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Page 47

and Langeberg ranges, respectively, were later re-

ported by Endro¤dy-Younga (1988). In October 1977,

Endro¤dy-Younga found live specimens of C. izardi but

baited ground-traps failed to attract these beetles.

Subsequently, he did obtain live specimens of C. izardi

and C. montisatris which were observed for up to

84 days in the laboratory. The habitat of C. montisatris

in the Swartberg was limited to a discrete strip of about

400 m horizontally and 20 m vertically, while the

C. izardi population in the Langeberg occurred at sites

8 km apart and differing by about 400 m in elevation.

The latter species is known from �ve additional local-

ities at increasing distances from the observation area

(Endro¤dy-Younga 1988: Fig 10). The adult activity

period for C. izardi was from October to February

(Barnard 1929; Endro¤dy-Younga 1988). Endro¤dy-

Younga (1988) commented on the slow undirected

movement of males of both species in the �eld. Aspects

of the ecology of Colophon species were reported (in

Japanese) by Mizukami and Kawai (1997).

No ecological �eld studies have been done on any of

the Cape stag beetles. As a result, no published infor-

mation is available on population sizes, rates of pop-

ulation turnover, predation rates and ecological

requirements, although Bartolozzi and Werner (2004)

do refer to their respective conservation status, most

regarded as rare. It is thus dif�cult to assess the con-

servation status and the threat to these species

(Anonymous 1994). All species of Colophon have

highly restricted ranges, some more than others. Holm

in Anonymous (1994) refers to C. montisatris as

occurring at only one locality of a few ha in extent, and

estimates its total population at less than 1,000


In view of limited information on the preferred

habitat and behaviour of these beetles (Anonymous

1994) and concern over possible requests for erecting

repeater stations, the Western Cape Department of

Environmental and Cultural Affairs, Directorate Nat-

ure Conservation and Museums, granted special per-

mission to collect and study these beetles from speci�c

localities so as to contribute to the existing information


During January 1996, 26 individuals of C. stokoei

Barnard, 6 C. westwoodi Gray and 12 C. neli Barnard

were collected in the Hottentots Holland Mountains,

on Table Mountain and in the Riversdale Mountains of

the Western Cape Province, respectively. In all in-

stances, a small number of the beetles were observed

and some were collected (to protect populations in the

wild, precise locations have been withheld). Live

specimens were taken to the laboratory in Stel-

lenbosch, Western Cape, to study their behaviour.

Collected specimens are deposited in the insect col-

lection of the Department of Conservation Ecology

and Entomology, University of Stellenbosch (USEC).

Habitats of C. stokoei, C. neli and C. westwoodi

Localities for these beetles are usually given as Hot-

tenhots Holland Mountains or Swartberg Mountains

(Endro¤dy-Younga 1988). However, the habitat of these

beetles is far more restricted and usually con�ned to

glades at, or near the top of these mountains, but

sometimes at considerably lower elevations. The hab-

itat for C. stokoei in the Hottentots Holland Mountains

(1,150�1,550 m asl), and C. neli in the Swartberg

Mountains ( < 1500 m asl), consisted of sloping �at-

lands, ranging from horizontal to about a 30� inclina-
tion. These were covered with typical, but rather short,

montane fynbos vegetation, interspaced with rocks.

The habitat (1,000 m asl) of C. westwoodi was fairly

marshy �atland, with taller restioid vegetation (30�

50 cm; dominated by Elegia spp.) on dark, poorly-

drained soil presumably with a high humus content. In

1992, specimens of C. westwoodi were also observed

near the De Villiers Reservoir on Table Mountain in a

similarly vegetated habitat. During one visit in mid

January 1996, 12 trampled specimens of C. westwoodi

were found on a path often used by tourists, close to a

favourable habitat. On Table Mountain, little evidence

of lizard predation was found, perhaps due to the

unsuitability of the habitat for lizards. However, a scat

of a predator, most probably that of the Small Gray

Mongoose [Galerella pulverulenta (Wagner)], contain-

ing the remains of C. westwoodi, was found.

The habitat of C. montisatris, at 2,085 m asl, con-

sisted of a tiny area of about 400 m horizontally and

20 m vertically in the Swartberg mountain range.

Outside this area not even fragments of this species

could be found and it is likely that this species is on the

verge of natural extinction unless the regional climate

soon turns more humid and cooler. In the Langeberg

Mountain range (at between 1300 and 1600 m asl),

C. izardi was found at sites 8 km apart and differing in

altitude by about 400 m. Of the two ranges, the

northern (Swartberg) is the drier, with less frequent

cloud cover, higher summer temperatures and a long

dry spell in summer. Environmental differences,

including that of vegetation, soils, exposure, etc., are

considerable between these two habitats (Endro¤dy-

Younga 1988). Although Colophon species are con-

�ned to certain localities, C. oweni occurs sympatrically

with C. izardi on the east side of Tradouw�s Pass in the

Riversdale Mountains of the Western Cape Province

44 J Insect Conserv (2007) 11:43�46


Page 48

and C. thunbergi is found on the west side of this pass

(Bartolozzi 1995). The contribution by Mizukami and

Kawai (1997) provide good pictures of the most of the

habitats of various Colophon species; unfortunately,

their detailed ecological notes on the genus remain

obscure, being presented in Japanese.


The adult activity period for Colophon ranges mainly

from October to March (Barnard 1929; Endro¤dy-

Younga 1988, own observations), i.e., the whole of

summer. Females were rarely collected: �ve males and

one female C. westwoodi, 12 males C. neli and 3 fe-

males and 23 males C. stokoei were collected. The ratio

of females to males is extremely low. Possibly males

are more active and thus more easily spotted than fe-

males, or sex ratios �uctuate during the season. It is

also possible that females spend more time in hiding,

while males search actively for females. In view of the

long emergence period and life span, this is the most

likely explanation. All beetles were collected in the

middle of their emergence period, and it may well be

that females appear earlier than males. Endro¤dy-

Younga (1988) found specimens, especially females,

with strongly eroded fore tibiae, indicating a long life

span similar to that shown in captive Tenebrionidae.

A C. stokoei larva was found about 20 cm depth in a

mixture of washed-down soil and nearly completely

rotten plant material, retained by and lodging against

the rocks on Table Mountain. Rearing it to the adult

stage proved unsuccessful. Two third-instar larvae of

C. neli were previously found by one of the authors

(CRO) in humus-rich soil. The larva of C. neli was

described subsequently by Scholtz and Endro¤dy-

Younga (1994).

Activity patterns

Adult beetles generally became active from late

afternoon (17 h) to early nightfall (20 h), but appeared

earlier when mountains were covered in cloud, or

misted over due to the south-easterly winds experi-

enced during the hot, dry summer months in the

Western Cape Province. When inactive, beetles hid in

the vegetation adjoining vertical rock faces and were

sometimes found under rocks. Movement in vegetation

was slow, making the observation of beetles dif�cult.

However on open ground movement was fairly rapid

and occurred in random directions until shelter was

found. This concurs with the observations of Endro¤dy-

Younga (1988).

Colophon species studied elsewhere have also been

found to be decidedly diurnal in activity. Specimens of

C. izardi have been observed in thick fog after sunset

or at about 16 h in drizzling rain and biting cold wind

(Endro¤dy-Younga 1988). Most specimens have been

seen during the late afternoon hours, often after sunset

but before dark (Endro¤ dy-Younga 1988). This con-

trasts with the observations of Barnard (1929) who

collected most of his specimens between 6 and 8 h and

at 14 h in the hot midday sun.

Behaviour and longevity of captive beetles

Beetles collected in the �eld were placed outdoors in 5l

plastic containers, half-�lled with compost-rich garden

soil during mid January 1996. Shelter, provided by

supported tile shards, was rapidly occupied by beetles

during the day. Locomotory activity, at ambient tem-

peratures of 17�22�C, commenced from 17 h onwards
and was directed away from the provided shelter. Males

encountering other males would attempt to push them

aside, or use their mandibles to attack them. Many

specimens, both in the �eld and in captivity, had lost

parts of their legs and antennae during such encounters.

When confronted by a female, mating was attempted.

The male, or males, would approach the female from

the rear, using their strong fore legs as pincers to hold

the female between the pro- and mesoterga. Coupling

lasted from a few minutes to more than two hours.

However, no eggs were produced. Endro¤dy-Younga

(1988) reported that attempts were made to provide

food, including a range of plants from their habitat, but

none of the captive specimens were observed to feed,

unless on the vegetation debris that was richly supplied

in the soils brought with them (Endro¤dy-Younga 1988).

Whilst conducting our research, beetles were only

interested in droplets of water. No feeding was ob-

served on the supplied compost from their localities,

but one beetle showed some interest in a slice of apple.

It could well be that in nature they ingest organic soil

matter as it is most unlikely that these beetles would

survive for months without food.

Colophon westwoodi beetles, collected on 6 January

1996, died after 18�20 days (2 females) and between 23

and 32 days (4 males)(mean 26.25 days), respectively.

Six males of C. neli, collected on 7 January 1996, died

after 24�32 days (mean 26.50 days), respectively.

C. stokoei were collected on 1 and 22 January 1996, of

which males died after 11�56 days (mean 32.48 days,

n = 23) and females after 29�61 days (mean

41.67 days, n = 3). Endro¤dy-Younga (1988) kept

C. montisatris alive for 84 days.

J Insect Conserv (2007) 11:43�46 45


Page 93

are posing to the ecological services coccinellids pro-

vide. In fact there may be evidence that adventives have

actually helped coccinellids regulate aphids in managed

systems. Two recent studies have documented long-

term declines in aphid densities since the establishment

of an adventive species (Alyokhin and Sewell 2004;

Evans 2004). Moreover, one adventive coccinellid, H.

axyridis, is regarded as one of the only established

natural enemies capable of regulating a major recent

adventive pest in North American agriculture (Rutl-

edge et al. 2004). More work is needed throughout

native ecosystems to understand what additional risks

adventives may pose to coccinellids and their functions

in these less well-studied environments.

Our case studies on the decline of A. bipunctata

and C. novemnotata illustrate that at least some spe-

cies in some systems have recently undergone exten-

sive declines. Additional circumstances such as

changing land use practices may have helped exacer-

bate situations, but in both cases it seems extremely

likely that adventive species played a role. What is

needed now is a better understanding of why these

two species have undergone more substantial changes

than other natives. Are there speci�c characteristics in

their physiology, ecology, or habitat use that put

coccinellid species more at risk, or are there speci�c

types of habitats, ecosystems, or landscapes that put

any species more at risk than others? While these are

not new questions to conservation or risk assessment,

coccinellid ecology would bene�t greatly from the

application of these other disciplines� current tools

and theories. Current invasions of coccinellids in

Europe, Asia, and Australia highlight the need to

understand what risks are posed to native coccinellids

and what can be done to emasculate the negative

effects of adventive species.

Acknowledgements The authors wish to thank Timothy New
for the opportunity to publish within this special section. We also
wish to acknowledge Rachel Clancy for her assistance in gath-
ering data and Tony Ives for his statistical advice. JPH was
funded by NSF grant EF-0313737.


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