Afr. J. Ecol. 1997, Volume 35, pages 10–38
Climatic and biogeographical associations of southern
African dung beetles (Coleoptera: Scarabaeidae s. str.)
ADRIAN L. V. DAVIS 1
Department of Zoology, University of Cape Town, Rondebosch, 7700 South Africa
Summary
Climatic and biogeographical associations of southern African dung beetles
(Scarabaeinae, Coprinae) were analysed from a collection amassed between 1971
and 1986. Endemism to Africa south of 15)S was much greater in southwesterly
climates (winter rainfall, bimodal spring/autumn rainfall, arid late summer
rainfall) than to the north-east in the moist, mid-summer rainfall region. Major
biogeographical groups centred to the south-west comprised predominantly
southern African endemics, Western/Eastern Cape coast, Karoo, Karoo/Namib,
northern Namibia and the south-western Kalahari. Biogeographical groups
centred on the south-eastern highlands and the subtropical east coast (midsummer rainfall) also comprised predominantly southern African endemics. All
other major groups centred to the north-east in the mid-summer rainfall region
comprised predominantly species with widespread tropical biogeographical
affiliations, pan-southern Africa but centred in the east, pan-mid-summer rainfall
region, eastern mid-summer rainfall region, tropical east Zimbabwe/central
Moçambique, subtropical/tropical game reserves (non-ruminant dung specialists). There were cross-climatic differences in taxonomic composition of the
fauna. Within the winter rainfall region, percentage species composition of
Scarabaeinae was greater whereas that of the coprine tribe, Onthophagini, was
lower than within the other three climatic regions. Percentage species richness of
most other tribes of Coprinae differed little between most climates but that of
Scarabaeinae declined from west to east (Canthonini, Scarabaeini), east to west
(Sisyphini) or to either side of the late summer rainfall region (Gymnopleurini).
Key words: Africa, biogeographical, climatic, dung, Scarabaeidae
Résumé
Les associations climatiques et biogéographique des bousiers (Scarabaeinae,
Coprinae) d’Afrique du sud furent analysées à parir d’une collection constituée
entre 1971 et 1986. L’endémisme en Afrique au sud de 15) était bien plus élevé
sous les climats du sud-ouest (pluies hivernales, pluies bimodales au printemps et
en automme, pluies tardives lors d’étés arides) qu’au nord-est dans la région
humide aux chutes de pluie de la mi-été. Des groupes biogéographiques majeurs
centrés au sud-ouest comprenaient principalement des endémiques d’Afrique
du sud, la côte du Cap Oriental et Occidental, L Karoo, L Karoo/Namib, la
1
Present address: Department of Zoology and Entomology, University of Pretoria, Pretoria 10002
South Africa
? 1997 East African Wild Life Society.
Associations of southern African dung beetles 11
Namibie septentrionale et le sud-ouest Kalahari. Des groupes biogéogrphiques
centrés sur les régions montagneuses du sud-est et la côte est subtropicale (chutes
de pluies de la mi-été) comprenaient également surtout des endémiques d’Afrique
du sud. Tous les autres groupes majeurs centrés au nord-est dans la région aux
chutes de pluie de la mi-été comprenaient principalement des espèces avec
affiliations biogéographiques tropicales étendue à l’ensemble de la région sud
africaine, mais centrées dans la région oriental à cutes de pluie de la mi-été, au
Zimbabwe oriental tropical/Mozambique central, et aux réserves de faune
sauvage subtropicales/tropicales (spécialistes des bouses de non-ruminants). On a
trouvé des différences dans la composition taxonomique de la faune quel que soit
le climat. Dans la région à chutes de pluie hivernales, le pourcentage d’espèces de
Scarabeinae dans la composition était plus grand alors que elle de la tribu des
coprinés, Ontophagini, était plus faible que dans les trois autres régions
climatiques. Le pourcentage de la richesse en espèces de la plupart des autres
tribus de Coprinae différait peu entre la plupart des climats mais celui des
Scarabaeinae diminuait d’ouest en est (Canthonini, Scarabaeini), d’est en ouest
(Sisyphini), ou en quelle que direction de la région à chutes de pluies estivales
tardives (Gymnopleurini).
Introduction
Dung-burying beetles (Scarabaeidae s. str.: Scarabaeinae, Coprinae) show a
relatively specialist spatial distribution (Davis, 1994a). Variables which influence
spatial specialization include climate (Kirk & Ridsdill-Smith, 1986; Davis &
Dewhurst, 1993), soil (Nealis, 1977; Cambefort, 1991a; Doube, 1991), vegetation
(Cambefort, 1982; Doube, 1983) and food type (Fincher et al., 1970; Davis,
1994b). This biogeographical study of southern African dung beetles considers
both the current and historical influence of these variables.
The prevailing climatic system over southern Africa developed in the late
Pliocene, c. three million years ago (Deacon, 1983). This system involves
interplay between three cells of air currents (Tyson, 1986) comprising the
rain-bearing westerlies centred over the south Atlantic Ocean, the rain-bearing
easterlies centred over the Indian Ocean and a cell of dry air centred over the
cold Benguela current in the Atlantic Ocean to the west of southern Africa.
During winter, the westerlies and the dry cell expand north-eastwards bringing a
winter peak in rainfall to the Western Cape, bimodal autumn and spring peaks
in rainfall to the Eastern Cape and dry season conditions to the remainder of
southern Africa. During summer, the easterlies expand sequentially southwestwards bringing a mid-summer peak in rainfall to the moist, easterly, climate
types and a late summer peak in rainfall to the arid, westerly, climate types.
Concurrent with the expansion of the easterlies, the cell of dry air contracts
southwards bringing dry summers to the winter rainfall region.
Geomorphologically, the subcontinent comprises a tilted plateau which is
bordered by coastal mountain ranges. A Permian (Deacon, 1983) or Triassic age
(Dingle et al., 1983; Tyson, 1986) is claimed for the Cape fold mountain system
in the Western and Eastern Cape whereas the south-eastern highlands are
younger and originate from the asymmetric raising and tilting westwards of the
plateau during the mid-Tertiary and the Pliocene. This second uplift is more or
less coeval with major climatic changes.
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
12
A. L. V. Davis
Fig. 1. Climatic regions of southern Africa after Walter & Lieth (1964) and modifications by Davis (1987).
Overall, the disposition of air currents and topographical features over the
subcontinent results in 25 different climatic regions (Walter & Lieth, 1964). These
climate types have been reduced to the four major climatic regions of winter,
bimodal, late summer and mid-summer rainfall by Davis (1987, 1993). In the
present study, the mid-summer rainfall region is divided into five subregions
(Fig. 1, Table 1).
Soil, vegetation and dung types show clear distribution patterns. A swathe
of late Cretaceous to mid-Tertiary sands (Tyson, 1986) dominate the central
Kalahari basin from the Northern Cape to southern Angola. Miocene to
Pliocene sands are also concentrated around the coastline. In contrast, finergrained soils dominate in the south-eastern highlands with a patchwork of
finer-grained soils and sand outliers elsewhere. Three principal vegetational
divisions (Werger, 1978) largely coincide with climatic regions. The shrublanddominated Cape floristic and Karoo/Namib divisions roughly coincide, respectively, with the winter/bimodal and the arid, late summer rainfall regions. The
highland grassland and lowland woodland of the southern Sudano-zambezian
division roughly coincide with the mid-summer rainfall region. Fragmentation in
the distribution of woody vegetation has occurred over much of the subcontinent
although it is particularly influential on dung beetle occurrence in the winter
rainfall region (Davis, 1993). The range of large indigenous mammals is also
fragmented so that high densities of non-ruminant dung are mostly restricted to
game reserves whereas ruminant dung remains widespread.
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 13
Table 1. Dung beetle collection data for the 25 climatic regions of southern Africa (Walter & Lieth, 1964)
south of 15)S
Number of collections on:
Soil type
Food type
Vegetation type
Number
Shrubland/
Non- Omnivore
Coarse
of
open
dung/ Pasture/
Climate
collection
Fine grained Ruminant ruminant
carrion grassland woodland Forest localities*
dung
type
dung
grained (sand)
Winter rainfall
III3
0
III4
5
III(IV)a
11
III(IV)b
8
IV
6
3
21
38
3
4
0
20
52
14
18
2
4
1
1
0
1
1
0
0
0
0
13
32
5
10
3
12
12
7
1
0
0
0
0
0
3
28
54
15
21
Bimodal rainfall
III5
29
IV(V)
31
V
22
Late summer rainfall
II4a
28
II(III)a
9
II(III)b
1
III1
6
III2
1
6
7
3
37
50
42
1
4
1
0
0
0
9
34
16
18
3
4
0
4
0
38
60
49
4
56
3
36
16
30
59
3
27
9
2
4
1
9
4
1
0
0
4
1
10
18
0
7
8
17
48
3
36
8
0
0
0
0
0
35
73
4
46
18
Mid-summer rainfall
Kalahari
II4b
17
II4c
59
29
92
46
137
1
20
0
10
27
12
19
130
0
1
50
183
Highveld
II3a
II3c
144
97
24
4
140
134
3
6
1
1
125
101
19
2
3
0
211
150
Transitional
II3d
106
35
156
6
6
85
56
2
197
East coast
II(I)a
II2a
II3e
II3f
35
5
28
4
46
6
43
4
19
0
23
0
13
0
2
0
31
4
15
0
20
2
37
3
29
1
6
0
144
7
108
4
13
25
60
71
83
189
12
1
80
8
1
10
32
32
58
29
49
110
15
0
23
108
113
349
29
0
32
0
Subtropical/tropical
II2b
61
II3b
53
II3g
117
*Discrepancies between the number of collection localities and collection data totals are due to missing
information.
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
14
A. L. V. Davis
The present study examines both climatic and biogeographical relationships
of the southern African dung beetle fauna recorded mainly on large herbivore
dung in unshaded situations. Regional trends in biogeographical and tribal
composition of the fauna have been determined. Multivariate analytical techniques have been used to identify major climatic and geographical groups. The
possible influence of local factors (soil, vegetation, dung type) on these groups
has been assessed. Local geographical distribution patterns and implications for
routes of dispersal are discussed.
Methods
Dung beetle distribution data for Africa south of latitude 15)S were drawn from
the reference collection of the former Australian CSIRO Dung Beetle Research
Unit (DBRU) which was amassed between 1971 and 1986. Distribution data
were plotted for the 220 species recorded in >15 separate collections out of 2068
made in southern Africa. Both geographical and climatic data matrices were
created at optimal scales for minimizing noise and maximizing biogeographical
information content. On all geographical matrices, species data were plotted
according to presence (1) or absence (0) from the 76#4) squares of latitude and
longitude from which dung beetle collections were made. For the climatic matrix,
calculations were made of the percentage occurrences of species across eight
major climatic regions of southern Africa (Table 2). These quantitative climatic
data were derived from a presence or absence data matrix comprising 264 entries
for each species. These entries comprised records for 202#1) squares of latitude
and longitude from which dung beetles were collected with the balance comprising separate records for each portion of a degree square partitioned by climatic
boundaries. Presence records for each species were summed for each of the eight
climatic regions. These summed records were divided by the total number of
degree squares in which collections were made in that region. These numbers
were converted to percentage distributions across the eight climatic regions. The
treatment of the data gave equal weighting both to each species, whose ranges
differed in size, and to each of the climatic regions which differed in area.
The climatic distribution of the 220 species was classified using cluster
analysis. A similarity matrix was calculated using the Bray-Curtis (non-metric)
similarity coefficient and this matrix was subjected to analysis by the aglomerative clustering technique, group average linking. The results were summarized as
a dendrogram (not illustrated) from which seven species clusters of similar
climatic distribution were defined at the 50% level of similarity. One of these
clusters (F) was subdivided at the 65% level of similarity.
The results of the climatic analysis (Table 2) were used to define four
supergroups of species whose geographical distribution was also analysed using
cluster analysis. These groups comprised western climatic specialists (B, C, H, 47
spp.), eastern climatic specialists (A, D, E, G, 71 spp.), subtropical/tropical
generalists (F1, F3, 74 spp.), and temperate to tropical generalists (F2, F4, F5, 28
spp.). Geographical distribution matrices for each group were each analysed
using the same clustering methods as in the climatic analysis. Results were
depicted as dendrograms from which species clusters of similar geographical
distribution were defined at either the 35% (Figs 2, 5) or 40% levels of similarity
(Figs 7, 8).
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 15
Table 2. List of 220 species of dung beetles grouped according to the results of cluster analysis of their
percentage distributions across eight climatic regions of southern Africa
Group
Species
A
Garreta malleolus (Kolbe)
Sisyphus impressipennis Lansberge
Copris armiger Gillet
Onitis cryptodus Gillet
Onitis inversidens Lansberge
Diastellopalpus infernalis Lansberge
Diastellopalpus quinquedens Bates
Onthophagus anomalus (Klug)
Onthophagus jeaneli d’Orbigny
Mean
B
Scarabaeus rugosus (Hausman)
Copris anceus Olivier
Copris capensis Waterhouse
Epirinus aeneus Wiedeman
Epirinus flagellatus (Fabricius)
Onthophagus minutus Hausman
Onitis confusus Bohaeman
Onthophagus cameloides d’Orbigny
Scarabaeus suri (Hausman)
Chironitis scabrosus (Fabricius)
Metacatharsius latifrons Harold
Mean
C
Onthophagus giraffa Hausman
Onitis aygulus (Fabricius)
Mean
D
Neosisyphus mirabilis Arrow
Sisyphus sp. nr sordidus
Catharsius sp. nr pandion
Copris inhalatus santaluciae
Nguyen-Phung
Onthophagus sp.
Onthophagus sp. nr sugillatus
Proagoderus aureiceps d’Orbigny
Onthophagus juvencus Klug
Proagoderus aciculatus Fahraeus
Copris puncticollis Boheman
Metacatharsius pseudoopacus Ferreira
Litocopris muticus Boheman
Onthophagus beiranus Péringuey
Sisyphus sordidus Boheman
Mean
Percentage frequency in climatic region
WR
BR
Late SR
Kala.
Highv.
II3d
EC
Trop.
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
100·0
100·0
100·0
100·0
100·0
100·0
100·0
100·0
100·0
100·0
100·0
100·0
91·6
61·0
62·7
73·1
69·7
77·9
84·5
42·8
36·5
72·7
0·0
0·0
8·4
20·4
28·8
26·9
16·0
15·9
15·5
27·6
0·0
14·5
0·0
0·0
0·0
18·6
0·0
0·0
8·4
6·2
0·0
26·7
28·0
8·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
30·5
2·7
0·0
0·0
0·0
0·0
8·5
0·0
5·9
0·0
0·0
2·9
4·9
2·8
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
24·5
9·7
17·1
60·2
44·8
52·5
0·0
23·3
11·7
4·3
6·8
5·6
11·1
15·4
13·3
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
100·0
100·0
100·0
100·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
9·8
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
100·0
100·0
94·1
95·8
96·3
90·2
90·3
76·3
84·3
84·9
93·7
0·0
0·0
5·9
4·3
3·8
3·5
0·0
23·8
15·8
15·1
5·2
Table 2 continued on next page
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
16
A. L. V. Davis
Table 2. continued from previous page
Percentage frequency in climatic region
Group
Species
WR
BR
Late SR
Kala.
Highv.
II3d
EC
Trop.
E
Scarabaeus natalensis zur Strassen
Copris jacchoides Nguyen-Phung
Onthophagus obtutus Péringuey
Copris corniger Sahlberg
Xinidium dentilabris Harold
Neosisyphus barbarossa Wiedeman
Onitis perpunctatus Balthasar
Onthophagus vigens d’Orbigny
Copris caelatus Fabricius
Sisyphus caffer Boheman
Proagoderus lanista Castelnau
Epirinus obtusus Boheman
Onthophagus fritschi d’Orbigny
Onthophagus cyaneoniger d’Orbigny
Onthophagus lugubris Fahraeus
Copris antares Ferreira
Neosisyphus macroruber Paschalidis
Euoniticellus triangulatus (Harold)
Euoniticellus africanus (Harold)
Sisyphus costatus Thunberg
Onthophagus asperulus d’Orbigny
Onthophagus bubalus Harold
Litocopris simplex Harold
Onitis pecuarius Lansberge
Drepanocerus sulcicollis (Castelnau)
Oniticellus pictus (Hausman)
Onthophagus binodis Thunberg
Gymnopleurus leei (Fabricius)
Mean
0·0
0·0
0·0
0·0
0·0
10·4
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
21·4
13·9
0·0
0·0
0·0
0·0
0·0
0·0
5·6
8·8
0·0
2·2
0·0
0·0
8·3
9·9
0·0
19·0
7·0
0·0
0·0
0·0
0·0
25·8
25·6
17·5
28·0
27·0
0·0
28·6
21·2
32·2
26·4
22·1
51·2
47·0
36·2
46·0
42·9
37·7
20·0
6·6
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
3·9
26·9
17·8
18·2
0·0
3·5
18·8
0·0
12·2
0·0
0·0
13·4
0·0
0·0
0·0
0·0
2·8
6·5
4·6
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
20·6
0·0
4·8
0·0
0·0
2·1
0·0
0·0
0·0
0·0
0·0
0·0
1·0
64·8
74·2
72·9
65·6
54·0
49·0
36·0
39·3
49·3
91·9
83·1
48·1
56·6
64·3
72·0
69·5
53·2
22·3
37·5
35·6
46·7
37·9
37·6
34·5
40·0
30·1
39·5
41·5
52·9
28·6
16·4
18·8
22·5
33·4
21·6
47·7
60·7
50·8
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
12·2
4·8
18·3
18·0
16·7
0·0
0·0
0·0
11·6
6·1
14·3
14·4
0·0
9·5
0·0
0·0
9·7
0·0
9·2
0·0
0·0
0·0
9·9
0·0
0·0
0·0
0·0
0·0
0·0
14·1
5·6
10·6
3·5
4·8
11·2
18·5
23·8
6·7
0·0
0·0
4·9
0·0
0·0
0·0
2·0
3·0
0·0
0·0
0·0
0·0
8·1
3·1
0·0
0·0
0·0
0·0
0·0
7·4
1·5
0·0
3·3
5·4
3·0
0·0
0·0
0·0
0·0
0·0
0·0
1·3
F1
Pachylomerus femoralis (Kirby)
Kheper lamarcki (M’Leay)
Scarabaeus goryi Harold
Onthophagus vinctus Erichson
Onitis deceptor Péringuey
Heliocopris andersoni Bates
Heliocopris japetus Klug
Copris elphenor Klug
Metacatharsius troglodytes Boheman
Onitis uncinatus Klug
Copris evanidus Klug
Allogymnopleurus thalassinus Klug
Heliocopris faunus Boheman
Scarabaeus galenus (Westwood)
Onthophagus pallidipennis Fahraeus
Scarabaeus zambesianus Péringuey
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
7·8
8·7
2·6
0·0
8·9
5·8
6·7
6·7
3·0
7·2
0·0
7·1
20·6
15·5
2·6
0·0
28·5
23·8
25·0
14·9
9·7
12·6
19·5
25·5
29·8
17·7
17·6
21·2
18·7
16·9
30·6
37·3
0·0
0·0
0·0
2·1
0·0
0·0
0·0
4·7
2·1
3·8
3·8
10·1
0·0
0·0
0·0
0·0
22·8
19·0
22·2
19·8
19·5
25·2
29·2
29·1
26·4
31·4
35·2
26·7
29·9
27·1
33·4
24·9
26·4
33·0
32·2
38·2
33·8
29·2
11·3
8·4
7·7
15·9
17·0
18·1
17·3
23·5
19·3
28·8
14·4
15·4
18·0
25·0
28·1
27·3
33·4
25·6
31·0
24·1
26·4
16·9
13·5
17·1
14·1
9·0
Table 2 continued on next page
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 17
Table 2. continued from previous page
Group
Species
F1 continued
Onthophagus signatus Fahraeus
Pedaria sp.
Neosisyphus confrater Kolbe
Onitis picticollis Boheman
Proagoderus brucei Reiche
Drepanocerus fastiditus Péringuey
Onthophagus obtusicornis Fahraeus
Neosisyphus spinipes Thunberg
Proagoderus loricatus Klug
Onthophagus stellio Erichson
Proagoderus tersidorsis d’Orbigny
Hyalonthophagus alcyonides
(D’Orbigny)
Gymnopleurus virens Erichson
Copris amyntor Klug
Drepanocerus laticollis Fahraeus
Phalops smaragdinus Harold
Euonthophagus carbonarius Klug
Copris denticulatus Nguyen-Phung
Onthophagus fimetarius Roth
Caccobius nigritulus Klug
Phalops flavocinctus Klug
Oniticellus formosus Chevrolat
Onthophagus pugionatus Fahraeus
Onthophagus flavolimbatus d’Orbigny
Onthophagus rasipennis d’Orbigny
Catharsius philus Kolbe
Sisyphus seminulum Gerstaecker
Onthophagus ebenus Péringuey
Tiniocellus spinipes (Roth)
Heliocopris neptunus Boheman
Neosisyphus fortuitus Péringuey
Pedaria sp.
Pedaria sp.
Phalops ardea Klug
Onthophagus depressus Harold
Onthophagus bicavifrons d’Orbigny
Onthophagus lamelliger Gerstaecker
Phalops boschas Klug
Neosisyphus calcaratus Klug
Kheper nigroaeneus (Boheman)
Copris mesacanthus (Harold)
Sisyphus gory Harold
Onthophagus aeruginosus Roth
Euonthophagus sp.
Catharsius sesostris Waterhouse
Onitis fulgidus Klug
Percentage frequency in climatic region
WR
BR
Late SR
Kala.
Highv.
II3d
EC
Trop.
0·0
0·0
0·0
0·0
0·0
7·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
4·2
0·0
0·0
0·0
0·0
0·0
11·3
0·0
0·0
0·0
0·0
13·5
2·2
0·0
0·0
0·0
0·0
3·4
0·0
0·0
0·0
0·0
0·0
34·4
12·0
0·0
0·0
0·0
0·0
9·1
1·3
8·5
4·4
0·0
2·9
0·0
15·5
21·4
24·1
25·2
14·2
21·3
11·6
0·0
0·0
8·5
11·3
9·8
38·4
26·5
24·8
26·0
29·2
29·2
30·8
33·9
35·2
39·2
34·9
22·8
27·8
30·7
28·7
30·1
33·8
21·2
38·6
39·3
50·9
45·3
40·4
19·5
0·0
21·5
22·4
18·7
15·8
15·8
6·5
18·4
9·5
7·1
10·5
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
4·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
6·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
1·2
0·0
0·0
1·7
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
1·2
0·0
4·0
0·0
0·0
8·2
5·1
12·0
6·8
11·4
4·7
8·0
4·8
6·1
7·5
2·7
7·2
7·7
11·5
3·5
3·4
0·0
0·0
0·0
0·0
0·0
11·8
8·2
12·0
15·4
14·6
11·2
9·9
4·5
8·0
9·9
4·3
2·9
0·0
5·3
3·3
0·0
2·9
3·0
2·0
10·4
0·0
7·9
9·7
3·5
0·0
0·0
2·5
4·5
4·4
4·2
3·5
4·4
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
9·6
11·6
10·4
14·6
5·6
11·2
5·4
40·8
41·0
40·1
36·0
36·5
37·4
37·5
38·4
40·5
37·5
53·6
47·9
46·2
45·9
42·0
40·9
39·4
42·6
40·9
53·6
42·3
56·8
54·9
48·1
44·0
45·5
44·9
19·8
27·0
26·8
28·2
34·7
34·6
33·1
23·6
23·7
23·2
26·1
21·1
28·9
18·6
22·2
28·1
26·0
24·8
27·7
26·7
22·2
32·4
31·6
30·4
30·9
31·5
20·7
26·2
11·0
19·1
13·9
15·3
18·8
19·5
28·6
26·0
21·7
26·1
20·1
20·1
28·7
22·1
28·8
24·6
24·3
28·0
27·0
24·2
34·6
17·5
17·6
15·5
17·3
19·4
18·0
17·7
19·7
26·0
23·1
17·2
25·8
28·6
20·5
17·8
26·0
25·4
21·1
24·3
32·1
30·8
31·9
21·3
31·3
31·2
32·8
Table 2 continued on next page
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
18
A. L. V. Davis
Table 2. continued from previous page
Group
Species
Percentage frequency in climatic region
WR
BR
Late SR
Kala.
Highv.
II3d
EC
Trop.
F1 continued
Heteronitis castelnaui (Harold)
Onitis mendax Gillet
Garreta nitens (Olivier)
Anachalcos convexus Boheman
Onitis viridulus Boheman
Onthophagus ?sugillatus Klug
Kheper subaeneus (Harold)
Hyalonthophagus alcedo (d’Orbigny)
Caccobius ferrugineus Fahraeus
Gymnopleurus humanus M’Leay
Mean
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·1
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·4
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
6·5
0·0
1·9
3·8
0·0
8·6
9·2
1·3
7·6
3·8
6·2
7·1
14·4
9·8
0·0
0·0
2·8
0·0
6·6
0·0
0·0
0·0
0·0
0·0
4·6
30·7
27·8
25·9
27·5
30·7
30·2
30·5
49·3
42·5
28·7
34·5
26·7
32·2
30·0
31·9
35·6
35·0
35·4
0·0
8·2
0·0
25·3
38·8
40·1
32·7
31·4
25·8
27·2
30·3
44·5
35·8
56·9
23·5
F2
Catharsisus tricornutus de Geer
Digitonthophagus gazella (Fabricius)
Euoniticellus intermedius (Reiche)
Liatongus militaris (Castelnau)
Drepanocerus kirbyi (Kirby)
Oniticellus planatus Castelnau
Onthophagus sp. nr sugillatus
Onitis caffer Boheman
Chironitis sp. nr scabrosus
Onitis alexis Klug
Chironitis hoplosternus (Harold)
Drepanocerus patrizii (Boucomont)
Phalops dregei Harold
Sarophorus costatus (Fahraeus)
Heliocopris hamadryas (Fabricius)
Neosisyphus ruber Paschalidis
Onthophagus cribripennis d’Orbigny
Onitis tortuosus Houston
Copris obesus Boheman
Caccobius obtusus Fahraeus
Cyptochirus ambiguus Kirby
Copris macer Péringuey
Onthophagus parumnotatus Fahraeus
Garreta unicolor (Fahraeus)
Mean
9·8
4·2
8·0
0·0
0·0
0·0
0·0
13·8
0·0
0·0
10·1
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
1·9
12·7
20·6
12·9
17·0
16·6
23·5
22·7
21·6
15·2
9·3
4·7
7·1
7·3
13·3
17·1
7·8
0·0
0·0
0·0
11·0
19·0
0·0
0·0
0·0
10·8
0·0
6·0
7·2
1·4
0·0
0·0
5·5
5·3
7·9
9·1
14·6
3·7
11·4
1·7
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
3·1
3·0
8·0
10·5
3·0
1·6
2·1
5·2
3·6
12·3
14·6
10·6
8·0
14·6
1·9
3·9
5·4
2·9
3·5
7·2
0·0
1·8
0·0
0·0
0·0
5·2
13·3
8·5
14·9
19·3
18·3
20·0
23·3
20·6
24·0
16·7
20·6
15·6
16·1
4·9
10·1
24·2
33·2
32·9
31·2
32·4
32·6
22·4
27·3
17·5
20·8
20·5
20·4
20·9
26·8
31·5
20·6
24·0
23·1
19·7
16·4
31·8
48·2
41·6
45·4
38·9
32·1
34·2
32·3
38·6
37·6
28·7
52·0
48·3
54·1
32·8
21·4
18·6
13·3
20·7
21·9
19·1
11·9
8·4
5·7
16·3
0·0
0·0
0·0
21·9
9·0
14·5
13·2
21·4
5·6
14·5
16·6
10·0
7·0
23·5
13·1
19·3
13·7
12·4
11·8
10·2
14·8
7·4
3·7
15·1
17·7
7·7
17·4
9·0
10·9
21·0
16·1
16·5
10·0
17·4
4·5
1·3
15·6
17·4
4·9
12·3
F3
Onthophagus albipodex d’Orbigny
Onthophagus quadrinodosus Fahraeus
Onthophagus ?pullus Roth
Mean
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
5·8
0·0
0·0
1·9
0·0
0·0
0·0
0·0
69·3
67·0
76·2
70·8
0·0
12·9
14·7
9·2
25·0
20·1
9·2
18·1
F4
Copris fidius Olivier
0·0
30·1
0·0
0·0
5·5
51·4
9·9
3·1
Table 2 continued on next page
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 19
Table 2. continued from previous page
Group
Species
Percentage frequency in climatic region
WR
BR
Late SR
Kala.
Highv.
II3d
EC
Trop.
F5
Caccobius viridicollis Fahraeus
Scarabaeus bohemani Harold
Mean
0·0
0·0
0·0
0·0
0·0
0·0
8·3
22·4
15·4
27·1
20·4
23·8
11·7
5·3
8·5
36·1
49·0
42·6
10·4
0·0
5·2
6·5
2·9
4·7
G
Scarabaeus deludens zur Strassen
Proagoderus bicallosus Klug
Onthophagus lacustris Harold
Onitis robustus Boheman
Caccobius sp.
Copris bootes Klug
Proagoderus dives Harold
Milichus apicalis Fahraeus
Neosisyphus infuscatus Klug
Metacatharsius exiguus Boheman
Onitis obscurus Lansberge
Onitis reichei Lansberge
Onitis westermanni Lansberge
Onthophagus plebejus Klug
Proagoderus rectefurcatus Fairmaire
Copris sp. nr macer
Catharsius heros Boheman
Kheper cupreus (Castelnau)
Onthophagus interstitialis Fahraeus
Scarabaeus funebris (Boheman)
Mean
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
10·2
0·0
0·0
0·0
0·0
0·0
0·0
8·9
0·0
2·4
18·5
2·0
7·7
13·6
0·0
0·0
12·8
0·0
4·1
0·0
3·7
22·2
31·6
11·0
13·2
0·0
0·0
10·0
19·5
29·0
0·0
10·1
9·4
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
9·5
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
36·5
26·1
3·6
0·0
0·0
0·0
0·0
0·0
0·0
16·3
23·7
14·7
0·0
0·0
0·0
0·0
0·0
0·0
20·0
0·0
0·0
10·3
0·0
4·3
53·5
47·2
54·6
51·7
59·4
65·8
47·2
54·9
51·0
51·5
0·0
25·5
15·3
39·1
34·9
23·2
22·5
22·3
41·6
23·4
39·2
38·9
39·2
45·4
48·3
27·8
34·2
32·4
21·4
21·2
16·1
68·4
63·5
71·5
60·9
65·1
46·9
49·1
48·7
9·3
21·9
41·5
H
Scarabaeus viator Péringuey
Euonthophagus vicarius Péringuey
Chironitis audens (Péringuey)
Phalops euplynes Bates
Gymnopleurus sericatus Erichson
Metacatharsius marani Balthasar
Onthophagus probus Péringuey
Gymnopleurus asperrimus Felsche
Gymnopleurus andreaei Ferreira
Gymnopleurus aenescens Wiedeman
Onthophagus verticalus Fahraeus
Onitis obenbergeri Balthasar
Metacatharsius opacus Waterhouse
Copris laioides Boucomont
Heliocopris atropos Boheman
Pachylomerus opaca Lansberge
Scarabaeus satyrus (Boheman)
Scarabaeus ambiguus (Boheman)
Phalops wittei Harold
17·5
0·0
0·0
4·6
6·3
10·4
17·1
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
14·1
0·0
0·0
32·2
35·4
22·2
12·7
11·5
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
6·5
0·0
0·0
50·3
64·6
57·8
55·1
60·1
74·8
73·9
100·0
100·0
4·3
6·3
0·0
0·0
0·0
0·0
51·2
37·2
25·6
31·6
0·0
0·0
6·3
14·4
13·1
10·9
9·0
0·0
0·0
60·4
41·1
47·9
42·3
58·1
57·4
20·9
22·1
20·9
34·4
0·0
0·0
0·0
6·2
4·2
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
27·1
15·7
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
18·6
27·4
38·3
0·0
0·0
0·0
27·9
14·8
13·9
8·1
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
7·9
0·0
13·1
0·0
19·0
0·0
0·0
0·0
0·0
0·0
0·0
13·7
6·9
4·7
3·9
0·0
0·0
0·0
16·8
17·3
13·8
44·7
41·9
23·7
0·0
5·3
12·6
10·2
Table 2 continued on next page
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
20
A. L. V. Davis
Table 2. continued from previous page
Percentage frequency in climatic region
Group
Species
WR
BR
Late SR
Kala.
Highv.
II3d
EC
Trop.
H continued
Scarabaeus flavicornis (Boheman)
Copris cassius Péringuey
Kheper prodigiosus (Erichson)
Copris gracilis Waterhouse
Proagoderus sappharinus Péringuey
Copris subsidens Péinguey
Catharsius ulysses Boheman
Onthophagus quadraticeps Harold
Onthophagus sp. nr variegatus
Catharsius calaharicus Kolbe
Scarabaeus damarensis Janssens
Onthophagus sp.
Copris cornifrons Boheman
Metacatharsius sp.
Scarabaeus proboscideus Guérin
Mean
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
12·6
2·4
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
3·5
30·0
24·5
26·2
32·3
33·0
41·1
20·6
24·5
26·9
37·0
45·7
46·1
52·2
61·6
36·3
39·1
39·2
49·7
42·9
42·3
35·9
37·4
45·0
49·1
64·6
55·1
49·8
41·9
42·7
38·4
46·3
33·5
0·0
5·0
0·0
0·0
0·0
0·0
11·7
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
2·1
26·1
15·3
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
0·0
5·6
0·0
0·0
0·0
0·0
0·0
0·0
0·0
10·3
0·0
0·0
0·0
0·0
0·0
0·0
0·0
1·5
4·7
5·5
30·9
25·4
31·1
21·6
22·7
16·1
8·5
7·9
4·5
12·1
5·1
0·0
4·8
12·2
WR, winter rainfall; BR, bimodal rainfall; Late SR, late summer rainfall; Kala., Kalahari; Highv.,
highveld; EC, east coast; Trop, north-east subtropical/tropical—see Fig. 1.
Clusters of similar geographical distribution from this serial climatic/
geographical analysis were assessed for validity by comparison with the results
from a second analysis conducted solely on geographical distribution. Classification of the geographical distribution of the 220 species was used to define four
subgroups of species whose geographical distribution matrices were classified
separately using the same clustering methods as in the climatic/geographical
analysis. The results are not illustrated. However, on the dendrograms constructed from the first analysis (Figs 2, 5, 7, 8), core species are marked with an
asterisk if they formed similar clusters to those shown in the second analysis.
Species which change in association between the two analyses are marked with
a cross. This assessment allows spurious groupings and species with outlier
distributions to be identified.
As collecting intensity on different soil, vegetation and dung types varied
regionally (Table 1), an assessment was made of its influence on distributional
results for the 220 species (Figs 2, 5, 7, 8). In each of the 25 climatic regions of
southern Africa (Walter & Lieth, 1964), the numbers of collections for each
species were summed for fine-grained vs. coarse-grained (sand) soils, for pasture/
grassland vs. shrubland/open woodland, and for ruminant vs. non-ruminant
herbivore dung types. The overall number of collections for each species on each
habitat variable were divided by the total number of collections made in the
climatic regions from which that species was collected. For each pair of variables,
these numbers were converted to percentages. Where possible, these standardized
results for collecting bias were compared with quantitative results for habitat
associations.
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 21
Table 3. Biogeographical composition of the total 582 species of dung beetles recorded and the 220 species
recorded most frequently in eight climatic regions of southern Africa
Percentage biogeographical composition of species
within each climatic region
Overall %
Mid-summer rainfall
Late
composition
Biogeographical
East Subtropical/ in southern
Winter Bimodal summer
distribution
tropical
rainfall rainfall rainfall Kalahari Highveld II3d coast
Africa
pattern*
220 species
1
2
3
4
5
6
No.species
80·6
3·2
12·9
3·2
0·0
0·0
31
71·2
6·8
16·9
3·4
0·0
1·7
59
72·9
8·2
10·6
5·9
2·4
0·0
85
48·1
21·8
16·5
11·3
1·5
0·8
133
59·1
13·6
19·1
7·3
0·0
0·9
110
45·9
21·8
19·5
9·8
1·5
1·5
133
40·7
24·1
22·1
10·3
1·4
1·4
145
44·3
24·7
18·4
10·3
1·1
1·1
174
55·0
20·0
15·0
8·2
0·9
0·9
220
582 species
1
2
3
4
5
6
No.species
90·5
1·6
6·3
1·6
0·0
0·0
63
79·1
4·7
11·6
2·3
1·2
1·2
86
77·6
6·0
7·8
4·3
4·3
0·0
116
60·3
16·3
12·0
8·2
2·2
1·1
184
69·9
9·8
13·7
5·9
0·0
0·7
151
58·4
17·3
13·4
7·4
1·0
2·5
202
55·5
19·7
15·1
6·7
0·8
2·1
238
57·6
17·7
11·1
7·6
2·2
3·8
368
71·8
11·7
7·4
5·0
1·5
2·6
582
*1, Recorded only in southern Africa south of latitude 15)S. 2, Recorded in southern, southern central and
east Africa. 3, Pan-African distribution, east, west/northern central, southern central, southern and,
infrequently, west central Africa. 4, Recorded in southern and southern central Africa. 5, Recorded in
southern, southern central and west central Africa. 6, Recorded in southern, southern central, west central
and east Africa.
Biogeographical relationships were assessed for all 582 species of dung beetles
which were recorded in southern Africa by the DBRU. Pan-African distribution
data were drawn from the reference collection of the former DBRU, Ferreira
(1972, 1978), Cambefort (1982, 1991a), Scholtz & Howden (1987a, b), Howden
& Scholtz (1987), Nguyen-Phung (1988a, b, c) and Nguyen-Phung & Cambefort
(1986, 1987). Biogeographical regions are as in Davis & Dewhurst (1993) except
that southern Africa is defined as lying south of 15)S instead of 20)S. The six
different biogeographical distribution patterns shown by the species are defined
in Table 3.
Cross-climatic variation in biogeographical composition of the dung beetle
fauna was assessed both for the total 582 species recorded and for the 220 species
showing the greatest frequency in collections. Within each of the eight climatic
regions, the numbers of species showing each biogeographical distribution
pattern were converted to percentage representation to remove species/area bias.
Cross-climatic variation in dung beetle community structure was assessed
using data for the entire 582 species recorded. The numbers of species recorded
in each of the 25 climatic regions (Walter & Lieth, 1964) were divided amongst
the nine tribes of dung beetles. These numbers were converted to percentage
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
22
A. L. V. Davis
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 23
tribal composition within each climatic region to remove species/area bias. Mean
percentage occurrence for each tribe was calculated for each of the four major
climatic regions of southern Africa (Davis, 1987). One-way analysis of variance
was used to test for statistical significance of cross-climatic differences in tribal
composition.
Results and discussion
The dung beetle fauna of southern Africa shows clear climatic and biogeographical associations. Increasing endemism to the south-west in both dung beetles
(Table 3) and Coleoptera as a whole (Endrödi-Younga, 1978) is associated
especially with arid late summer, winter and bimodal rainfall climate types.
Geographical groups of southern African endemics are centred to the south-west
(Figs 2–4) and also on the south-eastern highlands and the subtropical east coast
(Figs 4–6). However, most species groups centred to the north-east in the
mid-summer rainfall region comprise primarily widespread elements with broad
tropical biogeographical associations (Figs 6–8). Some widespread species
penetrate into the south-western climate types, chiefly across the highveld and
along the moist southern coastline. The numbers of widespread and highveld
species decline sharply to the west of George (34)00*S 22)15*E) (Figs 4, 6).
George is situated at the westerly limit of climate Type V (Walter & Lieth, 1964)
in which substantial bimodal rain falls throughout the year. Beyond George,
there is increasing emphasis on the spring peak in rainfall of the bimodal climate
Type IV(V). Most of the widespread, summer rainfall species, which penetrate
beyond George to the Western Cape, are active in the dry summer (Davis, 1987,
1993).
Increasing endemism to the arid west and cooler south-west is paralleled by
reduced species richness (Table 3) and changing community structure (Table 4;
Davis, 1994c). Functional complexity of the dung beetle fauna is reduced to the
south-west (Davis, 1994c) by the virtual loss of endocoprids (some Oniticellini),
which breed within dung in situ, and by the decline or loss of kleptocoprid species
(some Onthophagini) which use dung buried by other beetles. Tunnelling
Coprinae (Dichotomiini to Oniticellini) manifest few cross-climatic changes in
community structure except for reduced percentage occurrence of Onthophagini
to the south-west. This tribe includes many kleptocoprids in communities to the
Fig. 2. Dendrogram showing percentage geographical overlap between species of the western climatic
specialist group defined from Table 2. Species marked with an asterisk or a cross show, respectively,
similar or different cluster associations using different methods for analysing spatial distribution.
*Superscripted numbers represent the biogeographical distribution pattern shown by each species.
Numbers in box brackets are doubtful distribution patterns. **Standardized percentage of collection bias
to sand as opposed to finer-grained soils, numbers in brackets are percentage occurences of species
on sand in Gauteng from quantitative data (Davis, 1996), S, sand, C, clay, G, soil generalist,
asterisk=statistically significant relationship (P<0·05); **standardized percentage of collection bias to
pasture/grassland as opposed to shrubland/open woodland, numbers in brackets are percentage
occurences of species in pasture/grassland in the Western Cape and Gauteng from quantitative data
(Davis, 1993, 1996), P, pasture; S, shrubland; OW, open woodland; G, vegetation generalist, forest= >75%
of collections of species made in forest; **standardized percentage of collection bias to ruminant as
opposed to non-ruminant dung, numbers in brackets are percentage occurences of species on ruminant
dung in Gauteng from quantitative data (Davis, 1994b).
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
24
A. L. V. Davis
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 25
north-east. Increased representation by ball-rolling Scarabaeinae to the west
and south-west (Table 4) reflects the southern African centres of distribution in
both the Canthonini (Scholtz & Howden, 1987a, b; Howden & Scholtz, 1987)
and the Scarabaeini (Mostert & Scholtz, 1986). Sisyphini, and to a lesser extent,
Gymnopleurini, show an opposite trend.
There is also increased prominence of Gondwanaland Coleoptera to the
south-west and these are thought to be the oldest faunal members (EndrödiYounga, 1978). In Gondwanaland dung beetles (Halffter & Matthews, 1966;
Halffter, 1974), increased prominence to the south-west is true of the Canthonini
but not of the Dichotomiini, which are centred in the summer rainfall tropics
(Ferreira, 1972; Davis, 1993) south of the equator (Cambefort, 1991b) with a
taxonomic disjunction between tropical and winter/bimodal rainfall elements
(Davis, 1993). Southern Gondwanaland taxa may have a south temperate
(palaeantarctic) origin (Halffter, 1974; Endrödi-Younga, 1978) with tropical
relicts resulting from the 15–18) northward drift of the African continent from
the late Cretaceous to the Miocene (Axelrod & Raven, 1978). Alternatively,
tropical Gondwanaland taxa may have entered Africa via a tropical west African
route (Halffter, 1974). There is circumstantial support for Cambefort’s competitive exclusion hypothesis (1991b), which explains the distribution of African
Canthonini, since Anachalcos is the only widespread tropical canthonine genus
whereas Cape canthonines are maximized to cool seasonal conditions (Davis,
1993) which reduces overlap with other Scarabaeinae. Survival and radiation of
tropical Dichotomiini may result from dietary, behavioural (Cambefort, 1991b),
size (Davis, 1990) and distributional specializations.
Endrödi-Younga (1978) has defined seven biogeographical regions for southern Africa based on the distribution of ground Coleoptera, mainly Tenebrionidae.
The present study on dung beetles partly supports this zonation although some
modifications are proposed. These are based on species distribution of major
dendrogram clusters (Figs 2–8, Table 5) with minor clusters ignored. Members of
the western (Fig. 2) and eastern climatic specialists (Fig. 5) mostly form similar
clusters in both the climatic/geographical and the purely geographical analyses.
However, there are greater differences between clusters defined from the
subtropical/tropical (Fig. 7) and temperate to tropical generalists (Fig. 8) by the
two analyses. Even so, most species added to clusters in the climatic/geographical
analysis (7C, 7D, 8A) generally comprise blocks of species from no more than
one or two different clusters in the purely geographical analysis.
The winter and bimodal rainfall cluster (2B) is primarily restricted to sand on
the western and southern coastline of South Africa (Figs 2, 3, Table 5). Its
distribution largely conforms to the Cape Zone (Endrödi-Younga, 1978) except
that it includes Namaqualand but excludes the east coastline beyond East
London (33)01*S, 27)58*E). A few species occupy the entire winter and bimodal
rainfall regions, Copris capensis, Epirinus flagellatus. However, most species are
restricted to subregional distribution patterns. One subgroup is associated with
Fig. 3. Geographical distribution of species clusters of dung beetles defined from Fig. 2 (the numbers of
species occurring in each 4) square of latitude and longitude are size-coded: largest size= >75% of species
comprising each dendrogram cluster, second largest=50–75% of species, second smallest=25–50% of
species, smallest size= <25% of species comprising each dendrogram cluster).
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
26
A. L. V. Davis
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Fig. 4. Geographical distribution of species clusters of dung beetles defined from Figs 2 and 5 (see notes in legend to Fig. 3 for key
to species numbers).
Associations of southern African dung beetles 27
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Fig. 5. Dendrogram showing percentage geographical overlap between species of the eastern climatic specialist group defined from
Table 2 (*,** see notes in legend to Fig. 2).
28
A. L. V. Davis
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Fig. 6. Geographical distribution of species clusters of dung beetles defined from Figs 5, 7 and 8 (see notes in legend to Fig. 3 for
key to species numbers).
Associations of southern African dung beetles 29
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Fig. 7. Dendrogram showing percentage geographical overlap between species of the subtropical/tropical climatic specialist
group defined from Table 2 (*,** see notes in legend to Fig. 2).
30
A. L. V. Davis
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Fig. 8. Dendrogram showing percentage geographical overlap between species of the temperate to tropical climatic generalist
group defined from Table 2 (*,** see notes in legend to Fig. 2).
Associations of southern African dung beetles 31
Table 4. Percentage subfamily and tribal composition of the dung beetle fauna in four major climatic
regions of southern Africa
Mean percentage species composition&S.D.
in climate types comprising each climatic region
Subfamily/
tribe
Subfamily
Scarabaeinae
Coprinae
Tribes
Scarabaeini
Gymnopleurini
Sisyphini
Canthonini
Dichotomiini
Coprini
Onitini
Onthophagini
Oniticellini
Total number of
species in region
Winter
rainfall
Bimodal
rainfall
Late summer
rainfall
Mid-summer
rainfall
Resul;ts of
ANOVA (F)
(d.f.=3,21)
45·6&17·3b
54·4&17·3a
22·5&2·2a
77·5&2·2b
26·7&7·9a
73·3&7·9b
20·7&3·4a
79·3&3·4b
8·09***
8·09***
28·0&15·5b
0·9&1·7a
1·3&1·6a
15·4&5·6c
4·0&2·2
13·3&2·6
12·4&6·9
15·7&8·0a
9·0&5·2ab
4·7&2·1a
2·8&1·3ab
6·1&4·7ab
8·9&3·6bc
5·6&2·4
9·9&4·4
18·8&4·8
28·5&3·4b
14·6&3·8b
15·8&7·5ab
7·2&1·7b
1·1&1·4a
2·6&1·3ab
4·5&2·6
12·6&5·6
16·3&4·0
35·5&2·2bc
4·4&3·1a
7·1&5·0a
4·3&3·1ab
7·4&3·4b
1·9&1·5a
5·5&2·5
14·9&2·2
13·1&4·0
38·8&4·7c
6·9&3·1a
6·99**
4·89**
6·72**
21·99***
0·48
1·48
1·43
21·37***
4·38*
63
86
116
517
*P<0·05, **P<0·01, ***P<0·001. In each line, values followed by a different letter differed significantly
(P<0·05, Tukey’s HSD).
the dry west coastal sands of South Africa. Several species occur along the entire
coastline, Scarabaeus rugosus, Kheper bonellii M’Leay. Others appear in the
moister regions to the south, Copris anceus, Epirinus scrobiculatus Harold,
E. granulatus Scholtz & Howden, Neosisyphus quadricollis Gory. Another
subgroup extends from the south-western coast of the Western Cape along the
southern coastline to the Eastern Cape, Sarophorus tuberculatus (Castelnau),
Onthophagus giraffa, O. minutus, with further species restricted to the southern
coastline, Copris jacchus (Fabricius), Onitis minutus Lansberge, Scarabaeus
savignyi M’Leay. A number of species also occupy restricted or wider distributions in the south-west of the Western Cape, Onthophagus immundus
Péringuey, Epirinus comosus Péringuey, Scarabaeus spretus zur Strassen. Finally,
small numbers of species occupy arid ranges in Namaqualand, Scarabaeus
alienus Péringuey, Byrrhidium namakwensis Scholtz & Howden, B. ovale Harold,
or ranges restricted to the central Eastern Cape, Scarabaeus ambulans, Chironitis
sp., Epirinus striatus Scholtz & Howden, at the limits, respectively, of the winter
and bimodal rainfall regions.
Cluster 2A comprises a loose assemblage of species ranging from the winter
and bimodal rainfall regions across the arid karoo into the late summer rainfall
region (Figs 2, 3, Table 5). These species comprise both spring-active elements,
Epirinus aeneus, Onthophagus cameloides (Davis, 1993), which have probably
expanded their ranges to the north-east, and summer-active elements, Chironitis
scabrosus, Onitis aygulus, which have expanded their ranges to the south-west.
There are also some karoo-centred elements, Scarabaeus viator, Euonthophagus
vicarius, Onthophagus albipennis Péringuey, O. suturalis Péringuey. This range
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
32
A. L. V. Davis
Table 5. Climatic distributions of 14 major geographical species groups defined from dendrograms (Figs
2, 5, 7, 8)
Mean standardized percentage climatic distribution of species&S.D.
Species
clusters
Winter
rainfall
Bimodal
rainfall
2A
2D
34·8
&28·1
75·8
&23·3
6·6
&6·3
0·0
29·4
&9·5
19·5
&18·4
6·6
&7·7
0·0
2E
0·0
0·0
2F
0·0
5A
3·8
&10·0
0·0
5B
0·0
0·0
5D
0·0
0·0
5G
7A
2·2
&5·2
0·0
20·1
&16·7
0·0
7C
0·0
0·0
7D
0·2
&1·2
2·2
&4·2
0·1
&0·7
15·0
&6·0
2B
2C
8A
0·0
Late
summer
rainfall
31·6
&19·8
1·0
&2·8
69·9
&20·6
24·5
&17·7
13·7
&11·4
38·6
&11·3
0·0
0·8
&2·6
0·0
3·6
&6·5
6·5
&4·8
3·4
&3·1
0·1
&0·4
2·8
&3·3
Mid-summer rainfall
Subtropical/
tropical
Number
of
species
Kalahari
Highveld
II3d
East
coast
1·1
&2·5
0·5
&1·4
9·5
&7·0
45·9
&8·8
46·8
&13·0
41·1
&8·9
3·7
&5·4
5·6
&9·8
0·7
&2·6
0·3
&1·0
29·8
&5·3
14·4
&7·5
6·0
&5·0
5·0
&4·1
3·0
&5·6
3·2
&4·3
1·3
&2·3
0·0
0·0
0·0
0·0
6
0·0
0·0
0·0
8
1·8
&4·9
0·0
0·0
8
7·5
&10·3
6·0
&10·0
1·5
&4·7
2·2
&6·8
0·0
5·0
&7·0
0·8
&2·8
18·4
&23·2
37·7
&23·7
95·1
&5·7
6·6
&9.4
28·6
&3·8
20·8
&9·8
25·4
&6·4
15·9
&5·0
4·3
&4·3
29·6
&8·8
22·1
&10·7
7·9
&8·1
76·4
&25·4
53·7
&25·4
3·7
&5·4
1·4
&2·5
15·3
&3·6
28·3
&6·0
23·7
&5·8
11·9
&5·4
4·8
&9·2
2·0
&4·3
0·0
0·0
0·5
&1·7
50·6
&17·6
0·0
3·9
&4·6
5·5
&6·3
19·7
&8·2
15·3
&16·5
19·8
&5·3
29·2
&5·4
39·1
&9·7
27·4
&8·0
0·0
3
8
13
11
11
13
27
5
13
34
16
largely equates to the southern part of the central arid zone (Endrödi-Younga,
1978).
Cluster 2C is centred on the Karoo and arid regions of Namibia (Figs 2, 3,
Table 5). This region equates to the south-western zone plus the southern
and western parts of the central arid zone (Endrödi-Younga, 1978). It also
equates to the arid late summer rainfall region where 69% of observations were
made. Various species extend throughout the entire Karoo/Namib system,
Chironitis audens, Phalops euplynes, Gymnopleurus sericatus, although blue
and green varieties of G. sericatus appear in the southern Karoo in contrast
to the more widespread copper-red variety. Other distribution patterns
include, south-western Angola to the Northern Cape, Gymnopleurus andreaei,
Onthophagus semiflavus Boheman, and southern Namibia to the Karoo,
Onthophagus probus, O. ochropygus d’Orbigny. The relict species, Gymnopleurus
asperrimus, is centred on the Namibian/Northern Cape border zone whereas
? East African Wild Life Society, Afr. J. Ecol., 35, 10–38
Associations of southern African dung beetles 33
Phalops densegranosus d’Orbigny and Onthophagus phalopsides Frey have
only been recorded in arid climate type III2 at the edge of the Namib Desert
(latitudes 20)–25)S). Despite some regionalism, the composite parts of the
Karoo/Namib system probably form a distinct biogeographical entity in terms of
dung beetles.
Although the south-western zone (Endrödi-Younga, 1978) holds good for
Tenebrionidae which have diversified in arid climates, it is less valid in terms of
dung beetles which have limited diversification in deserts. There is a flightless
group of related species (formerly the genus Pachysoma, now synonymized with
Scarabaeus (Mostert & Holm, 1982) which is endemic to the arid west coastal
sands from the Western Cape to Swakopmund (22)33*S, 14)35*E) in Namibia.
However, cluster analysis (Davis, 1990) of numerical taxonomic data for these
species (Holm & Scholtz, 1979) shows that there are three distinct taxonomic
groups. Essentially, these are centred on the south-west of the Western Cape
(winter rainfall), Namaqualand/southern Namibia (winter/summer rainfall), and
southern Namibia (summer rainfall).
Cluster 2D occurs mainly on finer-grained soils in the moister northern
Namibia limited by arid climate to the west/south-west and by lower-lying
Kalahari sands to the east/south-east (Figs 2, 3, Table 5). Other species showing
a north Namibian distribution include, Onitis mnizechi Lansberge, O. obscurus,
Phalops prasinus Erichson, P. pyroides d’Orbigny. Some of these species show
a distribution extending into Angola, O. obscurus, P. prasinus. A few species
occurring as rarities in Etosha National Park, Neosisyphus macroruber,
Proagoderus lanista, show a disjunct distribution from their principal range,
respectively, in the Northern Cape and the highveld, and on the highveld
southwards to the Eastern Cape. A similar disjunct distribution is shown by
Scarabaeus ambiguus and Catharsius ulysses (cluster 2E), which are abundant
at the edge of the Kalahari both in northern Namibia and in South Africa.
These northern Namibian outliers may be relicts of Pleistocene northwards
expansion.
The biogeographical status of the Kalahari region is debatable. It may be the
north-eastern part of the central arid zone (Endrödi-Younga, 1978) or a poorly
differentiated subregion of the summer rainfall, savanna region (Holm &
Scholtz, 1983, 1984). Although it cannot be regarded as a distinct biogeographical region (Barker, 1993), there is a definite cluster (2F) of sand specialist dung
beetle species centred to the south-west of the Kalahari in the Northern Cape
and south-western Botswana (Figs 2, 4, Table 5). This suggests that the arid
south-west does act as a biogeographical focal point combining the selective
factors of climate and soil type. Furthermore, this centring to the south-west has
been noted in other families of Coleoptera including, Buprestidae, Carabidae
and Trogidae (Holm & Scholtz, 1983). Although zoogeographical affinities of
these families are to the north-east (Holm & Scholtz, 1984), at least one group
of dung beetles has speciated between the Kalahari (Scarabaeus flavicornis),
southern Namib (S. fritschi Harold) and Namaqualand/Western Cape regions
(S. canaliculatus Fairmaire) (Davis, 1993). Similar observations have been made
for south-western Kalahari Tenebrionidae whose ancestral relationships are
largely to the north-east with a minority showing south-westerly Namaqualand
affinities (Penrith, 1984). Endemicity is low, as in grasshoppers (Acrididae)
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A. L. V. Davis
(Barker, 1993), since most of the Kalahari-centred dung beetles show distributions outside of the Kalahari as defined by Barker (1993). Extended distributions occur, especially, in sand outliers in Gauteng and Northern Province,
Pachylomerus opaca, Copris cassius, Proagoderus sappharinus; southwards
through Namaqualand along the west coast of the Western Cape, Scarabaeus
proboscideus, Metacatharsius latifrons; and northwards into Angola, Proagoderus
sappharinus var., Copris cassius ssp. angolensis Ferreira.
Clusters 2E and 7A are loose groupings of species which each occur in three
or four different clusters in the purely geographical analysis. Their circumKalahari or northern KwaZulu/Natal centres of distribution (Table 5) reflect a
bias to greater frequency on tropical sands. Most species show a relatively broad
distribution in the mid-summer rainfall region whereas some with more widespread biogeographical affiliations are found at the northernmost limits of the
mega-Kalahari sands (Thomas, 1988) in Zaı̈re, Pachylomerus femoralis, Kheper
lamarcki (Walter, 1978).
The highveld/bimodal rainfall (5G) and east coast (5D) clusters (Figs 4–6,
Table 5) are centred, respectively, in the eastern part of the mountain zone and
the south-eastern coastal region of the tropical zone of Endrödi-Younga (1978).
Several highveld species are widespread beyond the regional centre from the
Northern Province/Mapumalanga highlands to climate Type V in the bimodal
rainfall region, Euoniticellus africanus, Onthophagus asperulus. Other species
occurring the entire length of the region are either distributed along the high
rainfall zone to the east of Lesotho, Sisyphus costatus, Litocopris simplex, or
largely distributed in the drier regions to the west of Lesotho, Copris antares,
Gymnopleurus leei. Further species are restricted to the summer rainfall highlands occurring from Northern Province southwards beyond Lesotho, Copris
jacchoides, C. corniger. Elements from probable northwards expansion of the
Cape canthonine fauna occur in the extreme highland regions of the KwaZulu/
Natal Drakensberg and the Mapumalanga highlands, Epirinus asper Péringuey,
E. mucrodentatus Scholtz & Howden, and at lower altitudes in more southerly
latitudes, Epirinus obtusus. Few highveld species occurred in the eastern
highlands of Zimbabwe where endemism was limited to four species, Onitis
autumnalis Davis, Heliocopris marshalli Péringuey, Xinidium davisi Cambefort
and Copris sp. nr integer. The east coast endemics comprise mainly sand
specialists with a few clay specialists, Onthophagus beiranus, Kheper clericus
(Boheman) (Doube, 1991). Some of these species are distributed from southern
KwaZulu/Natal to central northern Moçambique, sand: Copris puncticollis,
Proagoderus aciculatus, clay: Onthophagus beiranus. However, most are
centred on the high rainfall coastline of northern KwaZulu/Natal and southern
Moçambique where there is appreciable overlap between species of the temperate
to tropical (5A, 5B, 7A, 7C, 7D, 8A) and highveld (5G) clusters. Species of the
east coast cluster show tropical or highveld affinities. For instance, Onthophagus
juvencus, is a pan-African tropical element and the dune forest endemic,
Scarabaeus bornemisszai zur Strassen, is closely related to a highveld species,
Scarabaeus caffer (Boheman) (zur Strassen, 1980). The presence of Copris
inhalatus Quedenfeldt ssp. santaluciae and Scarabaeus galenus var. in the
St Lucia (27)45*S 32)30*E) region also indicate recent links with the dry sandveld
of the interior.
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Associations of southern African dung beetles 35
Cluster 5B shows a widespread but fragmented distribution throughout the
mid-summer rainfall region (Fig. 6). Collections of these species were made
primarily in game reserves (71·3%, n=171) with an extreme bias to greater
frequency on non-ruminant dung (Fig. 5). Of the 28·8% of collections of these
species made outside of reserves, 12·1% (n=29) were made on non-ruminant
dung and 16·7% (n=40) on other food types. This distribution reflects the
fragmented range of non-ruminant mammals.
Clusters 5A, 5B, 7C, 7D and 8A show primarily (78·8%) broad tropical
biogeographical affiliations (Figs 4–8, Table 5). Most groups are centred on the
trans-Botswana transitional area and the tropical subregion of Endrödi-Younga
(1978) without any obvious faunal separation between the two regions. Cluster
8A comprises species showing pan-southern African or pan-eastern southern
African distributions which are considered to be mostly recent elements. There is
a reduction in the size of ranges from that occupied by cluster 8A to those of the
the pan mid-summer rainfall region (7C) and game reserve (5B) clusters. A
further reduction in range is shown by the largest cluster of species (7D) which
occupies the moist eastern part of the mid-summer rainfall region and by tropical
species (5A) which are largely centred on climate type II2B (45·8% of observations) in eastern Zimbabwe and central Moçambique. Thus, in widespread
species, there is a regression in distribution from tolerance of mild temperate
conditions to restriction to moist tropical regions.
The biogeographical analysis suggests that southern Africa comprises six
principal centres which focus regional distributions. These are, the southwesterly winter and bimodal rainfall zone comprising up to five subzones, the
western arid zone of the Namib, Namibia and the Karoo, the south-western
Kalahari arid zone, the south-eastern highland zone, the east coastal zone and
the warm temperate to tropical zone in the north and north-east. Some of these
zones are centres of generic endemism which is concentrated along the coastal
regions and at high altitude in the highlands. Endemic Gondwanaland genera
occur in the winter and bimodal rainfall regions, Dichotomiini: Macroderes
(Ferreira, 1972), Canthonini: Aphengoecus, Byrrhidium, Outenikwanus (Scholtz &
Howden, 1987b). Others occur in the Drakensberg, Canthonini: Peckolus
(Scholtz & Howden, 1987b) or in the northern highveld and eastern highlands
of Zimbabwe, Dichotomiini: Xinidium (Cambefort, 1985). Endemic onitine
genera occur on the subtropical east coast associated with non-ruminant dung,
Anonychonitis in Hluhluwe and Umfolozi Game Reserves, Tropidonitis in
north KwaZulu/Natal coastal sandveld. Endemism in the scarabaeine genus,
Drepanopodus, is centred around the south-western Kalahari, Namaqualand.
Biogeographical associations of the dung beetle fauna have probably been
shaped principally by the reduced influence of the easterlies and increased
influence of the westerlies since the Pliocene &3 My ago (Deacon, 1983).
Pleistocene climatic oscillation has been responsible for waves of penetration of
temperate elements to the north and tropical elements to the south (EndrödiYounga, 1978). However, winter rainfall climate is not thought to have ever
shifted more than 1) of latitude further north than at present (Deacon et al.,
1992). Northern expansion has occurred particularly along the cooler eastern
highlands and the coastline with outlier relicts of cooler Pleistocene climate
occurring in the eastern Zimbabwe highlands and the northen Namibian
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A. L. V. Davis
highveld. Southwards faunal expansion in the present interglacial, dating from
10 000 years, has probably been enhanced by recent fragmentation of the natural
shrubland habitat in the winter rainfall region (Davis, 1993). Biogeographical
group distributions from the present analysis (clusters 2D, 7C, 7D) suggest that
the arid west and the central Kalahari sandveld basin comprise barriers to
southward movement by all but dry-tolerant and psammophile species. The
eastern highveld is less of a barrier but has clearly filtered out distinctly tropical
elements. Northward movement is presently limited by the Karoo/Namib,
Kalahari and tropical northern lowlands.
Acknowledgments
I thank Dr George Bornemissza, Dr Bernard Doube, Dr Jane Wright and my
other colleagues at the Australian CSIRO Dung Beetle Research Unit whose
efforts were responsible for the reference collection on which this study is based.
The collection is now part of the National Collection of Insects, Pretoria, South
Africa. Dr John Hoffmann kindly criticized the manuscript.
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Adrian Davis