how localized are the distributions of australian scarabs (coleoptera, scarabaeoidea)?

Diversity and Distributions (1999) 5, 143–149

BIODIVERSITY RESEARCH

How localized are the distributions of Australian scarabs(Coleoptera, Scarabaeoidea)?

P. G. ALLSOPP Bureau of Sugar Experiment Stations, Private Bag 4, Bundaberg DC 4670, Australia, E-mail:[email protected]

Abstract. The degree of localization in the However, the Australian fauna lacks the highproportion of widespread species characteristic of thedistribution of Australian scarabaeine (320 spp.),

aphodiine (160 spp.), geotrupid (166 spp.) and West Palaearctic Scarabaeinae, Aphodiinae andGeotrupidae. This difference reflects the differentdynastine (182 spp.) scarabs is analysed. Localization

differs among these groups, appearing to increase ecological histories of the two regions.marginally from the Dynastinae, to the Aphodiinae,to the Scarabaeinae, to the Geotrupidae, but with Key words. Localized species distributions,

endemic, relictual, Australia, Aphodiinae, Dynastinae,significant differences within groups, e.g. thescarabaeine tribes Onthophagini and Scarabaeini. Geotrupidae, Scarabaeinae.

INTRODUCTION the range of localization of Australian scarabs isanalysed in the same three groups, the Scarabaeinae,

Species diversity is not uniformly distributed across Aphodiinae and Geotrupidae, and for comparison inthe planet’s surface, being characterized by disharmony one of the phytophagous scarab groups, theand impoverishment (Quammen, 1996). Many species Dynastinae.have very limited distributions. These species are oftensaid to be endemic (Horton, 1973; Lumaret & Lobo,

MATERIALS AND METHODS1996), but this term is misleading and misused. Firstly,because it is relative, depending on the spatial scale

The Australian (including Tasmania and nearby off-considered. Secondly, because endemism (originated in

shore islands) and the West Palaearctic (includingthe same place as it is found) is one of two routes by

Britain, Ireland and Mediterranean islands) regionswhich a species’ present distribution can be limited;

have similar land areas, about 7.7×106 andthe other is relictualism (surviving in a given place

12×106 km2, respectively. To allow direct comparisonwhilst disappearing elsewhere) (Quammen, 1996). A

with Lumaret & Lobo’s (1996) analyses, I includedbetter term to describe a species with an observed

Tasmania and other major Australian islands (Lumaretlimited distribution is localized.

& Lobo included Britain, Ireland and MediterraneanAs noted by Lumaret & Lobo (1996), the study of

islands) and defined (Fig. 1) the same six geographicallocalized species is of great use, both for reconstructing

range-size classes as used by them:the biogeographic history of life and areas and for

I species with a distribution in an area less than ormaking decisions about the conservation of areas andequal to 0.1% of the total surface area of Australiaspecies. Lumaret & Lobo (1996) analysed the range of(7.7×103 km2);localization in three groups of scarabaeoid dung beetles

II species with a distribution in an area between 0.1%from the West Palaearctic region and showed that thereand 0.5% of the total surface area of Australiawas significant variation among the groups. This they(3.9×104 km2);attributed to differences among groups in tolerance to

III species with a distribution in an area betweenenvironmental variables and dispersal capacity. Are0.5% and 1% of the total surface area of Australiathese differences the same in a different region with a

different biogeographic history? To help answer this, (7.7×104 km2);

143 1999 Blackwell Science Ltd. http://www.blackwell-science.com/dad

144 P. G. Allsopp

Fig. 1. Sizes of the first five range-size classes defined for Australia.

IV species with a distribution in an area between 1% arrived in Australia in the last 210 years were excluded.and 5% of the total surface area of Australia I used a system of paper chips to estimate range sizes,(3.9×105 km2); similar to that used by Lumaret & Lobo (1996).

V species with a distribution in an area between 5% Kolmogorov–Smirnov two-sample tests (Steel &and 10% of the total surface area of Australia Torrie, 1980) were used to assess the hypothesis of(7.7×105 km2); independence among geographical range-size classes

VI species with a distribution in an area greater than and each of the Australian groups (including a10% of the total surface area of Australia. comparison between the Onthophagini and

Scarabaeini). The test was also used to assess theLumaret & Lobo (1996) considered a species localizedhypothesis of independence for each of theif its distribution fell into classes I–IV.Scarabaeinae, Aphodiinae and Geotrupidae of theI took the ranges of the species from: AphodiinaeAustralian and West Palaearctic regions. The(Aphodiidae of Lumaret & Lobo, 1996) – Stebnicka &Kolmogorov–Smirnov test is preferable to the chi-Howden (1994, 1995, 1996, 1997) and Storey & Howdensquare test because it exploits information in the(1996); Dynastinae – Allsopp (1990, 1993), Allsopp &ordering of the categories as well as means andCarne (1986), Carne (1957, 1976, 1978, 1980, 1981,variances (Analytical Software, 1996). However, I used1985a,b), Carne & Allsopp (1987) and Endrodi (1977,the chi-square test to assess the distributions for a1978); Scarabaeinae (Scarabaeidae of Lumaret & Lobo,theoretical evenness of species among range-size classes1996) – Hill (1993), Matthews (1972, 1974, 1976),for each of the major Australian groups; in this caseMatthews & Stebnicka (1986), Storey (1977, 1984, 1986,the ordering of the classes is immaterial and the chi-1991) and Storey & Weir (1988, 1990); Geotrupidae –square test is less conservative than theCarne (1965), Howden (1974, 1975, 1979, 1985a, 1992),

Krikken (1976) and Nikolajev (1990). Species that have Kolgomorov–Smirnov test.

1999 Blackwell Science Ltd, Diversity and Distributions, 5, 143–149

Localized distributions of Australian scarabs 145

Fig. 2. Distribution of species belonging to the Australian Scarabaeinae, Aphodiinae, Geotrupidae and Dynastinae in the sixrange-size classes. The number of species in each class is given above the bar. Distributions for the West Palaearctic Scarabaeinae,Aphodiinae and Geotrupidae (from Lumaret & Lobo, 1996) are given as insets.

RESULTS In the Scarabaeinae, the observed distribution amongrange-size classes differs significantly from speciesdistributed evenly among classes (v2=47.08; d.f.=5;The distribution of species among the six range-size

classes differs significantly across the Australian groups P < 0.0001) (Fig. 2). The distribution among range-sizeclasses differs significantly from that shown by the(Fig. 2). Distribution of the Geotrupidae differs from

each of the other three groups (Geotrupidae- West Palaearctic Scarabaeinae (D=0.63, P < 0.0001)(Fig. 2), where the distribution is strongly skewed toScarabaeinae D=0.16, P=0.0062; Geotrupidae-

Aphodiinae D=0.17, P=0.016; Geotrupidae- the right. However, there is a significant difference inrange-size distributions between the two larger tribesDynastinae D=0.16, P=0.019). The distribution of

the Aphodiinae does not differ significantly from that of of Australian scarabaeines the Onthophagini andScarabaeini (D=0.38, P < 0.0001) (Fig. 3); the thirdthe Scarabaeinae (D=0.08, P=0.60) or the Dynastinae

(D=0.10, P=0.31), but there is a weak trend to a tribe, the Coprini, has too few species (29) to allowadequate comparison. The distribution of the classesdifference in class distribution between the

Scarabaeinae and the Dynastinae (D=0.11, P=0.097). in the Scarabaeini is strongly skewed to the left, whereas


146 P. G. Allsopp

significantly from that shown by the West PalaearcticGeotrupidae (D=0.21, P =0.043) (Fig. 2).

In the Dynastinae, the observed distribution amongrange-size classes differs significantly from speciesdistributed evenly among classes (v2=28.96; d.f.=5;P < 0.0001) (Fig. 2), but with no strong skewness.

DISCUSSION

The evolution of the Australian fauna is dominated bythree major influences (Main, 1987). Australia wasformerly part of Gondwanaland, but broke the lastconnection during the early Tertiary. It was isolatedfrom that time until its collision with Asia during theMiocene. After the Miocene, faunal elements of Asianorigin have become established in Australia. Duringthe late Mesozoic era Australia was characterized bythree or four large emergent areas which probablyrepresent centres of speciation of some of the olderelements of the scarabaeoid fauna (Howden, 1981).This changing archipelago, particularly during theCretaceous, can explain much of the earlier radiation,resulting in many endemic genera and species groups.Odd relicts found in some of the interior sandy areascan also be best explained by this scenario. Evolutionin Australia during the isolated Tertiary period washeavily influenced by the continent’s northward driftthrough different climatic zones (Heatwole, 1987),together with major transgressions and regressions ofthe sea (Howden, 1981). During the Palocene, climatevaried from cool to warm temperate with a high rainfall,

Fig. 3. Distributions of species belonging to the Australianto subtropical or tropical. During the Eocene, AustraliaOnthophagini and Scarabaeini in the six range-size classes.maintained widespread humid conditions, even in theThe number of species in each class is given above the bar.south. Marked cooling and a reduction in rainfallcharacterized the Oligocene. The Early Miocene waswarmer and moist, but in the Middle to Late Miocenethere is no strong skews in the distribution of the

Onthophagini or the Coprini. conditions became cooler and rainfall decreased. ThePliocene saw wide fluctuations in climate, with a brief,In the Aphodiinae, the observed distribution among

range-size classes differs significantly from species warm, moist period, followed by progressive aridityand perhaps coolness over most of the continent withdistributed evenly among classes (v2=20.80; d.f.=5;

P=0.0009) (Fig. 2), but with no strong skewness. The wetter and warmer conditions perhaps persisting in thesouthern coastal regions. The current relict pattern ofdistribution among range-size classes differs

significantly from that shown by the West Palaearctic many species supports the idea of persistence of habitatthrough the Tertiary period and the development ofAphodiinae (D=0.33, P < 0.0001) (Fig. 2), where the

distribution is strongly skewed to the right. flightlessness because of restrictions into relatively smallareas (Howden, 1981). The Pleistocene saw furtherIn the Geotrupidae, the observed distribution among

range-size classes differs significantly from species fluctuations, coincident with glacials and interglacialsat higher latitudes and altitudes, as well as the invasiondistributed evenly among classes (v2=27.20; d.f.=5;

P < 0.0001) (Fig. 2), but with no strong skewness. The of Oriental taxa as Australia neared the Asiancontinent. The different extents of these three majordistribution among range-size classes differs



influences are reflected in the different occurrence of plants nor to animal dung, soil type and rainfall appearto be the most limiting factors (Howden, 1981). Manylocalized species amongst the groups of scarabs that I

analysed. The degree of localization appears to increase of the species from the Eyrean subregion appear to berelicts, surviving over long periods as the continentmarginally from the Dynastinae, to the Aphodiinae,

to the Scarabaeinae, to the Geotrupidae. became drier (Howden, 1981) – this may account forthe high proportion of species in range-size class IV.Much of the Australian dynastine fauna appears to

have its origin in the pre-Jurrasic to Jurrasic periods The distribution of range-size classes of theAustralian and West Palaearctic faunas differs greatly.and is concentrated in the arid Eyrean subregion

(Carne, 1957; Howden, 1981; Allsopp, 1995). The much Why is this so? In Australia, the significant, but notsevere, fluctuations in climate during the Mesozoic andyounger and much smaller Malayan element is

concentrated in the north and north-east (Torresian Cenozoic eras presumably led to isolation ofpopulations and evolution of new species. In contrast,subregion) and derives from ancestors arriving in the

Cenozoic, presumably after the Miocene. Speciation in the West Palaearctic fauna was heavily influenced bythe severe glacials and interglacials of the Pleistocene,the older element presumably benefited from the

climate fluctuations during the Mesozoic and early which necessitated recolonization of large areas(Hanski, 1991). This is most noticeable in the cool-Cenozoic eras.

The Australian aphodiines were poorly studied until susceptible Scarabaeinae, which is dominated bywidespread, dispersing species; the more cool-tolerantrecently. Howden (1981) speculated that, because they

are generally the smallest Australian scarabs, they Aphodiinae has a much higher proportion of localizedspecies. The Geotrupidae is the least cool-tolerant ofare the more widely dispersed. My analysis does not

support this, and the revisions of the Australian the groups analysed by Lumaret & Lobo (1996) andits degree of localization appears to be heavilyaphodiines by Stebnicka & Howden (1994, 1995, 1996,

1997) recognized a large number of localized species. influenced by the flightless genus Thorectes and thegroup’s basal lineages in the western Mediterranean.Among the indigenous Australian Aphodiinae there

are a number of forms that do not appear to have A further factor that has undoubtedly influenced thedistributions of both faunas is the impact of humans.close relatives elsewhere. Species of both old and recent

elements are concentrated around the south-eastern Within Australia, habitat destruction has beenimportant in eliminating species from the Australianand eastern periphery of the continent (Stebnicka &

Howden, 1995). Many of these appear to have had a fauna and in endangering the survival of others(Heatwole, 1987). There is no evidence that anylong residency in Australia, similar to the Dynastinae.

The Scarabaeinae, as a whole, have a large Australian scarab has become extinct, but one canspeculate that large species of Scarabaeinae may haveproportion of localized species. However, differences

among groups, especially the Onthophagini and been associated with the dung of larger native mammalsin the same way that the large Heliocopris spp. areScarabaeini, are even more evident than in, say, the

Dynastinae. The extreme tendency towards localized associated with elephant dung in Africa. These largenative mammals went extinct at the close of thespecies within the Scarabaeini reflect the group’s long

residency in Australia and a high proportion of Pleistocene and any associated large dung beetles mayhave disappeared at the same time. However, otherflightless species, reflecting limitation of habitat area

coupled with environmental stability (Matthews, 1974; species have benefited from changes to habitat,becoming more common (e.g. scarabs associated withHowden, 1981). Concentration of species along the

periphery of the continent (Matthews, 1974; Allsopp, sugarcane, Allsopp & Logan, 1999), if not more widelydistributed. Similar changes have probably occurred1995) probably reflects early expansion and later

extinction in unfavourable habitats. Many of the in the West Palaearctic region.In summary, the occurrence of localized speciesOnthophagini, in contrast, seem to be good dispersers

and their ancestors apparently arrived much more differs among groups of Australian scarabs, reflectingthe continent’s past isolation and recent exchange fromrecently from the north in less than 34 separate

incursions (Matthews, 1972; Howden, 1981). Asia. The occurrence is very different from that in thesame groups of West Palaearctic scarabs, lacking theThe Geotrupidae generally has patterns like the older

elements of the scarabaeine subfamilies. Many species high proportion of widespread species. The comparisonillustrates Howden’s (1985b) contention that bothare cool or cold tolerant and occur in mesic and arid

environments. Because they are not tied to various evolutionary time and ecological attributes are


148 P. G. Allsopp

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how localized are the distributions of australian scarabs (coleoptera, scarabaeoidea)?

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