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Feeding and mating behavior of Dorcacerus barbatus (Olivier, 1790) (Coleoptera: Cerambycidae) on Lantana camara L. (Verbenaceae)

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In this study we describe the mating behavior of Dorcacerus barbatus (Olivier, 1790) and its predation on the reproductive parts of Lantana camara L. (Verbenaceae) in a restored area of Atlantic forest, in southeastern Brazil. Lantana camara is regarded as one of the world's ten worst weeds. We found three species of longhorn beetles on this plant: Trachyderes succintus duponti Aurivillius, 1912, Andraegodius rufipes zonatus (Dalman, 1823), and D. barbatus; this last species represented more than 95% of all individuals found. This is the first record of these three cerambycid species on L. camara. We observed a sequence of mating-behavior stages: jousting, antennation, holding, mounting, fighting, licking, abdomen bending, and copulation. D. barbatus destroyed 60% of the reproductive parts during the mating process. These primary observations indicate that D. barbatus may be considered as a potential biocontrol agent for this invasive species.
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Received: 04-IV-08
Accepted: 13-IV-09
Distributed: 18-IX-09
SHORT COMMUNICATION
Feeding and mating behavior of Dorcacerus barbatus
(Olivier, 1790) (Coleoptera: Cerambycidae) on Lantana camara
L. (Verbenaceae)
G. Wilson Fernandes & Jean C. Santos
Ecologia Evolutiva e Biodiversidade/DBG, ICB/Universidade Federal de Minas Gerais, CP 486, 30161-970 Belo Horizonte, Minas Gerais, Brazil.
Lundiana 9(2):155-158, 2008
© 2009 Instituto de Ciências Biológicas - UFMG
ISSN 1676-6180
Beetles of the family Cerambycidae have assumed increasing
prominence as pests of forest and shade trees, shrubs, raw wood
products, and as vectors of tree diseases (e.g., Berti-Filho, 1997;
Dall’Oglio & Peres-Filho, 1997; Hanks et al., 1998). Moreover,
cerambycids are also considered important herbivores of several
plant species (e.g., Monné 2001abc, 2002ab, 2004). Thus,
detailed knowledge is needed on the natural history, behavior
and ecology of most Neotropical species of this important insect
group. In this study, we describe the mating behavior of
Dorcacerus barbatus (Olivier, 1790) and its predation on the
reproductive parts of Lantana camara L. (Verbenaceae) in a
restored area of Atlantic rain forest, in southeastern Brazil.
Lantana camara is a native plant in tropical and subtropical
America and is now widely distributed throughout the tropics,
subtropics and warm temperate regions of the world (Broughton,
2000; Ghisalberti, 2000). Dutch explorers introduced it into the
Abstract
In this study we describe the mating behavior of Dorcacerus barbatus (Olivier, 1790) and its predation on
the reproductive parts of Lantana camara L. (Verbenaceae) in a restored area of Atlantic forest, in
southeastern Brazil. Lantana camara is regarded as one of the world’s ten worst weeds. We found three
species of longhorn beetles on this plant: Trachyderes succintus duponti Aurivillius, 1912, Andraegodius
rufipes zonatus (Dalman, 1823), and D. barbatus; this last species represented more than 95% of all
individuals found. This is the first record of these three cerambycid species on L. camara. We observed a
sequence of mating-behavior stages: jousting, antennation, holding, mounting, fighting, licking, abdomen
bending, and copulation. D. barbatus destroyed 60% of the reproductive parts during the mating process.
These primary observations indicate that D. barbatus may be considered as a potential biocontrol agent for
this invasive species.
Keywords: Biological control, invasive species, insect behavior, herbivory, Atlantic Rain Forest.
Netherlands from Brazil in the late 1600s and later explorers
from other countries took seeds to continental Europe, Great
Britain and North America. Following its introduction into
Hawaii as a garden flower, it soon spread to the islands of the
Pacific, Australia and southern Asia (Ghisalberti, 2000). It
rapidly escaped cultivation to become one of the most noxious
weeds of the world (Broughton, 2000). It infests millions of
hectares of grazing and cropping land in 47 countries
(Ghisalberti, 2000), and is regarded as one of the world’s 10
worst weeds (Ghisalberti, 2000; Thomas & Ellison, 2000).
Lantana thrives in a wide variety of environmental conditions
and invades riverbanks, mountain slopes and valleys, pastures,
and commercial forests where it forms impenetrable stands that
obstruct access and utilization. Through allelopathic suppression
of indigenous plant species, Lantana invasions also interrupt the
succession processes (Gentle & Duggin, 1997) and reduce the
biodiversity of natural ecosystems. Many parts of L. camara are
toxic (Morton, 1994) and if consumed can cause cattle
poisoning, with estimated annual stock loss to the farming
community in South Africa of about 1800 heads (Kellerman et
al., 1996). As an example, L. camara was introduced in India
156
from Australia as an ornamental plant and later on invaded the
majority of Indian pasture lands (13.2 million ha), besides other
areas. The cost of its control is US$ 70 per hectare. As 4% of
India’s land area is pasture, the damage from Lantana is
estimated to be US$ 924 million per year (see Pimentel et al.,
2001).
The difficulties and expenses incurred by chemical and
mechanical control measures and the rapid rate of the weed’s
spread fueled the initiation of a strong biological control
program in the early 1960s. However, L. camara biological
control attempts started in 1902, followed by others at intervals
throughout the century (Broughton, 2000; Thomas & Ellison,
2000). These investigations resulted in 36 insect species being
released in various countries (Thomas & Ellison, 2000).
However, effective control has not been achieved yet because
the combined impact remains insufficient to reduce the weed to
acceptable levels and the biocontrol of the weed is considered
negligible (Anonymous, 1999). Most of the established
biocontrol agents are leaf feeders, which directed the search
towards new candidate agents that feed on different parts of the
plant (Winder & Harley 1983, Cilliers & Neser, 1991; Palmer &
Pullen, 1995; Broughton, 1999; Broughton, 2000, Palmer et al.,
2000; Thomas & Ellison, 2000; Baars et al., 2003; Williams &
Madire, 2007) or pathogens (Thomas & Ellison, 2000; Pereira et
al., 2003).
Casual observations on the herbivore insect and their host
plants in the Fazenda Bulcão, municipality of Aimorés, Minas
Gerais state, southeast Brazil, revealed three species of longhorn
beetles associated with L. camara in November 2004:
Trachyderes succintus duponti Aurivillius, 1912; Andraegodius
rufipes zonatus (Dalman, 1823); and Dorcacerus barbatus
(Olivier, 1790). Dorcacerus barbatus represented more than
95% of all individuals found. To our knowledge, this is the first
record of these three cerambycid species on L. camara.
The Fazenda Bulcão has an area of approximately 676 ha in
which a large land restoration program was initiated in 2001.
The Aimorés mountain ridge was originally covered by Atlantic
Forest vegetation of which only 5%-7% remains. Lantana
camara occurs in small patches throughout the property and
neighboring areas. We selected five individuals of this plant for
the observations on the feeding and mating behavior of D.
barbatus. Field behavioral observations were conducted unaided
and with the naked eye, following “ad libitum” (Altmann,
1974). Beetles were observed only during three days, and then
they disappeared. The highest abundance of beetles occurred in
the first day. Mating behavior events were observed from 08:00
to 17:00. Voucher specimens of the cerambycids were deposited
at the Museu de Zoologia da Universidade de São Paulo
(MZUSP), São Paulo, Brazil.
We observed a sequence of stages, namely jousting,
antennation, holding, mounting, fighting, licking, abdomen
bending, and copulation. The Dorcacerus barbatus male
approachs the female, touching her with his antennae or tarsi,
and then hold and mount her while touching her elytra and
pronotum with hihs palpae. Fukaya (2003) has shown that
similar mating attempts by males of Anoplophora malasiaca
(Thomson) (Coleoptera: Cerambycidae) were repeated on a glass
dummy treated with the solvent extract of female elytra,
indicating the presence of a female sex pheromone perceptible
by direct contact. Males mounted the female usually from the
Figure 1 - Dorcacerus barbatus (Cerambycidae) on Lantana camara
(Verbenaceae). (A) Abdomen bending during mating. (B)
Feeding on inflorescences. (C) Feeding on fruits.
Fernandes & Santos
157
rear, yet, at times, male and female may fight. In these
situations, the male lowers his head to stroke the elytra of the
female with his maxillary palpi. This behavior has been argued
to have a calming effect on the female (Crook et al., 2004). Once
mounted on a motionless female, the male curled the tip of his
abdomen, extending his genitalia under the female’s abdomen.
This “abdomen bending” continued until copulation commenced
(Fig. 1A). Similar mating behavior was observed by Crook et al.
2004 for the longhorn beetle Dectes texanus texanus LeConte.
We observed 24 mating pairs and 27 solitary individuals of D.
barbatus on the five individuals of L. camara. In spite of not
being quantified, it is probable that the single
individualsconsisted, in large part, of males waiting to approach
and copulate with females.
Most males of D. barbatus copulated while females fed on
the reproductive parts (flower and/or fruits) of L. camara. Plants
had an average of 121.6 ± 46.17 ( ± SE) reproductive parts. D.
barbatus totally destroyed 60% of them during the mating
process (Tab. 1). However, single individuals could also attack
the reproductive parts (Fig. 1B-C). Overall, most reproductive
parts of L. camara were attacked (attacked reproductive parts:
73.8 ±. 34.45, ± SE, n = 5 individuals; unattacked repro-
ductive parts: 47.8 ±. 14.17, ± SE, n = 5 individuals). Because
of the high impact on reproductive parts, D. barbatus may be
considered as a potential agent for the biological control of L.
camara. On the other hand, detailed studies on its impact on the
lifetime fitness and host specialization on L. camara are needed.
Dorcacerus barbatus may have a specific relationship with L.
camara. Despite the fact that this species was only found
feeding on L. camara, more studies will be needed. However,
other cerambycidae species (Aerenicopsis championi and A.
mendosa), not listed in this study, were also reported on L.
camara (see Monné, 2004). Among the longhorn beetles
observed, only T. succintus duponti (previously recognized as T.
succintus) attacks plantations of Eucalyptus (Berti-Filho, 1997)
and rubber trees, Hevea brasiliensis (Willd. ex Adr. Juss) Müll.
Arg., in Brazil (Dall’oglio & Peres-Filho, 1997). Among the
several plant species available at the study area, D. barbatus
appears to have selected L. camara as mating site, and this
selection of host plant can be of fundamental importance to the
behavior and biology of many cerambycid species. Hence,
detailed studies on the behavior and ecology of D. barbatus are
needed to answer this question.
Acknowledgments
We thank L. R. Viana and two anonymous reviewers for
their encouragement and comments on earlier versions of the
manuscript, M. Monné for comments and papers, U. R. Martins
(MZUSP) for the identification of the Cerambycidae and the
Instituto Terra for providing financial and logistical support. We
also thank CNPq (30.9633/2007-9) and Fapemig (CRA 697/06),
and Planta Tecnologia Ambiental.
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Table 1 - Number and proportion of attacked and unattacked reproductive parts (flowers and/or fruits), and the number of mating and solitary
beetles of Dorcacerus barbatus (Cerambycidae) on Lantana camara (Verbenaceae).
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Fernandes & Santos
... These observations led us to call this series of male behavior appeasement. The so-called antennation behavior [30,31], a sword-battle-like contact of the antennae of male and female, was not observed in this species. Scratching behavior against the female pronotum was observed in one very large male of more than 33 mm body length (Table 1, M3; Table 2). ...
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The biology and host range of the cerambycid beetle Aerenicopsis championi Bates, a potential biological control agent for the weed Lantana camara L., were studied. A. championi is a univoltine species associated with L. camara, L. urticifolia, and L. hirsuta in Mexico and Central America. In Mexico, adult emergence occurred in May and June at the start of the rainy season. Larvae fed within the stems over a 9- to 12-month period and caused damage to the plant. The insect was imported into Australia, where a procedure for rearing it in the laboratory was developed. Host-range tests indicated that adults oviposited and larvae commenced development in L. camara and L. montevidensis but not in any of 57 other species tested. A CLIMEX model indicated that most areas infested with lantana in Australia would have a favorable climate for A. championi. Permission to release this insect in Australia was obtained and three small releases were made in southern Queensland in February 1995.
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Seven major types of sampling for observational studies of social behavior have been found in the literature. These methods differ considerably in their suitability for providing unbiased data of various kinds. Below is a summary of the major recommended uses of each technique: In this paper, I have tried to point out the major strengths and weaknesses of each sampling method. Some methods are intrinsically biased with respect to many variables, others to fewer. In choosing a sampling method the main question is whether the procedure results in a biased sample of the variables under study. A method can produce a biased sample directly, as a result of intrinsic bias with respect to a study variable, or secondarily due to some degree of dependence (correlation) between the study variable and a directly-biased variable. In order to choose a sampling technique, the observer needs to consider carefully the characteristics of behavior and social interactions that are relevant to the study population and the research questions at hand. In most studies one will not have adequate empirical knowledge of the dependencies between relevant variables. Under the circumstances, the observer should avoid intrinsic biases to whatever extent possible, in particular those that direcly affect the variables under study. Finally, it will often be possible to use more than one sampling method in a study. Such samples can be taken successively or, under favorable conditions, even concurrently. For example, we have found it possible to take Instantaneous Samples of the identities and distances of nearest neighbors of a focal individual at five or ten minute intervals during Focal-Animal (behavior) Samples on that individual. Often during Focal-Animal Sampling one can also record All Occurrences of Some Behaviors, for the whole social group, for categories of conspicuous behavior, such as predation, intergroup contact, drinking, and so on. The extent to which concurrent multiple sampling is feasible will depend very much on the behavior categories and rate of occurrence, the observational conditions, etc. Where feasible, such multiple sampling can greatly aid in the efficient use of research time.
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The ornamental hybrid shrub, Lantana camara L. (lantana), is a serious environmental weed and has been targeted for biological control in South Africa since 1961. The established biocontrol agents cause insufficient levels of damage and additional natural enemies are required to reduce the invasiveness of this weed. The lantana mirid, Falconia intermedia (Distant), is a promising new agent that was imported from the Caribbean for life history and host-range studies. The nymphs and adults are leaf-suckers that cause chlorotic speckling, which reduces the photosynthetic capacity of the plant. Biological studies indicate that F. intermedia has considerable biocontrol potential, in that it has a high intrinsic rate of increase, the potential for multiple generations a year, highly mobile adults, and a high level of damage per individual. Host-specificity trials indicated that the lantana mirid has a narrow host range, with L. camara being the most suitable host, but several indigenous African species in the closely related genus Lippia are suitable alternative host plants. Under multiple-choice conditions, adults showed a significant and strong oviposition preference for L. camara over the Lippia species. A risk assessment of potential nontarget effects indicated that three Lippia species could sustain damage levels in the field. The relatively low probability of damage to indigenous species was considered a justified trade-off for the potentially marked impact on L. camara. The regulatory authorities accepted the results of this study and F. intermedia was released against L. camara in South Africa in April 1999.
Article
The seed-fly Ophiomyia lantanae (Froggatt) (Dip.: Agromyzidae) is one of the most widely established biological control agents for lantana. Larvae of O. lantanae are found in the lantana fruit, in which they attack the fleshy endosperm and pupate in either the seed receptacle or a chamber within the fruit. This paper questions the impact of the seed-fly as a biological control agent in Australia. Fruit were dissected and tested with tetrazolium chloride to assess the viability of embryos. Of 3300 fruit dissected, none of the embryos were affected. Dissections and testing of fruit of different ages suggest that embryos are viable for a short period, regardless of whether the fruit is fly-infested.