 rays. Lower pharyn- aclima. The pale blue ground color with the geal bone triangular in outline (Stauffer and Hert absence of distinct black vertical bars distinguishes 1992). Scales along side ctenoid with 30-32 later- M. callainos (Fig. 19b) from all other members of al-line scales. First gill arch with 11-12 rakers on this genus, with the exception of M. benetos(Fig. the ceratobranchial, 1-3 on epibranchial, and 1 19a). M. callainos females are either light blue or between the epibranchial and ceratobranchial white (Fig. 19c), while Metriaclimabenetos females (Table 7). are brown with green highlights. In general, M. Breeding males have a light blue ground color- callainos have a greater number of dorsal-finspines ation with green highlights on dorsal one-third; (mode - 18) than M. benetos (mode - 17) (Table 7). head light blue with green highlights dorsally, with The first principal components of the meristic data white gular and black opercular spot. Occasional- were plotted against the sheared second principal ly, males will be either white or blue/white components of the morphometric data for M. blotched. Dorsal fin light blue; caudal fin light benetosand M. callainos. There was no overlap in blue; anal fin white proximally, light blue distally, the minimum polygon formed by M. benetoswith with 1-3 yellow ocelli; P2 with black rays and clear that formed by M. callainos (Fig. 20). Variables membranes; Pi clear with white leading edge. A that had the highest loadings on the shearedsecond few females are similarly colored, but not as principal components were horizontal eye diameter intense and lack ocelli; some females white overall (-0.439), caudal peduncle length (0.409), and pre- (Fig. 19c; Staufferand Hert 1992), and an occasion- orbital depth (0.313); while those with the highest al blue/white blotched female is observed loadings on the principal components of the (Schroder 1980). meristic data were number of dorsal spines (0.227), Distribution.-Metriaclima callainosisindigenous number of P1 rays (0.215), and number of teeth on to the Nkhata Bay area, but has been introduced the left side of the lower jaw (0.216). to Likoma, Namalenje, and Thumbi West islands Description.-Jaws isognathous (Stauffer and (Ribbink et al. 1983; Stauffer and Hert 1992). In PSEUD07ROPHEUSZEBRA COMPLEX 205

Nkhata Bay, M. callainos is sympatric with M. Thumbi West Island has diverged morphologically zebra. Schroder (1980) concluded that these two from the Nkhata Bay population, and specimens forms were heterospecific, because he observed have been observed that may be hybrids between assortative mating. The introduced population at the M. zebra and the introduced population of M.

0.08

0.06 *

0.04 / e 0.02 e U U 0 * / m

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-0.06

-0.08 -

-0.1

-0.12

-0.14 - -2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 Factor 1 Fig. 21. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriacdimaxanstomachus.

1.2

U * U 1 - *

t : .~~~~~~~U

X~ 0.8-

s ^ v~~~~~~~~~~~~~~~~~~~~~~~~~~Nakatenga Island] 0.6 -* KanjedzaIslandj

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0 2 4 6 8 10 12 Head Length

Fig. 22. Plot of the horizontal eye diameter/head length ratio versus head length for two allopatric populations of Metriaclimaxanstomachus, indicating allometric growth. 206 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE callainos (Stauffer and Hert 1992). There is an principal components of the morphometric data undescribed species in which all males and females were snout length (-0.421), caudal peduncle length are white at Ruarwe, which we consider to be (0.378), and preorbital depth (-0.304). There is heterospecific and will be describedin a subsequent clearly non-overlapping morphometrics for the paper. two populations of M. xanstomachus. The speci- Material Examined.-PSU 2542.1, PSU 2542.2, mens from Nakantenga Island are all larger than MFU 2, USNM 322426; Nkhata Bay, Lake those from Kanjedza Island; thus, differences Malawi, Malawi, Africa, 3-4 m, May, 1988. observed may be attributed to allometric growth. For example, when horizontal eye diameter/head Metriaclima xanstomachus (Stauffer and Boltz) length is plotted against head length allometric Diagnosis.-The moderately sloped head, swol- growth is indicated (Fig. 22). There are certainly len rostral tip of the neurocranium, isognathous links between the Metriaclimapopulations at the jaws, and the presence of bicuspid teeth in the Maleri Islands with those at Kanjedza Island. For outer rows of the jaws place this species in Metri- example M. barlowi occurs at both localities and aclima. The light blue body coloration with 6 there is a form with a red dorsal fin at both vertical bars coupled with a bright yellow gular islands, although in the latter case, we regard the and blue dorsal fin delimits M. xanstomachusfrom two red-top forms at distinct species (see below). all other known Metriaclimaspecies. It may be that the two populations of M. xanstom- Description.-Jaws isognathous (Stauffer and achusrepresent semi-species (sensu Mayr) or are, in Boltz 1989: 9); teeth on jaws in 3-5 rows; majority fact, heterospecific. of teeth in outer rows bicuspid, with a few posteri- MaterialExamined.-PSU 3060, 5; Nakantenga or unicuspid teeth; those in inner rows tricuspid; Island, Lake Mala'i, Malaw'i, Africa; 3-7 m, 9-11 teeth in outer row of left lower jaw. Dorsal March, 1991; PSU 3061, 20; Kanjedza Island, Lake fin with 16-18 spines and 8-10 rays. Pectoral fin Malaw'i,Mala'wi, Africa; 1-4 m, March, 1988. with 12-14 rays; anal fin with 3 spines and 7-8 rays. Lower pharyngeal bone triangular in outline Metriaclima pyrsonotos Stauffer,Bowers, Kellogg (Stauffer and Boltz 1989: 9). Scales along side and McKaye, n. sp. (Fig. 23) ctenoid with 30-33 lateral-linescales. First gill arch Pseudotropheuszebra 'red dorsal', Ribbink et al. 1983:162; with 9-12 rakers on ceratobranchial, 2-3 on epi- Konings1990: 454 (in part). branchial, and 1 between the epibranchial and HOLOTYPE.- PSU 3062, adult male, 74.1 mm, ceratobranchial (Table 8). Nakantenga Island (Maleri Islands), Lake Malawri, Breeding males with light blue ground color and Africa, 2-4 m, March, 1991 (Figs. 23 & 24). 6 vertical gray bars; head dark blue with 1 light PARATYPES.- PSU 3063, 8; USNM 341926, 4; blue interorbital bar, yellow gular and branchio- ANSP 176200, 3; MFU 7, 4; (56.9 - 76.6 mm), data stegal rays. Dorsal fin blue with gray flecks; anal as for holotype. fin blue with 3-5 yellow ocelli; P2 fins black Diagnosis.-The moderately sloped head, swol- anteriorly fading to yellow/brown posteriorly; P1 len rostral tip of the neurocranium, isognathous black (Stauffer and Boltz 1989). jaws, and the presence of bicuspid teeth in the Distribution.-Metriaclima xanstomachus was outer rows of the jaws place this species in Metri- originally thought to be endemic to the Maleri aclima. The red dorsal fin in combination with Islands (Ribbink et al. 1983, Stauffer and Boltz black bars on a blue ground color distinguish this 1989); however, analysis of shape and color pat- form from all other members of the genus except terns showed that the population inhabiting for Metriaclimasandaracinos (Mpanda Island and Kanjedza Island is also M. xanstomachus.The first Kanjedza Island) and Metriaclima emmiltos principal components of the meristic data were (Chilumba), which also have red dorsal fins. plotted against the sheared second principal com- Metriaclimapyrsonotosdiffers from all the other red ponents of the morphometric data (Fig. 21). The top BB forms in that the vertical black lateral bars minimum polygons formed by the populations infiltrate the proximal portion of the dorsal fin. In inhabiting Nakantenga Island (one of the three general, M. pyrsonotoshas fewer teeth in the outer Maleri islands) and Kanjedza Island were signifi- row of the left lower jaw (mode - 7) than M. cantly (P < 0.05) different. Variables that had the emmiltos from Chilumba (mode - 9), and M. highest loadings on the principal components of sandaracinos from Kanjedza (mode - 10), or the meristic data were teeth rows in the upper jaw Mpanda (mode - 12) islands (Table 9). (0.248), teeth rows in the lower jaw (0.244), and Description.-Jaws isognathous (Fig. 23); teeth number of scale rows on cheek (0.238), while those on jaws in 2-3 rows; majority of teeth in outer with the highest loadings on the sheared second rows bicuspid, with a few posterior unicuspid teeth; those in inner rows tricuspid; 7 teeth in out- PSEUDOTROPHEUSZEBRA COMPLEX 207

Fig. 2. Metiadimpyrsnotos,HOLOT_E, PU 306, adul male 74.1mm SL

Fig. 23. Metriaclimapyrsonotos, HOLOTYYPE, PSU 3062, adultmale, 74.1 mm SL.

to the caudal peduncle; belly anterior to the P2's black fadingto light blue posteriorly;head black with 2 light interorbitalbars; dorsal fin red/orange with verticallateral bars of side sometimespene- tratingthe fin; light blue lappets;caudal fin blue with some specimenshaving an orangecast; anal fin blue with 1-4 yellow ocelli; pelvic fins black with white leading edges. Females blue/gray laterally with faint barring. Some individuals almostentirely black; head blackwith gray inter- orbitalbars; dorsal fin with brown cast; anal fin and pelvic fins black. Distribution.- Endemicto the Maleriislands where it has been observedat both Nakantenga andMaleri islands. Ribbinket al. (1983)postulat- Fig. 24. Lower pharyngeal bone of the type specimen of ed that it was originallyrestricted to Metriaadimapyrsonotos, PSU 3062. Nakantenga Island,but was introducedto MaleriIsland. The er row of left lower jaw of holotype, 4-10 in minimum polygon clusters formed by the M. paratypes. Dorsal fin with 17 spines in the holo- pyrsonotosand M. sandaracinosinhabiting Mpanda type and 16-18 in paratypes; dorsal-fin rays 8 in andKanjedza islands (see below) were significantly holotype and 8-10 in paratypes. Pectoral fins with (P <0.05) different(Fig. 25). Variablesthat had 13 rays in holotype and 12-14 in paratypes;anal fin the highestloadings on the shearedsecond princi- with 3 spines and 7 rays in holotype and 6-8 in pal componentswere snout length (0.473), pre- paratypes. Lower pharyngeal bone triangular in orbitaldepth (0.373),and caudalpeduncle length (-0.332),while those the outline (Fig. 24). Scales along side ctenoid; holo- having highestloadings on type with 31 lateral-line scales and 30-32 in para- the meristicdata were numberof teeth on the left types. First gill arch with 10-14 rakers on the lower jaw (-0.366),number of gill rakerson the ceratobranchial,2-3 on epibranchial,and 1 between ceratobranchial(0.308), and numberof gill rakers on the epibranchial the epibranchial and ceratobranchial (Table 9). (0.273). Breeding males have a light blue ground color- Etymology.- The name pyrsonotos,from the ation laterally with 5-6 black vertical bars anterior Greek,meaning red-backed to note the red dorsal fin. 208 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE

0.2

0.15

A A 0.1A

on /X- AeA/ A^xX ~ - ~!~ Al. rtos j

xn \ \2 e - *~~~~~~~~~~~~~~~~~~~~~~AM sandaracino0s- Kan?jediza , O~/X \iK~ ~ ~~~x-; x w w AXl'.sandaracino0s -Alpandi

-ni 0.15

-0.15 - I I o A emmiltos -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 Factor 1

Fig. 25. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriaclima Xyrsonotos,Metriaclima sanlaracinos and Metriaclimaemmiltos.

Metriaclima sandaracinos Stauffer, Bowers, Kel- mean snout length expressed as percent HL of M. logg and McKaye, n. sp. (Fig. 26) sandaracinos(Kanjedza Island mode - 34.6, Mpanda Pseudotropheuszebra 'red dorsal', Konings 1990: 454 (in Island mode - 34.4) differs from that found in M. part). pyrsonotos(32.2) and from M. emmiltos Chilumba HOLOTYPE.-PSU 3064, adult male, 70.9 mm, (mode - 32.9). MpandaIsland (Nkudzi Bay), Lake Malawi, Africa, Description.-Jaws isognathous (Fig. 25); teeth 1-3 m, February 1991, (Figs. 26, 27). on jaws in 3-4 rows; majority of teeth in outer PARATYPES.- PSU 3065, 7; USNM 341927, 4; rows bicuspid, with a few unicuspid posterior ANSP 176201, 4; MFU 9, 4; (55.0 - 73.1 mm); data teeth; those in inner rows tricuspid; 9 teeth in as for holotype. PSU 3066, 7; USNM 341928, 4; outer row of left lower jaw of holotype, 7-13 in ANSP 176202, 4; MFU 8, 4; (54.0 - 69.0 mm); paratypes;dorsal-fin rays 9 in holotype and 8-10 in Kanjedza Island, Lake Mala'wi, Mala'w'i,Africa, paratypes. Pectoral fins with 13 rays in holotype January, 1991. and 12-17 in paratypes; anal fin with 3 spines and Diagnosis.-The moderately sloped head, swol- 7 anal rays in holotype and 6-8 in paratypes. len rostral tip of the neurocranium, isognathous Lower pharyngeal bone triangular in outline (Fig. jaws, and the presence of bicuspid teeth in the 26). Scales along side ctenoid; holotype with 32 outer rows of the jaws place this species in Metri- lateral-linescales and 29-32 in paratypes. First gill aclima. The red/orange dorsal fin in combination arch with 8-13 rakers on the ceratobranchial, 2-3 with black bars on a blue ground color distinguish on epibranchial, and 1 between the epibranchial this form from all other members of the genus and ceratobranchial (fable 9). except for the red dorsal BB forms inhabiting Laterally, the dorsal one-half of breeding males Chilumba and Nakantenga islands. The vertical has a blue ground color; ventral one-half is or- black lateral bars do not extend imto the dorsal fin ange/brown ground color; 6-7 black vertical bars. as they do in M. prysonotos. In general, Metri- Head black with two light blue interorbital bars aclima sandaracinos, has a greater number of teeth and green highlights on opercle; gular in the outer row of the left lower jaw (Kanjedza brown/orange. Dorsal fin red with scattered blue Island mode - 10; Mpanda Island mode - 12) than highlights in membranes between spines; caudalfin M. pyrsonotos (mode - 7) from Nakantenga Island brown fading to red posteriorly; anal fin blue/gray or M. emmiltos from Chilumba (mode - 9). The with 3 yellow ocelli; pectoral-fin spine and first ray PSEUDOTROPHEUSZEBRA COMPLEX 209

Fig. 26. Metriadima sandaracinos,HOLOTYPE, PSU 3064, adult male, 70.9 mm SL.

rakers on the epibranchial (0.273). Etymology.-The name sandaracinos, from the Greek, meaning orange-colored to note the orange I 5A coloration of the dorsal fin.

Metriaclima emmiltos Stauffer, Bowers, Kellogg and McKaye, n. sp. (Fig. 28) Pseudotropheuszebra 'red dorsal',Konings 1990:454(in part). HOLOTYPE.-PSU 3067, adult male, 84.1 mm, Mpanga Rocks, Chilumba, Lake Mala'wi,Mala -wi, Africa, 1-3 m, April, 1991 (Figs. 28 & 29). PARATYPES.-PSU3068, 7; USNM 341929, 4; ANSP 176203, 4; MFU 10, 4; (59.3 - 84.2 mm); data as for holotype. Diagnosis.-The moderately sloped swol- Fig. 27. Lower pharyngeal bone of the type specimen of head, Metriadima sandaracinos,PSU 3064. len rostral tip of the neurocranium, isognathous jaws, and the presence of bicuspid teeth in the black, remainderred/orange; pelvic fins clear. outer rows of the jaws place this species in Metri- Femalesbrown laterallywith 7 darkvertical bars. aclima. The red/orange dorsal fin in combination Head brown. Dorsal fin brown with orange with black bars on a blue ground color distinguish lappets; caudal fin brown/black; anal fin this form from all other members of the genus brown/black with posteriad distal portion of except for M. pyrsonotosand M. sandaracinos,from membrane orange, no ocelli; pectoral-finrays Nakantenga and Kanjedza and Mpanda islands, black, membranesclear; pelvic-fin spine and first respectively. The vertical black lateral bars do not ray black, remainderdark orange. extend into the dorsal fin as they do in M. pyrso- Distribution.-Occupies the rocky shores of notos. In general, M. emmiltos has fewer teeth in KanjedzaIsland and ChiromboBay, and Mpanda the outer row of the left lower jaw (mode - 9), Islandand the rock outcroppingsof Nkudzi Bay. than the populations of M. sandaracinos. The The minimum polygon clusters formed by M. mean snout length expressed as percent HL of M. sandaracinos,M. pyrsonotosand M. emmiltoswere emmiltos (34.9) differs from that found in M. significantly(P < 0.05)different (Fig. 25). Variables sandaracinos(Kanjedza Island mean - 34.6, Mpanda that hadthe highestloadings on the shearedsecond Island mean - 34.4). The minimum polygon principalcomponents were snout length (0.473), clusters formed by M. emmiltos, M. sandaracinos preorbital depth (0.373), and caudal pedunde and M. pyrsonotos were significantly (P <0.05) length (-0.332),while those having the highest different (Fig. 25). Variables that had the highest loadingson the meristicdata were number of teeth loadings on the sheared second principal compo- on the left lowerjaw (-0.366),number of gill rakers nents were snout length (0.473), preorbital depth on the ceratobranchial(0.308), and numberof gill (0.373), and caudal peduncle length (-0.332), while 210 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

Fig. 28. Metriadima enmmitos,HOLOTYPE, PSU 3067, adultmale, 84.1 mm SL.

type with 31 lateral-line scales and 30-32 in para- types. First gill arch with 10-14 rays on the ceratobranchial,2-3 on epibranchial,and 1 between

E - *s~~AA the epibranchial and ceratobranchial (Table 9). Laterally, breeding males are dark blue dorsally, fading to light blue ventrally with 7 dark vertical bars. Head dark blue with two light blue interor- bital bars; green highlights on cheek and opercle with dark blue opercular spot; gular pale blue. Dorsal fin orange with light blue lappets; caudal fin orange rays with blue membranes; anal fin dark blue/gray anteriorly fading to pale blue posteriorly with 6-7 ocelli; pectoral fins with black rays and clear membranes; spine and first two rays of pelvic fin black, remainder clear. Some females similarly colored, but flecked with green highlights and pale green interorbital bars; some females with brown Fig. 29. Lower pharyngeal bone of the type specimen of ground color; no ocelli on anal fin. Metriaclimaemmiltos, PSU 3067. Distribution.-Inhabits Mpanga Rocks at Chilumba. those havingthe highestloadings on the meristic Etymology.-The name emmiltos, from the data were numberof teeth on the left lower jaw Greek, meaning red-tingedto note the color of the (-0.366), numberof gill rakerson the ceratobranch- dorsal fin. ial (0.308),and numberof gill rakerson the epi- branchial(0.273). Metriaclima mbenjii Stauffer,Bowers, Kellogg and Description.-Jaws isognathous (Fig. 28); McKaye, n. sp. (Fig. 30) teeth on jaws in 2-3 rows; majorityof teeth in Pseudotropheus'zebra mbenji', Konings 1990: 452. outer rows bicuspid,with a few unicuspidposteri- HOLOTYPE.-PSU 3069, adult male, 88.0 mm, or teeth; those in inner rows tricuspid;9 teeth in Mbenji Island, Lake Mala'wi, Malawi, Africa, outer row of left lower jaw of holotype, 8-12 in January, 1991 (Figs. 30 & 31). paratypes. Dorsal fin with 17 spinesin the holo- PARATYPES.-PSU3070, 3; USNM 341930, 2; type and 17-18in paratypes;dorsal-fin rays 9 in ANSP 176204, 2; MIFU 29, 2; (72.4 - 88.3 mm) holotypeand 8-10in paratypes.Pectoral fins with data as for holotype. PSU 3071, 10; (71.6 - 81.9 13 raysin holotypeand 12-13in paratypes;anal fin mm); Mbenji Island, Lake Malawi, Malawi, Africa, with 3 spines and 7 rays in holotype and 6-8 in April, 1991; PSU 3072, 4; (72.9 - 84.1); Mbenji paratypes. Lower pharyngealbone triangularin Island, Lake Malawi, Malawi, Africa, April, 1991. outline (Fig. 29). Scalesalong side ctenoid;holo- Diagnosis.-The moderately sloped head, swol- PSEUDOTROPHEUSZEBRA COMPLEX 211

Fig. 30. Metriadima mbenjii, HOLOTYPE, PSU 3069, adult male, 88.0 mm SL.

peduncle length (0.373); while those with the highest loadings on the principal components of the meristic data were teeth rows on the upper jaw (0.471), scale rows on cheek (0.463), and teeth rows on the lower jaw (0.427). Description.-Jaws isognathous (Fig. 30); teeth on jaws in 2-4 rows; majority of teeth in outer rows bicuspid, with a few posterior unicuspid teeth; those in inner rows tricuspid; 12 teeth in outer row of left lower jaw of holotype, 4-12 in paratypes. Dorsal fin with 18 spines in the holo- type and 17-18 in paratypes; dorsal-fin rays 8 in holotype and 8-10 in paratypes. Pectoral fin with 14 rays in holotype and 12-14 in paratypes; anal fin with 3 spines and 6-8 rays. Lower pharyngeal bone triangular in outline (Fig. 31). Scales along side ctenoid; holotype with 31 lateral-line scales Fig. 31. Lower pharyngeal bone of the type specimen of and 30-32 in paratypes. First gill arch with 11-13 Metriaclimambenjii, PSU 3069. rakers on ceratobranchial,2-3 on epibranchial, and len rostraltip of the neurocranium,isognathous 1 between the epibranchial and ceratobranchial jaws, and the presenceof bicuspidteeth in the (Table 10). outer rows of the jaws place this speciesin Metri- Most breeding males have a light blue ground aclima. The paleblue groundcoloration with very coloration with 6-8 faint vertical bars, and white faintvertical bars, in conjunctionwith a reddorsal belly; head below eye dark blue/gray; dorsal to fin (Fig. 32a), delimitsM. mbenjiifrom all other eye, pale blue with one dark blue/gray interorbital Metriaclimaspecies except for Metriaclimagreshakei bar, green opercular spot, and white gular. Dorsal (Meyerand Foerster). The first principalcompo- fin red orange with blue lappets; caudal fin with nents of the meristicdata were plottedagainst the blue rays and orange membranes; anal fin with shearedsecond principal components of the mor- orange proximal membranes, clear mid-fin mem- phometric data for M. mbenjiiand M. gresbakei branes, and blue tips; anal spines blue; posterior (Fig. 33). There was no overlapin the minimum anal-fin membranes red orange with 3 yellow polygon clustersformed by M. greshakeiwith that ocelli; P2 spine and first ray blue/gray with white formed by M. mbenjii. Variablesthat had the leading edge, remainder clear; P1 membranes highest loadingson the shearedsecond principal red/orange proximally, clear distally. A few componentswere snout length (-0.439),distance breeding males' dorsal one-half has a orange between the posterior anal-fininsertion and the ground color with dark blue blotches (Fig. 32b), dorsal caudal-fin insertion (0.419), and caudal and ventral one-half a blue ground coloration, scales are outlined in orange with blue blotches; 212 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

Fig. 32a. Metriadima mbenjii, blue form, from Mbenjii Island.

LiX~~~~~~~~~~~~~~~~~~~~~~~~~~~~p

Fig. 32b. Metraclima mbenjii, orange-blotch form, from Mbenjii Island. head dorsal to eyes is orange with dark blue and dear membranes. Most femaleswith brown blotchesand 2 light blue interorbitalbars; ventral ground color laterallywith green highlightsand to eyes is gray with purple,blue, and greenhigh- 6-8 verticalbars and graybelly; head brown with lights, gularlight blue/purple. Dorsalfin orange faint green interorbitalbar; dorsal fin brown; with light blue anddark blue blotcheswith several caudalfin light red/orangedistally; anal fin gray small orangespots in rayedportion of the mem- with 3 pale yellow ocelli; P2 spine and first ray branes;caudal fin orangewith light blue and dark black,remainder clear; P1 clear. Some femalesare blue blotches;anal fin orangeproximally and clear orange laterally with blue highlights and spots distally with three yellow ocelli; P2 dear with with blue/green belly;' head orange with occasionalorange spots; P1 with pale orangerays blue/greenopercle; dorsal and caudalfins orange PSEUD07ROPHEUS ZEBRA COMPLEX 213

0.15

0.1

Ci 0.05 A Q~~~~~~~~~~~~~ * Al mbenjii - OB \LI. A greshekai

0 A Al. mbenjii - Blue

j~~~~~ -0.05 7

-0.1 -3 -2 -1 0 1 2 Factor 1

Fig. 33. Plot of the sheared second principal components (morphometric data) and the first factor score (meristic data) for Metriaclima mbenjii, both the orange-blotch and blue forms, and Metriaclimagreshakei.

with dark orange spots scattered throughout; anal caudal peduncle length (0.373); while those with fin orange anteriorly and distally, clear proximally, the highest loadings on the principal components and 7-9 ocelli located posteriorly; P2 with first 2 of the meristic data were teeth rows on the upper membranes orange and the remainder clear with jaw (0.471), scale rows on cheek (0.463), and teeth orange spots; P1 pale orange with distal portion of rows on the lower jaw (0.427). some membranes black. Description.-Jaws isognathous; teeth on jaws in Distribution.-Metriaclima mbenjiiis endemic to 2-3 rows; majority of teeth in outer rows bicuspid, Mbenji Island. with a few posterior unicuspid teeth; those in Etymology.-The name referringto the fact that inner rows tricuspid; 3-12 teeth in outer row of M. mbenjii is endemic to Mbenji Island. left lower jaw. Dorsal fin with 17-18 spines and 8-9 rays. Pectoral fin with 11-14 rays; anal fin Metriaclima greshakei (Meyer and Foerster) with 3 spines and 6-7 rays. Lower pharyngeal Diagnosis.-The moderately sloped head, swol- bone triangular in outline. Scales along side len rostral tip of the neurocranium, isognathous ctenoid with 30-33 lateral-line scales. First gill arch jaws, and the presence of bicuspid teeth in the with 8-12 rakers on the ceratobranchial, 2-3 on outer rows of the jaws place this species in Metri- epibranchial, and 1 between the epibranchial and aclima. The pale blue ground coloration with very ceratobranchial (Table 10). faint vertical bars in conjunction with a red dorsal Breeding males have a blue/gray ground color fin, delimits M. greshakeifrom all other Metriaclima dorsally, fading to gray ventrally; live males are species except for M. mbenjii. The first principal cobalt with 6-8 faint vertical bars; head blue/gray components of the meristic data were plotted with blue opercle, green highlights and a white against the sheared second principal components of gular. Dorsal fin is dark rust colored proximally, the morphometric data for M. mbenjii and M. and orange/red distally with white lappets; cau- greshakei (Fig. 33). There was no overlap in the dal-fin rays are white with orange/red membranes; minimum polygon clusters formed by M. greshakei anal fin clear with 7 yellow ocelli; P2 with white with that formed by M. mbenjii. Variables that leading edge, clear rays, and gray/black mem- had the highest loadings on the sheared second branes. Females similarly colored, but sometimes principal components were snout length (-0.439), have a brown/rust ground color and the anal fin distance between the posterior anal-fin insertion has as many as 10 ocelli. and the dorsal caudal-fin insertion (0.419), and Distribution.-Metriaclima gresbakeiis restricted 214 J. R. STAUFFER,JR., N. J. BOWERS,K. A. KELLOGGAND K. R. MCKAYE

_F'

Fig. 34. Metriaclima thapsinogen,HOLOTYPE, PSU 3074, adult male, 86.4 mm SL. to the rock reefs south of Boadzulu Island, Lake Malawri(Konings 1990). Material examined.-PSU 3073, 20; (60.5 - 94.1 mm); Makokola Reef off southwestern end of Boadzulu Island, Lake Mala'wi, Malaw'i, Africa, 25-35 m, May, 1991.

Metriaclima thapsinogen Stauffer, Bowers, Kel- logg and McKaye, n. sp. (Fig. 34) HOLOTYPE.-PSU 3074, adult male 86.4 mm, Eccles Reef, Lake Malawri,Malawi, Africa, March, 1995, (Figs. 34 & 35). PARATYPES.-PSU 3075, 8; USNM 341931, 2; - 86.4 mm), data ANSP 176205 2; MFU 30, 2; (65.5 Fig. 35. Lower pharyngeal bone of the type specimen of as for holotype. Metriadima thapsinogen,PSU 3074. Diagnosis.-The moderately sloped head, swol- len rostral tip of the neurocranium, isognathous 6 black vertical bars; head dark blue with 2 light jaws, and the presence of bicuspid teeth in the blue interorbital bars, opercle with green high- outer rows of the jaws place this species in Metri- lights, and yellow gular. Dorsal fin orange with aclima. The blue ground coloration with distinct blue lappets; caudal fin gray in proximal se- vertical bars in conjunction with the orange dorsal ven-eighths, distal one-eighth orange; anal fin gray fin and yellow chin delimits M. thapsinogenfrom with 2 yellow ocelli; P2 orange/brown; P1 with all other species in Metriaclima. black rays and clear membranes. Females green Description.-Jaws isognathous (Fig. 34); teeth laterally with scales outlined in yellow; head gray on jaws in 2-4 rows; majority of teeth in outer with 2 green interorbital bars and pale yellow rows bicuspid, with a few posterior unicuspid gular; dorsal fin red; caudal fin gray; anal fin teeth; those in inner rows tricuspid; 9 teeth in proximal one-quarter gray, distal three-quarters outer row of left lower jaw of holotype, 7-11 in orange/brown with no ocelli; P2 orange/brown; P1 paratypes. Pectoral fin with 14 rays in holotype, with gray rays and clear membranes. 13-15 rays in paratypes. Lower pharyngeal bone Distribution.-Metriaclima thapsinogen was triangular in outline (Fig. 35). Scales along side collected only at Eccles Reef. ctenoid; holotype with 31 lateral-line scales and Etymology.-The name thapsinogenwas respelled 30-32 in paratypes. First gill arch with 11-13 from the Greek thapsinogenion, meaning yellow rakers on the ceratobranchial,2-3 on epibranchial, chin to note the yellow color of the gular and and 1 between the epibranchial and ceratobranchial branchiostegal rays. gable 11). Breedingmales have a blue groundcolor with PSEUD07ROPHEUSZEBRA COMPLEX 215

DISCUSSION the reductionof the left ovary is evidencewhich aligns the genus, Metriaclima,with the other We agree with Mayden and Wood (1995) and groupsof rock-dwellingcichlids of Lake Mala'wi. Mayden (in press), that the Evolutionary Species The presence of bicuspid teeth in the anterior Concept, sensu Wiley (1978) is the only valid portion of the outer rows of both the upper and theoretical concept; however it is non-operational lower jawsdistinguishes Metriaclima from Cyatho- and must be used in conjunction with other chromis,Cynotilapia, Gephyrochromis, and Labido- surrogate species concepts (Stauffer et al. 1997). chromis. The isognathousjaws of Metriaclima When delimiting sympatric species, differentiation delimits it from Genyochromis,and its terminal of morphological traits, including color patterns, mouth discriminatesMetriaclima from Labeotroph- are considered to be indicative of reproductive iso- eus. Withinwhat was historicallyconsidered to be lation (biological species concept; Mayr and Pseudotropheus,the moderatelysloped ethmo-vom- Ashlock 1991) or specific mate-recognition system erineblock and swollen rostral tip to the neurocra- (recognition species concept; Paterson 1993). The nium separatesMetriaclima from the P. elongatus, designation of allopatric populations as either P. tropheops,and P. williamsi complexes. De- being conspecific or heterospecific is more prob- scribedspecies that arenot discussedin this paper, lematical. but which are characterizedby a moderately As indicated by Lewis (1982), the acquisition of slopedethmo-vomerine block and swollen rostral reproductive isolation without significant morpho- tip, and thus belong to Metriaclimainclude: logical change makes it difficult to delimit African Metriaclimaaurora, Metriaclimabarlowi, Metri- haplochromine cichlids. Additionally, the use of aclima elegans,Metriaclima estherae, Metriaclima starch gel electrophoresis has been inconclusive for hajomaylandi,Metriaclima lombardoi, Metriaclima delimiting species (Kornfield 1978), although DNA lanisticola,Metriaclima livingstonii, andMetriaclima sequencing of mtDNA may be adequatefor distin- pursus. guishing among Lake Malawi cichlids in some lin- WithinMetriaclima, the speciesdescribed herein eages (Bowers et al. 1994). Rapid speciation of representthose that have a blue lateral ground Malaw'iancichlids, however, may have prevented color with blackvertical bars, or have secondarily sorting of mitochondrial lineages, permitting dis- lost these barsand appearto be solid blue. This tantly related species to share mtDNA polymorph- blue/blackcolor patternis presentin many of the isms derived from a common ancestor (Moran and other genera of rock-dwellingcichlids, which Kornfield 1993). Many morphologically and inhabit Lake Mala'wi,including Labidochromis genetically similar species have been separated zebroides Lewis, Petrotilapia "likoma barred" based on breeding coloration and behavioral (Ribbinket al. 1983),Cynotilapia afra (Gunther), characteristics (Ribbink et al. 1983, McKaye and Pseudotropheuselongatus Fryer, Pseudotropheuscf. Stauffer 1986, Stauffer et al. 1993). For example, williamsi'namalenge', Pseudotropheus microstoma behavioral observations were used to hypothesize Trewavas (P. tropheopscomplex), and Labeotropheus that the blue-black color form (M. zebra) was trewavasaeFryer. The blue/black color pattern reproductively isolated from the blue color morph can also be found in many of the sand-dwelling (M. callainos) at Nkhata Bay (Holzberg 1978, fishessuch as Protomelas similis (Trewavas), Copadi- Schroder 1980). chromiscyaneus (Trewavas), and Trematocranus Many of the forms described herein, were first microstomaTrewavas. Thus, Staufferet al. (1995) hypothesized to be undescribed species based on speculatedthat the blue/black color is pleisio- differences in color pattern. Unique color patterns morphicbased on eitherthe commonalityprinci- were used in conjunction with the discontinuity of ple (Watrousand Wheeler 1981) or outgroup morphological differences to determine specific comparisons(Smith and Koehn 1971). status. For example, if a series of allopatric popu- Metriaclimazebra is consideredto be the most lations demonstrated gradualvariation throughout primitivespecies in the genus. Certainlyit is the its range (i.e. M. zebra; Figs. 6a, 6b, 6c) they were mostwidespread, ranging from CrocodileRocks at considered to be conspecific. If however, similarly the southernend of Lake Malawiito Chilumba, colored forms exhibited discontinuities in morpho- which is located in the north. Breedingmales logical characteristicsthroughout their range, they exhibitthe blue/blackcolor pattern,and there are were regardedas heterospecific populations (i.e. M. no otherpigmentation patterns (e.g. red dorsal fin, pyrsonotos,M. sandaracinos,M. emmiltos;Fig. 25). yellow throat, extension of lateral vertical bars The combination of the following: 1) presence onto the dorsal fin), which are interpretedas of many small scales in the nape region, 2) prefer- apomorphiccharacters. ence for boulder and rocky substrate, and 3) and Surely, allopatric speciation and subsequent 216 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE dispersalhas occurredwithin Metriaclima. For Intra- and inter-specificmitochondrial DNA sequence example,the occurrenceof M. malabranchionat variation within two species of rock-dwelling cichlids both MitandeRocks and Domwe Islandat Cape (Teleostei: Cichlidae) from Lake Malawi, Africa. MaClear,can be explainedby dispersal.It is much Molecular Phylogenetics and Evolution 3: 75-82. 1997. more difficult,however, to believethat the species Bowers, N. J. and J. R. Stauffer, Jr. Eight new species of rock-dwelling cichlids of the genus Melano- with a red dorsalfin, M. emmiltos,which is endem- chromis (Teleostei: Cichlidae) from Lake Malawi, ic to Chilumbain the northernpart of the lake is Africa. Ichthyol. Explor. Freshwaters 8: 49-70. more closely related to other species with red Fryer, G. 1959. The trophic interrelationships and dorsalfins (i.e. M. sandaracinosand M. pyrsonotos) ecology of some littoral communities in Lake Nyasa, than it is to the M. zebrapopulation with which it with especial reference to the fishes, and a discussion is sympatric. of the evolution of a group of rock-frequenting Metriaclimaelegans, M. lanisticola,M. living- Cichlidae. Proc. Zool. Soc. Lond. 132: 153-281. stonii, M. pursusare the only describedmbuna Fryer, G. and T. D. Iles. 1972. The cichlid fishes of the species,which have secondarilyinvaded the sand. Great Lakes of Africa. TFH Publications, Neptune, All of these speciesare associated with emptysnail USA. Holzberg, S. 1978. A field and laboratory study of the shells(Lanistes spp.) either as adultsor duringtheir behaviour and ecology of Pseudotropbeus zebra ontogeny. This behaviorcharacter may in factbe (Boulenger), an endemic cichlid of Lake Malaw4i, an apomorphic character,in which case these Africa. Zeitschr. Zool. Syst. Evolutg. Forsch. 16: 171- speciesare more closelyrelated to eachother than 187. to other sympatricMetriaclima species, or it may Humphries, J., F. Bookstein, B. Chernoff, G. Smith, R. be due to convergence.A detailedanalysis of this Elder, and S. Poss. 1981. Multivariate discrimination group must be completed,which includesat least by shape in relation to size. Systematic Zoology 30: two undescribedspecies endemic to Nakantenga 291-308. and Likomaislands. Konings, A. 1990. Cichlids and all the other fishes of Lake Mala4ii. T.F.H. Publications, Inc., Neptune City, New Jersey. ACKNOWLEDGEMENTS Konings, A. 1995. Malaw'i cichlids in their natural habitat. Cichlid Press, Germany. The authorswish to thank the governmentof Kornfield, I. 1978. Evidence for rapid speciation in Malawi for providing the necessarypermits to African cichlid fishes. Experientia 34: 335-336. collect fishes. L. Knappof the SmithsonianSort- Lewis, D. S. C. 1982. Problems of species definition in ing Center(USNM) provided monies for shipping Lake Mala4zi cichlid fishes (Pisces: Cichlidae). a portion of the specimensback to the United Ichthyological Bulletin of the J. L. B. Smith Institute States. M. Katzretouched the photographsof the of Ichthyology 23: 1-5. type specimens.This work was fundedin partby Mayden, R. L. in press. A hierarchy of species concepts: the deouement in the of the the StatesAgency for InternationalDevel- saga species problem. In: United M. F. Claridge, H. A. Dawah, and M. R. Wilson opment (GrantNo. 11.204;DHR-5600-G-1043-00, (eds.). The units of biodiversity - species in practice. Programin Scienceand Technology Cooperation, Systematics Association, London. Office of Sciences Advisor), National Science Mayden, R. L. and R. M. Wood. 1995. Systematics, Foundationgrant BNS 86-06836),and two Ful- species concepts, and the evolutionarily significant brightResearch Awards (Council for International unit in biodiversity and conservation biology. Amer. Exchangeof Scholars)to JRS. Ad Koningssup- Fish. Soc. Sym., 17: 58-113. plied the underwater photos of Metriaclima Mayr, E. and P. D. Ashlock. 1991. Principles of system- mbenjii. GeorgeTurner and T. D. fles provided atic zoology. McGraw-Hill, New York. valuablecomments, which greatlyimproved the McKaye, K. R., T. Kocher, P. Reinthal, R. Harrison, and I. Kornfield. 1984. Genetic evidence for allotropic manuscript. and sympatric differentiation among color morphs of a Lake Mala4ii cichlid fish. Evolution 31: 215-219. LITERATURE CITED McKaye, K. R. and J. R. Stauffer,Jr. 1986. Description of a gold cichlid, Pseudotropheusbarlowi (Teleostei: Barel, C., M. van Oijen, F. Witte, and E. Witte-Maas. Cichlidae), from Lake Malaw4i,Africa. Copeia 1986: 1977. An introduction to the taxonomy and mor- 870-875. phology of the haplochromine Cichlidae from Lake Meyer, M. K. and W. Foerster. 1984. Un nouveau Victoria. Neth. J. Zool. 27: 333-389. Pseudotropheusdu lac Malanviavec des remarques sur Bookstein, F., B. Chernoff, R. Elder, J. Humphries, G. le complexe Pseudlotropheus-Melanochromis(Pisces, Smith and R. Strauss. 1985. Morphometrics in Perciformes, Cichlidae). Revue fr. Aquariol. 10: Evolutionary Biology. Academy of Natural Sciences, 107-112. Spec. Publ. 15, Philadelphia, PA. Moran, P. and I. Kornfield. 1993. Retention of an Bowers, N. J., T. D. Kocher, and J. R. Stauffer,Jr. 1994. ancestral polymorphism in the mbuna species flock PSEUDOTROPHEUSZEBRA COMPLEX 217

(Pisces: Cichlidae) of Lake MalawiI. Molecular Biolo- 1993. Three new sand-dwelling cichlids from Lake gy and Evolution 10: 1015-1029. Malawi, Africa, with a discussion of the status of the Moran, P., I. Kornfield, and P. Reinthal. 1994. Molecu- genus Copadichromis(Teleostei: Cichlidae). Copeia lar systematics and radiation of the haplochromine 1993: 1017-1027. cichlids (Teleostei: Cichlidae of Lake Malawi. Copeia Trewavas, E. 1935. A synopsis of the cichlid fishes of 1994: 274-288. Lake Nyasa. Ann. Mag. Nat. Hist. (10)16:65-118. Oliver, M. 1984. Systematics of African fishes: determi- Watrous, L. E. and Q. D. Wheeler. 1981. The nation of the most primitive taxon, and studies on the out-group comparison method of character analysis. haplochromines of Lake Malaw'i (Teleostei:Cichlidae). Systematic Zoology 30:1-11. Ph.D. Dissertation, Yale University, New Haven, CT. Wiley, E. 0. 1978. The evolutionary species concept Oliver, M. L. and P. V. Loiselle. 1972. A new genus reconsidered. Systematic Zoology 27: 17-26. and species of cichlid of the mbuna group (Pisces: Cichlidae) from Lake Malawii. Revue de Zoologie et de Botanique Africaines 85: 309-320. Paterson, H. E. H. 1993. Evolution and the recognition concept of species. In: S. F. McEvey (ed.). Collected [TABLES 1-11 FOLLOW] writings of H. E. H. Paterson. Johns Hopkins Univer- sity Press, Baltimore, Maryland. Reyment, R., R. Blackith, and N. Cambell. 1984. Multivariate Morphometrics. Academic Press, New York, NY. Ribbink, A. J., B. A. Marsh, A. C. Marsh, A. C. Ribbink and B. J. Sharp. 1983. A preliminary survey of the cichlid fishes of the rocky habitats in Lake Malaw'i. S. Afr. J. Sci. 18:149-310. Schroder, J. H. 1980. Morphological and behavioural differences between the BB/OB and B/W colour morphs of Pseudotropheuszebra Boulenger (Pisces: cichlidae) Ztschr. Zool. Syst. Evolutg. Forsch. 18: 69-76. Smith, G. R. and A. K. Koehn. 1971. Phenetic and cladistic studies of biochemical and morphological characteristics of Catostomus. Systematic Zoology 20:282-297. Staeck, W. 1980. Pseudotropheusheteropictus n. sp. Aus dem Malaw4i-See(Pisces: Cichlidae). Senckenbergiana Biol. 60: 159-162. Stauffer, J. R., Jr. 1988. Three new rock-dwelling cichlids (Teleostei: Cichlidae) from Lake Malaw'i, Africa. Copeia 1988: 663-668. Stauffer, J. R. 1991. Description of a facultative cleaner- fish (Teleostei: Cichlidae) from Lake Malaw'i,Africa. Copeia. 1991: 141-147. Stauffer, J. R., Jr. and J. M. Boltz. 1989. Description of a rock-dwelling cichlid (Teleostei: Cichlidae) from Lake Malaw4i,Africa. Proc. Biol. Soc. Wash. 102: 8-13. Stauffer, J. R., Jr., J. J. Bowers, K. R. McKaye, and T. D. Kocher. 1995. Evolutionarily significantunits among cichlid fishes: The role of behavioral studies. Amer. Fish. Soc. Sym. 17:227-244. Stauffer, J. R., Jr. and E. Hert. 1992. Pseudotropheus callainos, a new species of mbuna (Cichlidae), with analyses of changes associated with two intra-lacus- trine transplantations in Lake Malaw'i, Africa. Ichthyol. Explor. Freshwaters 3(3):253-264. Stauffer, J. R., Jr., C. H. Hocutt, and R. L. Mayden. 1997. Pararhinichthys,a new monotypic genus of minnows (Teleostei: Cyprinidae) of hybrid origin from eastern North America. Ichthyol. Explor. Freshwaters 7: 327-336. Stauffer, J. R., Jr., T. J. LoVullo, and K. R. McKaye. 218 J. R. STAUFFER, JR., N. J. BOWERS, K. A. KELLOGG AND K. R. MCKAYE

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