cladistic, phenetic and biogeographical analysis of the flightless dung beetle genus, gyronotus van...

Journal of Natural History 2001 35 1607ndash1625

Cladistic phenetic and biogeographical analysis of the macr ightless dungbeetle genus Gyronotus van Lansberge (Scarabaeidae Scarabaeinae)

in threatened eastern Afrotropical forests

ADRIAN L V DAVIS CLARKE H SCHOLTZ andJAMES DU G HARRISON

Department of Zoology amp Entomology University of Pretoria Pretoria

0002 South Africa e-mail adaviszoologyupacza

(Accepted 15 April 2000)

By virtue of their low vagility ightless insects are useful indicators of biogeo-graphical history Relationships of the ightless dung beetle genus Gyronotusare of particular interest due to its Gondwanaland ancestry distinctive relictdistribution along the south-eastern seaboard of Africa and its restriction toforests which are seriously threatened by exploitation Because of the limitednumber of diagnostic morphological characters it was necessary to code morpho-metric data in order to conduct distance and cladistic parsimony analysis ofinterspeci c relationships in Gyronotus There was a good correlation betweenrelationships indicated by the dendrogramscladograms and those determined byan examination of aedeagus character states both of which indicate a disjunctionbetween south and east African species and a broad separation between northernand southern South African species Comparison of the bilaterally asymmetricalaedeagi of Gyronotus with the symmetrical aedeagi of the sister genus AnachalcosHope suggests geographical polarization of character states from greater plesi-omorphy in east African Gyronotus to greater apomorphy in South Africanspecies particularly in the southernmost element in which the aedeagus showsextreme asymmetry Furthermore body shape follows a similar geographicalgradient in that the three Gyronotus species of tropical east Africa are signi cantlymore elongate than the three ovoid lowlandafromontane species of South AfricaAn examination of historical factors suggests that this spatially-restricted distribu-tion is the relict of a very old tropical lowland pattern In extant taxa thephylogenetic polarization is towards one of ve main centres of afrotropicalforest biodiversity in the geologically old Eastern Arc and the adjoining lowlandforest (Swahili centre of endemism) Survivors from old lineages may be onereason for such centres of high biodiversity

Keywords Afrotropical biogeographical cladistic dung ightless forestphenetic Scarabaeinae

Introduction

Eleven of the 20 extant genera of canthonine dung beetles in Africa are restrictedto the south-eastern seaboard between the Western Cape in South Africa and the

Journal of Natural HistoryISSN 0022-2933 printISSN 1464-5262 online copy 2001 Taylor amp Francis Ltd

httpwwwtandfcoukjournals

A L V Davis et al1608

East Usambara Mountains in north-eastern Tanzania (Scholtz and Howden 1987)where they are primarily found in forest habitats (Davis unpublished ) These 11genera may be regarded as relicts since they belong to a tribe with ancientGondwanaland a liations (HalŒter and Matthews 1966 HalŒter 1974 Cambefort1991) and are all species-poor This relictual distribution pattern is presumablyrelated to historical events particularly to the Miocene and Plio-Pleistocene upliftin east central Africa which separated the forests of eastern Africa from those inWest Africa (Lovett 1993 Burgess et al 1998) In East Africa survival of wetrainforest and its associated faunal relicts may be a consequence of the relativelystable surface temperatures (0ndash2 szlig C variation) in the Indian Ocean (Prell et al 1980)and continued coastal and orographic capture of high rainfall (Gillman 1949)during the Pleistocene cycles of cool dry and warm wet climate

The genus Gyronotus is one of the most species-rich of these eastern seaboardforest relicts It comprises six known species showing disjunct or parapatric distribu-tions (Scholtz and Howden 1987 Davis et al 1999) extending from the marginsof the bimodal rainfall region in the Eastern Cape South Africa (Davis 1993 1997)to the East Usambara Mountains which lie towards the northern end of the easternarc mountain system in north-eastern Tanzania All Gyronotus species are ightlessand are known to occur predominantly on ner-grained soils in lower altitude forestswith tropical or afromontane a nities Such ightless taxa with their low vagilityhave been shown to be most useful indicators of evolutionary and biogeographicalhistory (Bruhl 1997)

Both northern tropical and southern temperate (Palaeantarctic) Gondwanalandorigins have been hypothesized for afrotropical members of the tribe Canthonini(HalŒter 1974) However subsequent intra-continental dispersal vicariance eventsand extinctions have presumably shaped the modern relictual distribution patternsof the genera Phenetic cladistic and morphological analyses were conducted onGyronotus in order to examine their interspeci c relationships in comparison to thebiogeographical a nities of the species The results suggest a tropical generic a li-ation with a change in body shape from the lowland forest of East Africa to thelowland and low-lying afromontane forests of South Africa These geographicaldistribution patterns are discussed in an historical context

The ndings describe the relict distribution of an old dung beetle lineage ineastern Afrotropical forests In the present day these eastern forest habitats comprisean archipelago (Hamilton 1989 Low and Rebelo 1996 Burgess et al 1998) ofboth natural and anthropogenically-induced fragments which are seriouslythreatened by continuing exploitation Barnes (1990) predicted that these easternAfrotropical forests would be reduced by 95 by the year 2040 However recentanalyses suggest that the coastal portion between Somalia and Mozambique isalready reduced to a few thousand square kilometres of small fragments (Burgesset al 1998) Furthermore at least one of the species of Gyronotus dung beetles inthe present study (G glabrosus Scholtz and Howden) already deserves criticallyendangered (CR) status ( IUCN 1994) and may well be extinct (EX ) (Daviset al 1999)

Methods

Geographical distribution data for Gyronotus were derived from material heldby the University of Pretoria (now lodged in the Transvaal Museum Pretoria) andmaterial loaned by seven southern African and three European museums and

Cladistic and biogeographical analysis of Gyronotus 1609

institutions Specimens of four species were well-represented in these collections G

pumilus (Boheman) (1281 ) G carinatus Felsche (53 1 ) G Wmetarius Kolbe (41)and G mulanjensis Davis Scholtz and Harrison (33) However the institutionsheld only limited numbers of G dispar (Felsche) (7) and G glabrosus Scholtz andHowden (6) A seventh example of G glabrosus in the Howden Collection atCarleton University Ottawa Canada is the only other known museum specimenof this species During its known activity period in February 1998 unsuccessfulattempts were made to collect further material of this highly endemic species at itsone remaining known relatively undisturbed locality the afromontane Woodbushindigenous forest (23 szlig 52frac34 S 29 szlig 56frac34 E )

The morphology of Gyronotus showed relatively little variation and providedextremely few characters useful for cladistic analysis of interspeci c relationshipsIn such situations Chapill (1989) suggested that one alternative is to code morpho-metric data for cladistic analysis Since the use of phenetic characters for phylogeneticanalyses has been criticized (Wiley 1981) the results were compared with characterstates shown by the aedeagi These characters were the only morphological featureswhich showed diŒerences that could be interpreted with con dence as having strongphylogenetic signi cance

Twenty-seven morphometric characters used for distance or parsimony analysisof Gyronotus are listed in table 1 Measurements were made of all available specimensof G dispar (three males four females) and G glabrosus (four males two females)whereas ten specimens were measured for each of the other four species G WmetariusG pumilus G carinatus ( ve males ve females) and G mulanjensis (six malesfour females) The morphometric data matrix was treated in two diŒerent ways inorder to change the relative in uence of size and shape on the analyses Themeasurements (mm) were either left untransformed or were log10 transformed Bothtreatments were double standardized (Somers 1989) by subtracting each value fromthe row mean and each of these standardized values from the standardized columnmean For each treatment the data for each morphometric character was tested forsigni cant diŒerences between the six species using one-way ANOVA (table 1)Multiple range tests were conducted on each character using Tukeyrsquos HSD forunequal sample size (SpjotvollStoline Test ) On the basis of the results for thesemultiple range tests each character was coded for cladistic analysis using the diver-gence coding method of Thorpe (1984) This method provides a diŒerent code foreach signi cantly diŒerent character state and develops a species Ouml character statedistance matrix coded in categorical terms

Both distance and cladistic parsimony analyses were conducted on each of thetwo divergence matrices (table 2) using PHYLIP version 35 (Felsenstein 1993) andPAUP version 31 (SwoŒord 1993) Distance analyses were conducted using all 27characters in each matrix and species were linked by neighbour joining In contrastonly informative characters were used for cladistic parsimony analysis In the matrixderived from raw double-standardized data there were only 11 informativecharacters whereas in that derived from log10 double-standardized data there were14 informative characters In each analysis the character states were treated asunordered and the cladograms were centre-rooted

Each of the morphometric matrices was also subjected to principal componentsanalysis (PCA) As the rst component frequently correlates to size whereas follow-ing components correlate primarily to shape (Somers 1989 Fairbairn 1992) theordinate values for the rst two factors from each analysis were tested for correlation


Table 1 List of morphometric characters for Gyronotus and the results of one-way ANOVAon interspeci c diŒerences between the characters either double-standardized rawmeasurements in mm or double-standardized log10 transformed measurements (seemethods)

F(5 47) F(5 47)(raw data)dagger ( log10 data)dagger

1 Distance between the tips of the clypeal teeth 17255 45972 Distance between the genal angles 6708 68373 Distance between the centre of the clypeal emargination and 10360 3422

a line drawn between the genal angles4 Distance between the centre of the clypeal emargination and 1384 1971

the posterior margin of the vertex5 Width between the anterior angles of the pronotal disc 4169 41236 Width between the lateral angles of the pronotal disc 3240 16667 Width between the posterior angles of the pronotal disc 493 20288 Length along the middle of the pronotal disc 4780 52659 Maximum depth of the pronotal disc between the dorsal 4695 3887

surface and the ventral surface of the profemora10 Depth of the pronotal disc between the dorsal surface and 1002 926

the pleural margin at the anterior end11 Depth of the pronotal disc between the dorsal surface and 1511 874

the pleural margin at the posterior end12 Distance from the centre of the posterior margin to a line 7113 2056

drawn between the posterior angles of the prothoraccic disc13 Length along the midline of the metasternum 665 105514 Distance between the left mesocoxa and left metacoxa 8648 43715 Distance between the right mesocoxa and the left metacoxa 1243 80316 Distance between the mesocoxae 1093 103517 Maximum depth from the elytra to the metasternum 3518 94618 Depth from the dorsal extremity of the left elytron to the 2037 889

pleural margin at the anterior end19 Maximum depth of the left elytron to the pleural margin 732 105820 Maximum height of the curvature along the lateral edge of 7638 1327

the left elytron above a line drawn between the anterior andposterior ends

21 Maximum width of the elytral pseudo-epipleurae 7702 280722 Maximum width of the right elytron Ouml 2 2496 388323 Width of the right elytron at the anterior end Ouml 2 1153 271824 Maximum length of the right elytron 5538 99225 Width of the pygidium along the basal ridge 1733 204226 Length of pygidium from the tip to the basal ridge 2326 227827 Length of the sternites along the midline of the abdomen 2709 1222

between the metasternum and the pygidium

daggerAll F numbers statistically signi cant plt0001

with both body length and the ratio of body length to maximum width of the elytraThese respective measurements were used as indices for body size and body shapeThe results provide some indication of the relative in uence of these two parameterson the four derived matrices used for distance and parsimony analyses

The relationships indicated by the cladograms are compared to characters shownby the aedeagi of Gyronotus those of Anachalcos Hope and that of Canthodimorpha

Davis Scholtz and Harrison which is a newly-discovered large-bodied monotypictropical genus from the northern coastline of Mozambique (Davis et al 1999)


Tab

le2

Div

erge

nce

mat

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s(T

horp

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Although Anachalcos has been identi ed as the sister group (Scholtz and Howden1987) it is probably quite distant from Gyronotus which is both ightless and hasasymmetrical aedeagi However it is a useful comparison since the aedeagi ofAnachalcos are similar to those of most other afrotropical Canthonini in as muchas the parameres are symmetrical (Scholtz and Howden 1987) and lack the mem-branous terminal process on the left paramere which is characteristic of all Gyronotus

(Davis et al 1999)

Results

Geographical distribution

The six Gyronotus species showed either disjunct or parapatric distributions fromthe Eastern Cape in South Africa to north-eastern Tanzania ( gure 1) Two SouthAfrican species G pumilus (Eastern Cape southern KwaZulu-Natal ) and G

carinatus (northern KwaZulu-Natal ) showed a parapatric distribution pattern larg-ely restricted to coastal hills below 320m ( gure 2) The southern distribution limitsof G pumilus and the northern distribution limits of G carinatus coincide with thesouthern and northern limits of Walter and Liethrsquos (1964) temperatesubtropicalclimate type II (I )a ( gure 2) The respective northern and southern distribution

Fig 1 Geographical distribution of Gyronotus species from east to southern Africa(T Tanzania M Malawi SA South Africa) (S Swahili centre of endemism SMSwahili-Maputaland transition zone (Burgess et al 1998) Ma Maputaland centre ofendemism P Pondoland centre of endemism which occupies a narrow coastal zone(van Rensburg et al 1999))


Fig 2 Geographical distribution of Gyronotus pumilus and G carinatus relative to altitudeand the subtropical coastal climate type II(I )a after Walter and Lieth (1964)

limits of G pumilus and G carinatus occur just north of Durban (29 szlig 51frac34 S 31 szlig 01frac34 E )at which latitude land greater than 1500 m in altitude swings away from the coastlineThe third southern African species was recorded only in a small localized region ofthe eastern escarpment of Northern Province in South Africa The combined distribu-tion of these species is sympatric with both coastal forest (particularly in the south)and the low-lying portion of forest classi ed as Afromontane (particularly in thenorth) (table 3) Distribution data for the east African species was geographicallymore limited ( gure 1) Most G Wmetarius were recorded from 215ndash850m in theEast Usambara Mountains of north-eastern Tanzania although a few specimenswere collected from the adjacent coastline The few G dispar were recorded fromsouth-eastern Tanzania around Lindi (10 szlig 00frac34 S 39 szlig 42frac34 E ) and along the RovumaRiver (11 szlig S 39 szlig E ) All G mulanjensis were recorded from 900ndash1000 m on MtMulanje (16 szlig 22frac34 S 35 szlig 07frac34 E ) in southern Malawi

Distance and cladistic analyses

The dendrograms and cladograms derived from the four diŒerent analyses areshown by gure 3 In essential details the inter-relationships among the species wereconsistent with their geographical distribution Each dendrogramcladogram showeda clear branching between east and southern African taxa and in each case there


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(gu

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Fig 3 Centre-rooted distance dendrograms and centre-rooted consensus parsimony clado-grams for inter-relationships between Gyronotus species (A B) Cladistic matrix derivedfrom double-standardized raw morphometric measurements by divergence coding(Thorpe 1984) (A) Distance dendrogram based on 27 characters (B) Parsimonycladogram based on 11 informative characters tree length 5 28 CI 5 0893 (C D)Cladistic matrix derived from double-standardized log10 transformed morphometricmeasurements by divergence coding (Thorpe 1984) (C ) Distance dendrogram basedon 27 characters (D) Parsimony cladogram based on 14 informative characters treelength 5 36 CI 5 0889 The numbers are the branch lengths from the nodes forunordered character states

was a similar resolution of relationships between South African species with thesouthernmost species G pumilus distanced from the two more northerly speciesHowever the results for the east African species indicated three diŒerent hypotheticalinter-relationships between species Of the three diŒerent patterns one each wasyielded from the matrices derived from double-standardized raw data whereas thethird was common to the dendrogram and cladogram yielded by the matrices derived


from double-standardized log10 data These diŒerences may re ect the relativein uence of body size versus body shape according to the treatment of the datamatrices In the matrices derived from double-standardized raw data body size hasa stronger overall in uence than body shape (higher r2 and F numbers for regressionagainst Factor 1 which accounts for the greatest amount of variance in ordination(table 4) This emphasis is re ected by the closer linking of the two largest speciesG dispar and G Wmetarius Although both body size and body shape are de-emphasized in the matrices derived from double-standardized log10 data body shapehas the greater overall in uence (higher r2 and F numbers for regression againstboth Factors 1 and 2) This emphasis is re ected by the steps in common betweenthe elongate G dispar and G mulanjensis whereas the less elongate G Wmetarius isplaced on a separate branch ( gures 3C 3D)

Further phenetic analysis

As the cladograms were derived from morphometric data it is not surprisingthat the depicted interspeci c relationships are consistent with those indicated bythe three basic measurements of maximum length width and depth These measure-ments clearly separate the more elongate deeper east African species from the moreovoid atter southern African species (table 5) Ratios between the three dimen-sions indicate that width varies inversely with the other two dimensions whereaslength and depth vary proportionately to one another Thus the broadest species(G glabrosus) is also the attest and the most narrow species (G dispar) is alsothe deepest

Character states of aedeagi

F igure 4 illustrates the morphological diŒerences between the aedeagi ofGyronotus and those of large-bodied putative sister groups with primarily tropicaldistributions The parameres of tropical species in putative sister genera Anachalcos

procerus Gerstaecker and Canthodimorpha lawrencei Davis Scholtz and Harrisonare symmetrical and diverge at the tips Those of the east African Gyronotus alsodiverge at the tips but are asymmetrical due to a terminal process on the leftparamere The tips of the parameres of the southern African Gyronotus species aremore convergent and are also asymmetrical due to a terminal process projectingaway to the left of the left paramere

Discussion

Geographical polarization of characters

Comparison of Gyronotus aedeagi with those of the putative sister groupAnachalcos and that of Canthodimorpha suggests that bilateral symmetry anddivergence at the tips of the parameres are plesiomorphic character states ( gure 4)Although the aedeagi of the east African species of Gyronotus are characterized byasymmetry this is primarily due to the presence of a membranous terminal processwhich in ventral view projects to the right from the left paramere and occupies thespace between the divergent tips of the parameres ( gure 4) In contrast the SouthAfrican species are characterized by apomorphic convergence of the tips of theparameres ( gure 4mdashparameres slightly separated in the gure to facilitate drawing)In the two northernmost South African species asymmetry of the parameres ismainly due to the membranous terminal process which in ventral view projects


Tab

le4

The

corr

elat

ion

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sbo

dysi

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h)1

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63

173

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p

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141

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135

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77d

206

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159

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162

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26b

194

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(W)

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36b

900

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107

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911

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354

1

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027

b88

35

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atio

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W0

62Ocirc

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b0

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b0

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16

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Fig 4 Distal and ventral views of canthonine aedeagi showing diŒerences in parameremorphology between (A) Anachalcos procerus Gerstaecker (distal ) (B)Canthodimorpha lawrencei Davis Scholtz and Harrison (ventral) (C ) east AfricanGyronotus van Lansberge (ventral ) (C1 G Wmetarius Kolbe C2 G mulanjensis DavisScholtz and Harrison C3 G dispar (Felsche)) and (D) South African Gyronotus(ventral ) (D1 G carinatus Felsche D2 G glabrosus Scholtz and Howden D3G pumilus (Boheman)) Scale bars 5 1 mm

away to the left of the left paramere However in the southernmost species theasymmetry is more extreme in that each paramere is radically diŒerent in shape(Davis et al 1999) The left paramere is more or less straight and bears the leftwardsaligned terminal process the right paramere is curved towards the mid-point andboth are quite unlike those of the other ve species in which each paramere issimilarly curved close to the distal end Thus there is clear biogeographical polariza-tion of the character states of the aedeagi with greater plesiomorphy in eastAfrican species


Parallels between morphology biogeography and phenetic analysis of GyronotusThe interspeci c relationships indicated by the cladograms derived from phenetic

analysis largely corresponded to relationships suggested by morphological charactersof the aedeagi ( gure 4 Davis et al 1999) and these relationships were consistentwith the biogeographical distribution of the species along the south-eastern seaboardof Africa The clear separation of the elongate east African and more ovoid southernAfrican species is re ected by the diŒering position of the terminal process on theleft paramere of the aedeagi ( gure 4) The shape of this process in G Wmetarius

and in G mulanjensis diŒers only slightly in that the one is rounded and the otherangular Although it appears that these two species are very close the angularprocess of G dispar is su ciently similar that it is di cult to determine which ofthe cladograms if any re ects the true phylogenetic relationships between the eastAfrican species those strongly re ecting size and shape ( gures 3A 3B) or thosemore biased by shape ( gures 3C 3D) It is also possible that evolution of aedeagiand morphometric character states are divergent rather than parallel The relation-ships between the southern African species are resolved with greater consistencyThe shape of the parameres is similar in the two northerly species which diŒerprincipally only in the mono d shape of the outward projecting terminal process inG carinatus and the bi d shape of this process in G glabrosus ( gure 4) The moredistant phenetic separation of the southernmost species G pumilus is very stronglysupported by the characters of its aedeagus which diŒer radically from those of theother species However the outward projection of the terminal process indicates itscloser relationships to the other South African species than to the east Africanspecies

Biogeographical climatic and habitat associations of GyronotusSingle Gyronotus species are characteristic of several southern sub-centres of

endemism identi ed in mainland eastern coastal forests of Africa by Burgess et al(1998) These sub-centres occur in the East UsambaraTanga (G Wmetarius) Lindi(G dispar) and UdzungwaMulanje mountain regions (G mulanjensis) The coastalforests of southern Tanzania (Lindi subcentre) contain the most unique assemblageof endemic taxa along the east coast (Burgess et al 1998) and also harbour thelargest of the six known Gyronotus species (G dispar)

In terms of lowland versus afromontane forest association the biogeographicala nities of Gyronotus are dependent on the system used to classify oral zonationIn diŒerent regions altitudinal temperature zonation is modi ed by a combinationof factors including latitude rainfall regime aspect and the relative in uence ofmaritime versus continental climate (Hamilton 1989) Therefore instead of usinga single in exible classi cation system we have used three regional systems whichmake allowance for geographical diŒerences in the altitudinal occurrence of thesame temperature zones and their related ora These classi cation systems are thoseproposed by Lovett (1993) for the Eastern Arc Chapman and White (1970) forMalawi and Low and Rebelo (1996) for southern Africa Thus the East Africanspecies of Gyronotus may be described as elements of tropical lowland forests whichoccur below 1370 m in Malawi (Chapman and White 1970) and below 800m in theeastern arc (Lovett 1993) In the more southerly latitudes of South Africa mostlowland and highland forest is classi ed as afromontane (Low and Rebelo 1996)except for that on coastal sandveld of recent origin (Lawes 1990) AlthoughGyronotus have been recorded in coastal regions around East London (33 szlig 00frac34 S


27 szlig 55frac34 E ) none has been recorded during extensive trapping in coastal sand forestin the north of KwaZulu-Natal (Van Rensburg et al 1999) However during veryrecent trapping (Davis Scholtz van Aarde and Chown unpublished ) Gyronotus

have been recorded in old coastal dune forest near Richards Bay (28 szlig 30ndash31 frac34 S32 szlig 23frac34 E ) in southern Maputaland Thus the South African species of Gyronotus

may be described as occurring in low-lying forests with either southern coastalnorthern sand or afromontane a nities (Low and Rebelo 1996)

However according to a GIS overlay of vegetation-type distribution (Low andRebelo 1996) there was a poor correlation between Gyronotus grid references andeither coastal or afromontane forest Most localities (22 out of 26) coincided withbushveld (17) or grass (5) This included some known forest localities VernonCrookes Nature Reserve (30 szlig 16frac34 S 30 szlig 29frac34 E ) Dwesa Forest (32 szlig 15frac34 S 28 szlig 49frac34 E ) SilakaForest (31 szlig 33frac34 S 29 szlig 30frac34 E ) Woodbush Forest (23 szlig 52frac34 S 29 szlig 56frac34 E ) Others (3) coincidedwith coastal forest Ntsubane Forest (31 szlig 27frac34 S 29 szlig 44frac34 E ) Dwesa Forest (32 szlig 17frac34 E28 szlig 50frac34 E ) Only one coincided with afromontane forest Ngoye Forest (28 szlig 50frac34 S31 szlig 43frac34 E ) This may be due to various factors including inaccurate locality labelsandor inaccurate grid references in relation to the patchy occurrence of forestfragments Clearance of forest subsequent to collection may also have in uencedresults since a number of the localities are for material collected in the late 1800s orearly 1900s

Available habitat data and circumstantial evidence suggest that Gyronotus com-prise entirely forest specialists (table 3) Furthermore trap data for Gyronotus Wmet-

arius G pumilus and G mulanjensis suggest that these species are also microhabitatspecialists occurring in cool or cool wet spots within the forest Traps placed atvarious sites in the East Usambara Mts failed to record G Wmetarius outside offorest or at drier sites within forest It was recorded at only one site adjoining astream (215m) (Newmark Davis and Rickart unpublished ) Similar data fromgrassland and forest in Vernon Crookes Nature Reserve shows that G pumilus isrestricted to forest patches where it was trapped only at wet spots next to a streamor in a patch characterized by a high density of worm casts (Davis Scholtz andChown unpublished ) Furthermore one of only two localities for G mulanjensis

occurs along Muloza Stream on Mt Mulanje in Malawi However very recentrecords (Davis Scholtz van Aarde and Chown unpublished ) show that G carinatus

occurs in relatively cool coastal dune forest in southern Maputaland as well as on ner-grained soils in forests of the coastal hills

Gyronotus species also show geographical diŒerences in body shape These diŒer-ences may be related to climate since the elongate Gyronotus species are found intropical east and east-central Africa whereas the ovoid species are found in lowlandand low-lying afromontane forest in South Africa A statistical test of correlationbetween body shape measurements and climate cannot be attempted at present inthe absence of suitable forest temperature data However the proportionally deepestand most elongate species (G dispar) occurs in low altitude tropical forest of south-east Tanzania The proportionally least deep and least elongate tropical species (G

Wmetarius) occurs primarily in the cool wet forests of the East Usambara Mts Theproportionally attest most ovoid of the southern African species (G glabrosus)shows a northerly inland distribution in forests with afromontane a nities Theproportionally deepest least ovoid of the southern African species (G pumilus)occupies a southerly but coastal distribution


Distributional hypotheses

Burgess et al (1998) have suggested ve hypotheses to account for the distribu-tional patterns of biota in east African coastal forests which they de ne as occurringbetween Somalia and Mozambique encompassing the Swahili centre of endemismand the transitional region between the Swahili and Maputaland centres of endem-ism These hypotheses are equally applicable to the relict distribution of Gyronotus

which links the Swahili centre of endemism to those of Maputaland and Pondolandfurther to the south Hypothesis one suggests that distributional patterns result fromcollecting artifacts which may or may not account for the between ZambeziLimpopodisjunction in Gyronotus records Hypothesis two suggests that climatic history isresponsible for distributional patterns In Gyronotus combined tectonic and orogenicin uences on climatic history are probably responsible for the isolation of theirancestors in eastern forests during the Miocene and perhaps for later north-southspecies group separation Hypothesis three suggests that vicariant species evolutionis responsible for distribution patterns In Gyronotus some within-group speciationmay result from more recent forest fragmentation during the Pleistocene InGyronotus it is probable that human disturbance (hypothesis four) is responsiblefor increasing fragmentation of range localization of survivors and threat to theirsurvival F inally hypothesis ve suggests past sea-level changes have in uenceddistributional patterns In Gyronotus the distinctive relict distribution pattern at themargin of coastal lowlands might possibly be in uenced by recent change in coastlineposition due to PlioPleistocene intensi cation of polar glaciation and subsequentmarine regression These hypotheses are discussed in more detail below

Gyronotus and south-east African historical and vicariance events

The geographical distribution of Gyronotus may describe the relict of an oldtropical lowland distribution pattern which links wet tropical lowland forest of EastAfrica (Chapman and White 1970 Lovett 1993) to low altitude forest with coastalor afromontane a nities (Low and Rebelo 1996) in South Africa The northernlimits of their polarized distribution lie in the East Usambara Mts whose age isuncertain but probably originated from 290ndash180 MY ago with further uplift at7 MY (Gri ths 1993) The East Usambaras comprise part of the ancient easternarc mountain system which is the eastern-most of ve principal centres of forestbiodiversity stretching across central Africa (Hamilton 1989) Their exclusion fromthe East African highlands Ethiopia and Somalia is probably related to the recentPlio-Pleistocene uplift of the East African mountains (Baker and Wollenberg 1971Gri ths 1993) and increasing aridity of the coastal regions In these more northerlyregions canthonines are represented only by the species of the two most widespreadgenera Anachalcos Hope and Odontoloma Boheman (Ferreira 1972 Howden andScholtz 1987) The southern limits of Gyronotus distribution lies at the southernmargins of subtropical climate in South Africa Their exclusion from the more south-westerly winter and bimodal rainfall regions would be related to the northwardexpansion of the westerly system of air currents in the Pliocene (3 MY ) whichresulted in cooler climate (Deacon 1983 Deacon et al 1992) This event would beresponsible for the geographically distinctive nature (Griswold 1991 Davis 1997)and high level of endemism (Davis 1993 1994 Picker and Samways 1996) in thearthropod fauna of the Western and Eastern Cape where canthonines have radiatedinto non-forest habitats There is a wide separation between east African andsouthern African Gyronotus species in geographical phenetic and systematic terms


( gures 1 4 table 3) Between the old barriers of the broad hot dry valleys of theLimpopo and the Zambezi rivers their apparent non-occurrence may be due toeither actual absence or absence of collection It is predicted that a survey of forestson the lower eastern slopes of the East Zimbabwecentral west Mozambique high-lands might discover further Gyronotus species assuming that such forests remainextant

The tropical Gyronotus species of east Africa mostly occur in lowland forests(below 1370m in MalawimdashChapman and White 1970 below 800m in the easternarcmdashLovett 1993) These east African forests are associated with the orographiccapture of moisture by Eastern Arc and Malawi Rift mountains or with condensationof maritime or lacustrine evaporation (Gillman 1949) Therefore they would besensitive to cyclic climatic oscillation from warmer wetter climate during interglacialsand cooler drier climate during glacials of the Pleistocene There is a continuingdebate on the eŒects of Pleistocene expansion and contraction cycles in East Africanforests (Lawes 1990 Bruhl 1997 Roy 1997) Pre-Pleistocene events are probablyresponsible for most of the distribution patterns recorded for arthropods in afromon-tane forests (Griswold 1991 Bruhl 1997) although montane regions have beenidenti ed as centres of Pleistocene speciation in afromontane birds with older ances-tral species occurring in the adjoining lowlands (Roy 1997) Some within regiondivisions between Gyronotus species may also be related to recent vicariance eventsAlthough recent northsouth separation between the east African G Wmetarius andG mulanjensis is supported by their closely-similar aedeagi it is not supported byany of the inconsistent results shown by the cladograms based on phenetic data Aclose relationship between G carinatus and G glabrosus in South Africa is supportedby both character states of the aedeagi and by phenetic data However separationbetween the two species would predate the late Pleistocene (before 100000BP) asreconstruction of past forest distribution (Lawes 1990) suggests that there havebeen no recent links between the forests of Northern Province and Zululand

Acknowledgements

The authors thank Mr Riaan Stals (National Collection of Insects Pretoria)Dr Hamish Robertson (South African Museum Cape Town) Mr Malcolm Kerley(The Natural History Museum London) Dr Cornell Dudley (Museums of MalawiChichiri Blantyre) Dr Tanza Crouch (Durban Natural Science Museum Durban)Dr Frederick Gess (Albany Museum Grahamstown) Dr John Irish (NasionaleMuseum Bloemfontein) the late Dr Sebastian Endrody-Younga (TransvaalMuseum Pretoria) Dr Manfred Uhlig (Museum fur Naturkunde HumboldtUniversitat Berlin) and Dr Lothar Zerche (Deutsches Entomologisches InstitutEberswalde) for the loan of Gyronotus material Dr Paulette Bloomer and MsBettine Jansen van Vuuren kindly analysed the cladistic matrices whereas Mr HeathHull kindly produced the GIS distribution map for south-eastern South Africa andMs Marguerite Pienaar did the drawings We also thank Mr John Crawford-Brunt(SAF COL) for permission to place traps for Gyronotus in Woodbush indigenousforest

References

Baker B H and Wollenberg J 1971 Structure and evolution of the Kenya rift valleyNature 229 538ndash542


Barnes R F W 1990 Deforestation trends in tropical Africa African Journal of Ecology28 161ndash173

Bruhl C A 1997 F lightless insectsmdasha test-case for historical relationships of Africanmountains Journal of Biogeography 24 233ndash250

Burgess N D Clarke G P and Rodgers W A 1998 Coastal forests of EasternAfricamdashstatus endemism patterns and their potential causes Biological Journal of theLinnean Society 64 337ndash367

Cambefort Y 1991 Biogeography and evolution in I Hanski and Y Cambefort (eds)Dung Beetle Ecology (Princeton Princeton University Press) pp 51ndash67

Chapill J A 1989 Quantitative characters in phylogenetic analysis Cladistics 5 217ndash234Chapman J D and White F 1970 The Evergreen Forests of Malawi (Oxford

Commonwealth Forestry Institute)Davis A L V 1993 Biogeographical groups in a southern African winter rainfall dung

beetle assemblage (Coleoptera Scarabaeidae)mdashconsequences of climatic history andhabitat fragmentation African Journal of Ecology 31 306ndash327

Davis A L V 1994 Habitat fragmentation in southern Africa and distributional responsepatterns in ve specialist or generalist dung beetle families (Coleoptera) African Journalof Ecology 32 192ndash207

Davis A L V 1997 Climatic and biogeographical associations of southern African dungbeetles (Coleoptera Scarabaeidae s str) African Journal of Ecology 35 10ndash38

Davis A L V Scholtz C H and Harrison J Du G 1999 New and threatenedAfrotropical dung beetle taxa in the Gondwanaland tribe Canthonini (ColeopteraScarabaeidae Scarabaeinae) African Entomology 7 77ndash84

Deacon H J 1983 An introduction to the fynbos region time scales and palaeoenviromentsin H J Deacon Q B Hendey and J J Lambrechts (eds) Fynbos Palaeoecology APreliminary Synthesis South African National Scienti c Report no 75 1ndash20(Pretoria CSIR )

Deacon H J Jury M R and Ellis F 1992 Selective regime and time in R M Cowling(ed) The Ecology of Fynbos Nutrients Fire and Diversity (Cape Town OxfordUniversity Press) pp 6ndash22

Fairbairn D J 1992 The origins of allometry size and shape polymorphism in the commonwaterstrider Gerris remigis Say (Heteroptera Gerridae) Biological Journal of theLinnaean Society 45 167ndash186

Felsenstein J 1993 PHYLIP Phylogeny Inference Package Version 35 (Seattle Universityof Washington)

Ferreira M C 1972 Os escarabotildedeos de Africa (sul do Saara) Revista Entomologica deMocEuml ambique 11 5ndash1088

G illman C 1949 A vegetation-types map of Tanganyika Territory The GeographicalReview 39 7ndash37

Griffiths C J 1993 The geological evolution of east Africa in J C Lovett and S KWasser (eds) Biogeography and Ecology of the Rainforests of Eastern Africa (CambridgeCambridge University Press) pp 9ndash21

Griswold C E 1991 Cladistic biogeography of afromontane spiders Australian SystematicBotany 4 73ndash89

Halffter G 1974 Elements anciens de lrsquoentomofaune neotropical ses implications biogeog-raphiques Quaestiones Entomologicae 10 223ndash262

Halffter G and Matthews E G 1966 The natural history of dung beetles of thesubfamily Scarabaeinae (Coleoptera Scarabaeidae) Folia EntomoloAcirc gica Mexicana12plusmn 14 1ndash312

Hamilton A C 1989 Afromontane forests in H Lieth and M J A Werger (eds) Ecosystemsof the World 14B Tropical Rain Forest Ecosystems (Amsterdam Elsevier) pp 155ndash182

Howden H F and Scholtz C H 1987 A revision of the African genus OdontolomaBoheman (Coleoptera Scarabaeidae Scarabaeinae) Journal of the EntomologicalSociety of Southern Africa 50 155ndash192

InternationalUnion for the Conservation of Nature Species Survival Commission 1994IUCN Red List Categories as Approved by the 40th Meeting of the IUCN CouncilGland Switzerland (Gland The World Conservation Union)


Lawes M J 1990 The distribution of the Samango monkey (Cercopithecus mitis erythrarchusPeters 1852 and Cercopithecus mitis labiatus I GeoŒroy 1843) and forest history insouthern Africa Journal of Biogeography 17 669ndash680

Lovett J C 1993 Eastern Arc moist forest ora in J C Lovett and S K Wasser (eds)Biogeography and Ecology of the Rainforests of Eastern Africa (Cambridge CambridgeUniversity Press) pp 33ndash55

Low A B and Rebelo A G 1996 Vegetation of South Africa Lesotho and Swaziland(Pretoria Department of Environmental AŒairs and Tourism)

Picker M D and Samways M J 1996 Faunal diversity and endemicity of the CapePeninsula South Africamdasha rst assessment Biodiversity and Conservation 5 591ndash606

Prell W L Hutson W H Williams D F BeAacute A W H Geitzenhauer K andMalfino B 1980 Surface circulation of the Indian Ocean during the last glacialmaximum approximately 18000yr BP Quaternary Research 4 309ndash336

Roy M S 1997 Recent diversi cation in African greenbuls (Pycnotidae Andropadus)supports a montane speciation model Proceedings of the Royal Society of LondonSeries B 264 1337ndash1344

Scholtz C H and Howden H F 1987 A revision of the southern African Canthonina(Coleoptera Scarabaeidae Scarabaeinae) Journal of the Entomological Society ofSouthern Africa 50 75ndash119

Somers K M 1989 Allometry isometry and shape in principal components analysisSystematic Zoology 38 169ndash173

Swofford D L 1993 PAUP Phylogenetic Analysis Using Parsimony Version 31(Champaign Illinois Natural History Survey)

Thorpe R S 1984 Coding morphometric characters for constructing distance Wagnernetworks Evolution 38 244ndash255

Van Rensburg B McGeoch M A Chown S L and Van Jaarsveld A S 1999Conservation of heterogeneity among dung beetles in the Maputaland centre ofendemism South Africa Biological Conservation 88 145ndash153

Walter H and Lieth H 1964 Klimadiagramm-Weltatlas (Jena Gustav F ischer)Wiley E O 1981 Phylogenetics The Theory and Practice of Phylogenetic Systematics (New

York J Wiley amp Sons)


East Usambara Mountains in north-eastern Tanzania (Scholtz and Howden 1987)where they are primarily found in forest habitats (Davis unpublished ) These 11genera may be regarded as relicts since they belong to a tribe with ancientGondwanaland a liations (HalŒter and Matthews 1966 HalŒter 1974 Cambefort1991) and are all species-poor This relictual distribution pattern is presumablyrelated to historical events particularly to the Miocene and Plio-Pleistocene upliftin east central Africa which separated the forests of eastern Africa from those inWest Africa (Lovett 1993 Burgess et al 1998) In East Africa survival of wetrainforest and its associated faunal relicts may be a consequence of the relativelystable surface temperatures (0ndash2 szlig C variation) in the Indian Ocean (Prell et al 1980)and continued coastal and orographic capture of high rainfall (Gillman 1949)during the Pleistocene cycles of cool dry and warm wet climate

The genus Gyronotus is one of the most species-rich of these eastern seaboardforest relicts It comprises six known species showing disjunct or parapatric distribu-tions (Scholtz and Howden 1987 Davis et al 1999) extending from the marginsof the bimodal rainfall region in the Eastern Cape South Africa (Davis 1993 1997)to the East Usambara Mountains which lie towards the northern end of the easternarc mountain system in north-eastern Tanzania All Gyronotus species are ightlessand are known to occur predominantly on ner-grained soils in lower altitude forestswith tropical or afromontane a nities Such ightless taxa with their low vagilityhave been shown to be most useful indicators of evolutionary and biogeographicalhistory (Bruhl 1997)

Both northern tropical and southern temperate (Palaeantarctic) Gondwanalandorigins have been hypothesized for afrotropical members of the tribe Canthonini(HalŒter 1974) However subsequent intra-continental dispersal vicariance eventsand extinctions have presumably shaped the modern relictual distribution patternsof the genera Phenetic cladistic and morphological analyses were conducted onGyronotus in order to examine their interspeci c relationships in comparison to thebiogeographical a nities of the species The results suggest a tropical generic a li-ation with a change in body shape from the lowland forest of East Africa to thelowland and low-lying afromontane forests of South Africa These geographicaldistribution patterns are discussed in an historical context

The ndings describe the relict distribution of an old dung beetle lineage ineastern Afrotropical forests In the present day these eastern forest habitats comprisean archipelago (Hamilton 1989 Low and Rebelo 1996 Burgess et al 1998) ofboth natural and anthropogenically-induced fragments which are seriouslythreatened by continuing exploitation Barnes (1990) predicted that these easternAfrotropical forests would be reduced by 95 by the year 2040 However recentanalyses suggest that the coastal portion between Somalia and Mozambique isalready reduced to a few thousand square kilometres of small fragments (Burgesset al 1998) Furthermore at least one of the species of Gyronotus dung beetles inthe present study (G glabrosus Scholtz and Howden) already deserves criticallyendangered (CR) status ( IUCN 1994) and may well be extinct (EX ) (Daviset al 1999)

Methods

Geographical distribution data for Gyronotus were derived from material heldby the University of Pretoria (now lodged in the Transvaal Museum Pretoria) andmaterial loaned by seven southern African and three European museums and



























Tab

le2

Div

erge

nce

mat

rice

s(T

horp

e19

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for

dist

ance

and

clad

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anal

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betw

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Gyro

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Ch

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12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

27

Der

ived

fro

mdo

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-sta

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dize

dra

wph

enet

icm

easu

rem

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(mm

)C

har

in

form

ativ

eDaggeri

ii

ii

ii

ii

ii

G

pum

ilus

58

46

87

48

86

38

65

45

64

36

26

45

21

4G

ca

rinatu

s1

84

68

85

88

42

86

24

47

28

02

53

88

70

G

gla

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18

46

87

58

84

28

63

44

71

21

42

39

97

4G

dis

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a3

a7

20

40

09

80

5a

aa

08

6a

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30

37

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34

42

57

32

37

26

44

46

78

66

a9

43

14

G

Wm

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us

80

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23

53

44

84

16

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36

66

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57

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Der

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Ch

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88

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75

77

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88

84

36

46

67

69

13

43

89

80

G

gla

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17

87

74

38

85

34

65

66

71

32

30

28

98

6G

dis

par

97

96

75

a0

23

54

65

56

16

37

99

93

33

5G

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31

36

66

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66

68

69

99

33

26

G

Wm

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90

00

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33

8a

04

20

37

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8

dagger Rec

oded

fro

ma

rang

eof

Otilde5

to5

toa

rang

efr

om

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10(1

0re

cod

edas

lsquoarsquo

wh

ich

isre

adas

one

unit

grea

ter

than

9in

the

dist

ance

anal

ysis

)Dagger A

llch

arac

ters

used

for

dist

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anal

ysis

on

lyin

form

ativ

ech

arac

ters

(i)

used

for

pars

imo

nyan

alys

is



(Davis et al 1999)

Results











Tab

le3

Sum

mar

yof

know

nge

ogra

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aldi

stri

but

ion

and

habi

tat

asso

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ion

sof

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Sout

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Tan

zani

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Len

gth

Geo

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Alt

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s(m

m)dagger

(pat

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)H

abit

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Gyro

notu

sva

nL

ansb

erge

1874

South

Afr

ica

All

know

nre

cord

sin

fore

stG

pum

ilus

(Boh

eman

1857

)13

ndash15

Sout

h-ea

stco

asta

lhi

lls50

ndash450

For

est

469

of

128

info

rest

(gu

res

12

)G

ca

rinatu

sF

elsc

he19

1113

ndash15

Zul

ula

nd

coas

tal

hills

50ndash5

00F

ores

t5

7of

53in

fore

st(

gure

s1

2)

G

gla

bro

sus

Scho

ltz

and

Ho

wde

n19

8712

ndash12

5D

rak

ensb

erg

esca

rpm

ent

in78

0F

ores

t33

of

6in

fore

stN

ort

hern

Pro

vinc

e(

gure

s1

2)

Tro

pic

al

Afr

ica

Gm

ula

nje

nsi

sD

avis

Sch

olt

zan

dH

arri

son

118

ndash13

3M

tM

ulan

je

Mal

awi

(gu

re1

)90

0ndash10

00F

ores

tN

ot

reco

rded

F

ores

tan

d19

99op

enco

llect

ion

sm

ixed

DaggerG

Wm

etari

us

Ko

lbe

1894

125

ndash15

5M

ain

lyE

ast

Usa

mba

raM

ts50

ndash850

For

est

Fou

rtr

appe

dby

stre

amin

plu

sea

stco

ast

ofT

anza

nia

fore

stbu

tno

neou

tsid

e(

gure

1)

G

dis

par

(Fel

sche

1911

)16

ndash18

Sout

h-ea

stT

anza

nia

(gu

re1

)50

For

est

No

habi

tat

note

s

dagger Len

gth

wit

hhe

adde

exe

das

oppo

sed

tora

ised

asin

tabl

e5

Dagger C

L

Bel

lam

y(p

erso

nal

com

mun

icat

ion)











Discussion




Tab

le4

The

corr

elat

ion

ofG

yro

notu

sbo

dysi

ze(m

axim

umle

ngt

h)1

and

body

shap

e(r

atio

max

imu

mle

ngt

hm

axim

um

wid

thof

elyt

ra)2

wit

hor

din

ates

for

Fac

tors

1an

d2

deri

ved

fro

mpr

inci

pal

com

pon

ents

anal

ysis

ofm

orp

hom

etri

csda

tatr

eate

din

two

diŒ

eren

tw

ays

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

raw

mea

sure

men

tsra

wm

easu

rem

ents

log 10

mea

sure

men

tslo

g 10m

easu

rem

ents

(27

char

acte

rs)

(11

char

acte

rs)

(27

char

acte

rs)

(14

char

acte

rs)

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Pro

por

tion

ofov

eral

lvar

ianc

e(

)55

217

756

015

058

313

266

413

6R

2fo

rre

gres

sio

nof

body

size

on0

980

0002

092

000

20

060

230

080

24or

din

ates

1F

(15

1)fo

rre

gres

sio

nof

body

size

2870

60

0

0162

106

008

345

150

9

437

16

38

on

ordi

nat

es1

R2

for

regr

essi

on

ofbo

dysh

ape

on0

620

130

670

060

150

250

140

30or

din

ates

2F

(15

1)fo

rre

gres

sio

nof

body

shap

e83

55

7

55

103

79

3

358

63

173

1

873

22

26

on

ordi

nat

es2

p

lt0

05

p

lt0

02

p

lt0

001


Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References


































































Tab

le2

Div

erge

nce

mat

rice

s(T

horp

e19

84)dagger

for

dist

ance

and

clad

isti

cpa

rsim

ony

anal

ysis

ofin

ter-

rela

tio

nshi

ps

betw

een

Gyro

notu

ssp

ecie

s

Ch

arac

ter

12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

27

Der

ived

fro

mdo

uble

-sta

ndar

dize

dra

wph

enet

icm

easu

rem

ents

(mm

)C

har

in

form

ativ

eDaggeri

ii

ii

ii

ii

ii

G

pum

ilus

58

46

87

48

86

38

65

45

64

36

26

45

21

4G

ca

rinatu

s1

84

68

85

88

42

86

24

47

28

02

53

88

70

G

gla

bro

sus

18

46

87

58

84

28

63

44

71

21

42

39

97

4G

dis

par

a3

a7

20

40

09

80

5a

aa

08

6a

a1

30

37

9G

m

ula

nje

nsi

s5

34

42

57

32

37

26

44

46

78

66

a9

43

14

G

Wm

etari

us

80

41

23

53

44

84

16

43

48

36

66

84

57

9

Der

ived

fro

mdo

uble

-sta

ndar

dize

dlo

g 10tr

ansf

orm

edra

wm

easu

rem

ents

Ch

ar

info

rmat

iveDagger

ii

ii

ii

ii

ii

ii

ii

G

pum

ilus

57

46

75

38

88

43

68

76

64

47

34

35

32

5G

ca

rinatu

s1

75

77

a4

88

84

36

46

67

69

13

43

89

80

G

gla

bro

sus

17

87

74

38

85

34

65

66

71

32

30

28

98

6G

dis

par

97

96

75

a0

23

54

65

56

16

37

99

93

33

5G

m

ula

nje

nsi

s5

24

42

57

31

36

66

44

66

68

69

99

33

26

G

Wm

etari

us

90

00

01

33

33

8a

04

20

37

37

34

43

37

8

dagger Rec

oded

fro

ma

rang

eof

Otilde5

to5

toa

rang

efr

om

0to

10(1

0re

cod

edas

lsquoarsquo

wh

ich

isre

adas

one

unit

grea

ter

than

9in

the

dist

ance

anal

ysis

)Dagger A

llch

arac

ters

used

for

dist

ance

anal

ysis

on

lyin

form

ativ

ech

arac

ters

(i)

used

for

pars

imo

nyan

alys

is



(Davis et al 1999)

Results











Tab

le3

Sum

mar

yof

know

nge

ogra

phic

aldi

stri

but

ion

and

habi

tat

asso

ciat

ion

sof

Gyro

notu

ssp

pbe

twee

nth

eE

aste

rnC

ape

Sout

hA

fric

aan

dno

rth

-eas

tern

Tan

zani

a

Len

gth

Geo

grap

hic

alra

nge

Alt

itu

din

alG

enu

san

dsp

ecie

s(m

m)dagger

(pat

chy

occu

rren

ce)

rang

e(m

)H

abit

atN

ote

s

Gyro

notu

sva

nL

ansb

erge

1874

South

Afr

ica

All

know

nre

cord

sin

fore

stG

pum

ilus

(Boh

eman

1857

)13

ndash15

Sout

h-ea

stco

asta

lhi

lls50

ndash450

For

est

469

of

128

info

rest

(gu

res

12

)G

ca

rinatu

sF

elsc

he19

1113

ndash15

Zul

ula

nd

coas

tal

hills

50ndash5

00F

ores

t5

7of

53in

fore

st(

gure

s1

2)

G

gla

bro

sus

Scho

ltz

and

Ho

wde

n19

8712

ndash12

5D

rak

ensb

erg

esca

rpm

ent

in78

0F

ores

t33

of

6in

fore

stN

ort

hern

Pro

vinc

e(

gure

s1

2)

Tro

pic

al

Afr

ica

Gm

ula

nje

nsi

sD

avis

Sch

olt

zan

dH

arri

son

118

ndash13

3M

tM

ulan

je

Mal

awi

(gu

re1

)90

0ndash10

00F

ores

tN

ot

reco

rded

F

ores

tan

d19

99op

enco

llect

ion

sm

ixed

DaggerG

Wm

etari

us

Ko

lbe

1894

125

ndash15

5M

ain

lyE

ast

Usa

mba

raM

ts50

ndash850

For

est

Fou

rtr

appe

dby

stre

amin

plu

sea

stco

ast

ofT

anza

nia

fore

stbu

tno

neou

tsid

e(

gure

1)

G

dis

par

(Fel

sche

1911

)16

ndash18

Sout

h-ea

stT

anza

nia

(gu

re1

)50

For

est

No

habi

tat

note

s

dagger Len

gth

wit

hhe

adde

exe

das

oppo

sed

tora

ised

asin

tabl

e5

Dagger C

L

Bel

lam

y(p

erso

nal

com

mun

icat

ion)











Discussion




Tab

le4

The

corr

elat

ion

ofG

yro

notu

sbo

dysi

ze(m

axim

umle

ngt

h)1

and

body

shap

e(r

atio

max

imu

mle

ngt

hm

axim

um

wid

thof

elyt

ra)2

wit

hor

din

ates

for

Fac

tors

1an

d2

deri

ved

fro

mpr

inci

pal

com

pon

ents

anal

ysis

ofm

orp

hom

etri

csda

tatr

eate

din

two

diŒ

eren

tw

ays

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

raw

mea

sure

men

tsra

wm

easu

rem

ents

log 10

mea

sure

men

tslo

g 10m

easu

rem

ents

(27

char

acte

rs)

(11

char

acte

rs)

(27

char

acte

rs)

(14

char

acte

rs)

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Pro

por

tion

ofov

eral

lvar

ianc

e(

)55

217

756

015

058

313

266

413

6R

2fo

rre

gres

sio

nof

body

size

on0

980

0002

092

000

20

060

230

080

24or

din

ates

1F

(15

1)fo

rre

gres

sio

nof

body

size

2870

60

0

0162

106

008

345

150

9

437

16

38

on

ordi

nat

es1

R2

for

regr

essi

on

ofbo

dysh

ape

on0

620

130

670

060

150

250

140

30or

din

ates

2F

(15

1)fo

rre

gres

sio

nof

body

shap

e83

55

7

55

103

79

3

358

63

173

1

873

22

26

on

ordi

nat

es2

p

lt0

05

p

lt0

02

p

lt0

001


Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References










































(Davis et al 1999)

Results











Tab

le3

Sum

mar

yof

know

nge

ogra

phic

aldi

stri

but

ion

and

habi

tat

asso

ciat

ion

sof

Gyro

notu

ssp

pbe

twee

nth

eE

aste

rnC

ape

Sout

hA

fric

aan

dno

rth

-eas

tern

Tan

zani

a

Len

gth

Geo

grap

hic

alra

nge

Alt

itu

din

alG

enu

san

dsp

ecie

s(m

m)dagger

(pat

chy

occu

rren

ce)

rang

e(m

)H

abit

atN

ote

s

Gyro

notu

sva

nL

ansb

erge

1874

South

Afr

ica

All

know

nre

cord

sin

fore

stG

pum

ilus

(Boh

eman

1857

)13

ndash15

Sout

h-ea

stco

asta

lhi

lls50

ndash450

For

est

469

of

128

info

rest

(gu

res

12

)G

ca

rinatu

sF

elsc

he19

1113

ndash15

Zul

ula

nd

coas

tal

hills

50ndash5

00F

ores

t5

7of

53in

fore

st(

gure

s1

2)

G

gla

bro

sus

Scho

ltz

and

Ho

wde

n19

8712

ndash12

5D

rak

ensb

erg

esca

rpm

ent

in78

0F

ores

t33

of

6in

fore

stN

ort

hern

Pro

vinc

e(

gure

s1

2)

Tro

pic

al

Afr

ica

Gm

ula

nje

nsi

sD

avis

Sch

olt

zan

dH

arri

son

118

ndash13

3M

tM

ulan

je

Mal

awi

(gu

re1

)90

0ndash10

00F

ores

tN

ot

reco

rded

F

ores

tan

d19

99op

enco

llect

ion

sm

ixed

DaggerG

Wm

etari

us

Ko

lbe

1894

125

ndash15

5M

ain

lyE

ast

Usa

mba

raM

ts50

ndash850

For

est

Fou

rtr

appe

dby

stre

amin

plu

sea

stco

ast

ofT

anza

nia

fore

stbu

tno

neou

tsid

e(

gure

1)

G

dis

par

(Fel

sche

1911

)16

ndash18

Sout

h-ea

stT

anza

nia

(gu

re1

)50

For

est

No

habi

tat

note

s

dagger Len

gth

wit

hhe

adde

exe

das

oppo

sed

tora

ised

asin

tabl

e5

Dagger C

L

Bel

lam

y(p

erso

nal

com

mun

icat

ion)











Discussion




Tab

le4

The

corr

elat

ion

ofG

yro

notu

sbo

dysi

ze(m

axim

umle

ngt

h)1

and

body

shap

e(r

atio

max

imu

mle

ngt

hm

axim

um

wid

thof

elyt

ra)2

wit

hor

din

ates

for

Fac

tors

1an

d2

deri

ved

fro

mpr

inci

pal

com

pon

ents

anal

ysis

ofm

orp

hom

etri

csda

tatr

eate

din

two

diŒ

eren

tw

ays

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

raw

mea

sure

men

tsra

wm

easu

rem

ents

log 10

mea

sure

men

tslo

g 10m

easu

rem

ents

(27

char

acte

rs)

(11

char

acte

rs)

(27

char

acte

rs)

(14

char

acte

rs)

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Pro

por

tion

ofov

eral

lvar

ianc

e(

)55

217

756

015

058

313

266

413

6R

2fo

rre

gres

sio

nof

body

size

on0

980

0002

092

000

20

060

230

080

24or

din

ates

1F

(15

1)fo

rre

gres

sio

nof

body

size

2870

60

0

0162

106

008

345

150

9

437

16

38

on

ordi

nat

es1

R2

for

regr

essi

on

ofbo

dysh

ape

on0

620

130

670

060

150

250

140

30or

din

ates

2F

(15

1)fo

rre

gres

sio

nof

body

shap

e83

55

7

55

103

79

3

358

63

173

1

873

22

26

on

ordi

nat

es2

p

lt0

05

p

lt0

02

p

lt0

001


Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References














































Tab

le3

Sum

mar

yof

know

nge

ogra

phic

aldi

stri

but

ion

and

habi

tat

asso

ciat

ion

sof

Gyro

notu

ssp

pbe

twee

nth

eE

aste

rnC

ape

Sout

hA

fric

aan

dno

rth

-eas

tern

Tan

zani

a

Len

gth

Geo

grap

hic

alra

nge

Alt

itu

din

alG

enu

san

dsp

ecie

s(m

m)dagger

(pat

chy

occu

rren

ce)

rang

e(m

)H

abit

atN

ote

s

Gyro

notu

sva

nL

ansb

erge

1874

South

Afr

ica

All

know

nre

cord

sin

fore

stG

pum

ilus

(Boh

eman

1857

)13

ndash15

Sout

h-ea

stco

asta

lhi

lls50

ndash450

For

est

469

of

128

info

rest

(gu

res

12

)G

ca

rinatu

sF

elsc

he19

1113

ndash15

Zul

ula

nd

coas

tal

hills

50ndash5

00F

ores

t5

7of

53in

fore

st(

gure

s1

2)

G

gla

bro

sus

Scho

ltz

and

Ho

wde

n19

8712

ndash12

5D

rak

ensb

erg

esca

rpm

ent

in78

0F

ores

t33

of

6in

fore

stN

ort

hern

Pro

vinc

e(

gure

s1

2)

Tro

pic

al

Afr

ica

Gm

ula

nje

nsi

sD

avis

Sch

olt

zan

dH

arri

son

118

ndash13

3M

tM

ulan

je

Mal

awi

(gu

re1

)90

0ndash10

00F

ores

tN

ot

reco

rded

F

ores

tan

d19

99op

enco

llect

ion

sm

ixed

DaggerG

Wm

etari

us

Ko

lbe

1894

125

ndash15

5M

ain

lyE

ast

Usa

mba

raM

ts50

ndash850

For

est

Fou

rtr

appe

dby

stre

amin

plu

sea

stco

ast

ofT

anza

nia

fore

stbu

tno

neou

tsid

e(

gure

1)

G

dis

par

(Fel

sche

1911

)16

ndash18

Sout

h-ea

stT

anza

nia

(gu

re1

)50

For

est

No

habi

tat

note

s

dagger Len

gth

wit

hhe

adde

exe

das

oppo

sed

tora

ised

asin

tabl

e5

Dagger C

L

Bel

lam

y(p

erso

nal

com

mun

icat

ion)











Discussion




Tab

le4

The

corr

elat

ion

ofG

yro

notu

sbo

dysi

ze(m

axim

umle

ngt

h)1

and

body

shap

e(r

atio

max

imu

mle

ngt

hm

axim

um

wid

thof

elyt

ra)2

wit

hor

din

ates

for

Fac

tors

1an

d2

deri

ved

fro

mpr

inci

pal

com

pon

ents

anal

ysis

ofm

orp

hom

etri

csda

tatr

eate

din

two

diŒ

eren

tw

ays

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

raw

mea

sure

men

tsra

wm

easu

rem

ents

log 10

mea

sure

men

tslo

g 10m

easu

rem

ents

(27

char

acte

rs)

(11

char

acte

rs)

(27

char

acte

rs)

(14

char

acte

rs)

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Pro

por

tion

ofov

eral

lvar

ianc

e(

)55

217

756

015

058

313

266

413

6R

2fo

rre

gres

sio

nof

body

size

on0

980

0002

092

000

20

060

230

080

24or

din

ates

1F

(15

1)fo

rre

gres

sio

nof

body

size

2870

60

0

0162

106

008

345

150

9

437

16

38

on

ordi

nat

es1

R2

for

regr

essi

on

ofbo

dysh

ape

on0

620

130

670

060

150

250

140

30or

din

ates

2F

(15

1)fo

rre

gres

sio

nof

body

shap

e83

55

7

55

103

79

3

358

63

173

1

873

22

26

on

ordi

nat

es2

p

lt0

05

p

lt0

02

p

lt0

001


Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References


















































Discussion




Tab

le4

The

corr

elat

ion

ofG

yro

notu

sbo

dysi

ze(m

axim

umle

ngt

h)1

and

body

shap

e(r

atio

max

imu

mle

ngt

hm

axim

um

wid

thof

elyt

ra)2

wit

hor

din

ates

for

Fac

tors

1an

d2

deri

ved

fro

mpr

inci

pal

com

pon

ents

anal

ysis

ofm

orp

hom

etri

csda

tatr

eate

din

two

diŒ

eren

tw

ays

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

raw

mea

sure

men

tsra

wm

easu

rem

ents

log 10

mea

sure

men

tslo

g 10m

easu

rem

ents

(27

char

acte

rs)

(11

char

acte

rs)

(27

char

acte

rs)

(14

char

acte

rs)

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Pro

por

tion

ofov

eral

lvar

ianc

e(

)55

217

756

015

058

313

266

413

6R

2fo

rre

gres

sio

nof

body

size

on0

980

0002

092

000

20

060

230

080

24or

din

ates

1F

(15

1)fo

rre

gres

sio

nof

body

size

2870

60

0

0162

106

008

345

150

9

437

16

38

on

ordi

nat

es1

R2

for

regr

essi

on

ofbo

dysh

ape

on0

620

130

670

060

150

250

140

30or

din

ates

2F

(15

1)fo

rre

gres

sio

nof

body

shap

e83

55

7

55

103

79

3

358

63

173

1

873

22

26

on

ordi

nat

es2

p

lt0

05

p

lt0

02

p

lt0

001


Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References









































Tab

le4

The

corr

elat

ion

ofG

yro

notu

sbo

dysi

ze(m

axim

umle

ngt

h)1

and

body

shap

e(r

atio

max

imu

mle

ngt

hm

axim

um

wid

thof

elyt

ra)2

wit

hor

din

ates

for

Fac

tors

1an

d2

deri

ved

fro

mpr

inci

pal

com

pon

ents

anal

ysis

ofm

orp

hom

etri

csda

tatr

eate

din

two

diŒ

eren

tw

ays

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

Do

uble

-sta

ndar

dize

dD

oub

le-s

tand

ardi

zed

raw

mea

sure

men

tsra

wm

easu

rem

ents

log 10

mea

sure

men

tslo

g 10m

easu

rem

ents

(27

char

acte

rs)

(11

char

acte

rs)

(27

char

acte

rs)

(14

char

acte

rs)

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Fac

tor

1F

acto

r2

Pro

por

tion

ofov

eral

lvar

ianc

e(

)55

217

756

015

058

313

266

413

6R

2fo

rre

gres

sio

nof

body

size

on0

980

0002

092

000

20

060

230

080

24or

din

ates

1F

(15

1)fo

rre

gres

sio

nof

body

size

2870

60

0

0162

106

008

345

150

9

437

16

38

on

ordi

nat

es1

R2

for

regr

essi

on

ofbo

dysh

ape

on0

620

130

670

060

150

250

140

30or

din

ates

2F

(15

1)fo

rre

gres

sio

nof

body

shap

e83

55

7

55

103

79

3

358

63

173

1

873

22

26

on

ordi

nat

es2

p

lt0

05

p

lt0

02

p

lt0

001


Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References









































Tab

le5

Mor

pho

met

ric

mea

sure

men

tsfo

rsi

xsp

ecie

sof

Gyro

notu

s

Sout

hern

Afr

ican

spec

ies

Eas

tA

fric

ansp

ecie

sA

NO

VA

Mea

sure

men

tG

pum

ilus

G

cari

natu

sG

gla

bro

sus

G

dis

par

G

mula

nje

nsi

sG

Wm

etari

us

F(5

47)

Len

gth

(L)

147

2Ocirc0

64c

141

4Ocirc0

26cd

135

8Ocirc0

77d

206

4Ocirc0

41a

159

5Ocirc0

34b

162

0Ocirc0

26b

194

41

W

idth

(W)

905

Ocirc0

36b

900

Ocirc0

18b

944

Ocirc0

53b

107

4Ocirc0

42a

824

Ocirc0

22c

911

Ocirc0

46b

354

1

Dep

th(D

)6

02Ocirc

023

bd5

80Ocirc

021

cd5

57Ocirc

021

d8

24Ocirc

038

a6

09Ocirc

024

bc6

38Ocirc

027

b88

35

R

atio

sL

W0

62Ocirc

002

b0

64Ocirc

001

b0

70Ocirc

002

a0

52Ocirc

002

d0

52Ocirc

001

d0

56Ocirc

003

c87

16

L

D0

41Ocirc

001

a0

41Ocirc

001

a0

41Ocirc

002

a0

40Ocirc

002

ab0

38Ocirc

001

b0

39Ocirc

002

ab5

16

W

D0

67Ocirc

003

bc0

64Ocirc

002

c0

59Ocirc

002

d0

77Ocirc

004

ab0

74Ocirc

002

ab0

70Ocirc

003

b35

44

Sa

mp

lesi

ze10

106

710

10

Inea

chro

w

valu

esfo

llow

edby

adi

Œer

ent

lett

erdi

Œer

edsi

gni

cant

ly(T

ukey

rsquosH

SD

for

uneq

ual

nETH

plt

005

)

plt

000

1































Acknowledgements


References






































































Acknowledgements


References








































cladistic, phenetic and biogeographical analysis of the flightless dung beetle genus, gyronotus van...

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