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Tropical Zoology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ttzo20 Galling insects on neotropical species of Baccharis (Asteraceae) G. W. Fernandes a , M. A.A. Carneiro a , A. C.F. Lara a , L. R. Allain a , G. I. Andrade a , G. R. Julião a , T. R. Reis a & I. M. Silva a a Ecologia Evolutiva de Herbivoros Tropicais/DBG, CP 486, ICB/Universidade Federal de Minas Gerais, 30161-970, Belo Horizonte, MG, Brazil Published online: 01 Aug 2012.
To cite this article: G. W. Fernandes , M. A.A. Carneiro , A. C.F. Lara , L. R. Allain , G. I. Andrade , G. R. Julio , T. R. Reis & I. M. Silva (1996) Galling insects on neotropical species of Baccharis (Asteraceae), Tropical Zoology, 9:2, 315-332, DOI: 10.1080/03946975.1996.10539315 To link to this article: http://dx.doi.org/10.1080/03946975.1996.10539315
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Galling insects on neotropical species of Baccharis (Asteraceae)
G.W. FERNANDES, M.A.A. CARNEIRO, A.C.F. LARA, L.R. ALLAIN, G.I. ANDRADE, G.R. JuuA.o, T.R. REis and I.M. SILVA Ecologia Evolutiva de Herbivoros Tropicais/DBG, CP 486, ICB/Universidade Federal de Minas Gerais, 30161-970 Belo Horizonte, MG, Brazil
Received 29 April 1994, accepted 3 October 1995
We report on the insect galls on neotropical species of Baccharis (Asteraceae) and describe new kinds oi galls on Baccharis spp. of the cerrado and rupestrian fields of Brazil. Gall collections were made in several localities in Minas Gerais (Belo Hori zonte, Caratinga; Lagoa Santa, Moeda, Ouro Preto, and Serra do Cip6). Baccharis may support the richest galling fauna of the neotropics (121 galling species on only 40 species of Baccharis). Nevertheless, gallers were not evenly distributed across host plant species. Four species of Baccharis alone supported 46% of the galling fauna. The most diverse fauna occurred on B. dracunculifolia (17 galls), B. concinna (15 galls), B. salicifolia (13 galls) and Baccharis sp. 1 (11 galls). The most numerous galling taxa were, in decreasing order, Cecidomyiidae (Diptera), Lepidoptera (several Gelechiidae), Psyl lidae (Homoptera), and Tephritidae (Diptera). The most diverse gall communities were on Baccharil species that inhabit harsh environments, such as B. concinna in the rupestrian fields of Minas Gerais, B. salicifolia in the dry chaco vegetation of Argentina, and B dracunculi/olia and Baccharis sp. 1 of the cerrado of Lagoa Santa in Minas Gerais.
KEY WORDS: BacchariJ. insect community, insect galls, neotropics, species richness. Downloaded by [University of Sydney] at 13:01 26 August 2013
Introduction 315 Materials and method~ 316 Results and discussion 317 Acknowledgments 330 References 330
INTRODUCTION
Gall-forming insects are very common in all biogeographical regions of the world, except the poles (see MA~I 1964, GAGNE 1989, FERNANDES et al. 1994). They achieve the highest diversity in warm temperate ecosystems; i.e., between 25° and 40° north or south (PRICE et al. 1996). In the Neotropical region, they are most species-rich in 316 G. W. Fernandes et alii
the rupestrian fields and in the cerrado (savanna) vegetation of southeastern Brazil (FERNANDES & PRICE 1988, LARA & FERNANDES 1996a). The mechanisms that in fluence this global distributional pattern are still under investigation, but it appears that plant sclerophylly and its related emergent properties have influenced the evolution of galling insects (see FERNANDES & PRICE 1991, 1992; LARA & FERNANDES 1996b; PRICE et al. 1996). Other studies have addressed other hypotheses on the evolution of the galling habit (e.g., CoRNELL 1983, WARING & PRICE 1990, HARTLEY & LAWTON 1992, BLANCHE 1994, see review by PRICE et al. 1987). Studies on insect galls are now very common in ecology. Galls are often conspicuous and easy to find all year round, which limits methodological problems of seasonality and turnover rates characteristic of other insect herbivores (see LARA & FERNANDES 1996b). The majority of galling insects are host-plant and plant-organ specific, and gall morphology is specific to each inducer (FLOATE et al. 1996) making them excellent systems for the estimation of the patterns of abundance and richness (FERNANDES & PRICE 1988, FERNANDES et al. 1994). In addition. galls may be seen as extended phenotypes of their inducers (WEis et al. 1988). Several ecological hypoth eses have been generated from studies of gall communities on many temperate plants [e.g., Salix spp. (PRicE & RmNINEN 1993), Quercus (AsKEW 1980, CoRNELL 1983), Atriplex (HAWKINS 1984), Chrysothamnus (FERNANDES 1992a), and Solidago (GAGNE 1989). Tropical insect galling communities are, however, poorly known and mostly undescribed. We do not know of any community of tropical insect galls that has been studied in as much detail as the ones mentioned above. Neotropical studies on galling insects have been primarily descriptive (e.g., FERNANDES et al. 1988, GAGNE 1994 and references therein), or centred on the natural history of single species (e.g., FER NANDES et al. 1987, FERREIRA et al. 1990). We believe that a better understanding of gall distribution, gall ecology, and insect plant relationships would be achieved if diverse tropical communities were studied in detail over the long-term to enable appropriate comparisons with corresponding temperate systems. For this reason we have chosen to study the highly diverse insect gall community on the neotropical species of Baccharis Linnaeus (Asteraceae). In this study we present a list, partially extracted from the literature, of the insect galls on neotropical species of Baccharis and report for the first time new types of insect galls found on several species of Baccharis of the cerrado and rupestrian fields of Brazil. Downloaded by [University of Sydney] at 13:01 26 August 2013
MATERIALS AND METHODS
This paper is a combination of data extracted from the literature and data obtained by us during a 4 year study. Insect galls on B. concinna Barroso, B. dracunculifolia D.C., and on Baccharis sp. 2 to sp. 9 were collected from 1990 to 1993 by our research group while performing several studies on these plants. On B. concinna and B. dracunculifolia galls were counted on 200 plants twice a year from 1991 to 1993. The other species were surveyed once a year from 1990 to 1993. Surveys were performed in several localities, but primarily in the cerrado and rupestrian fields of the State of Minas Gerais (Belo Horizonte, Lagoa Santa, Maeda, Ouro Preto, and Serra do Cip6), and in the Atlantic rain forest (Caratinga) in southeastern Brazil. Baccharis sp. 2 was sampled in Belo Horizonte, while Baccharis Galling insects on Baccharis 317
sp. 3 to Baccharis sp. 9 were sampled from Lagoa Santa to Serra do Cip6 (between 19°10' and 19°40'S. and 43°30' and 43°55'W). Gall collection methods are reported in FERNANDES et al. (1988). Complementary bibliographical surveys utilized several compendiums and lists of galls on neotropical species of Baccharis as sources of additional information. These are listed in the reference column of Table 1 and additional field site description can be found in these references. Insect galls vary enormously in form, host organ, colour, number of internal larval chambers, and epidermal cover (MANI 1964), and we utilized these variable characters to describe them. Because of the many problems with the taxonomy of tropical insects, we identify most gallers only to family level. Whenever possible, we give the genus and species names. Insects are continuously being reared from galls and these will be identified to the species level in the near future. Some Baccharis were listed as sp. 1., sp. 2, etc. to sp. 9 due to taxonomical problems. The identification of Baccharis at the species level is generally complex due to the large number of species in the genus, approximately 500 (BoLDT 1989). Ninety percent of these species occur in South America (NESOl\1 1990), and more than 120 species are known to occur in southeastern Brasil alone (BARROSO 1976). In addition, hybridization is common in the genus (e.g., FAIN! et al. 1991) making identification complicated. Nevertheless, all plants are being identified by specialists and will be reported in the near future as studies develop.
RESULTS AND DISCUSSION
Baccharis may have the richest galling fauna of the neotropics. One hundred and twenty one species of galling insects were found on 40 species of Baccharis (Table 1). Gallers were not evenly distributed across host plant species. Four species of Baccharis (10% of the species studied) supported 46% of the galling fauna (56 galling species) (Table 1). The host plants with the most diversity were B. dracunculifolia (17 gall species), B. concinna (15 gall species), B. salicifolia (13 gall species) and Baccharis sp. 1 (11 gall species). We also found a gall induced by an unidentified species of fungus on B. concinna, while other authors reported unidentified insect galls on B. nitida, and B.
Downloaded by [University of Sydney] at 13:01 26 August 2013 poeppigiana (these galls were not used in the analysis). The diversity of galls on Baccharis spp. is certainly much higher as there are almost 500 Baccharis species in the neotropics (BARROSO 19 76, HELL WIG 1990) and practically no comprehensive attempt has been made to survey their galling insects. The data we present, beg the question of whether, despite having rich gall communities, gall abundance is equal across the species and the habitats occupied by them. Future studies will focus on comparisons of gall richness and abundance between B. concinna and B. dracunculifolia in Brazil. The most numerous galling taxa belonged to the cecidomyiids (Diptera Cecido myiidae), Lepidoptera (several Gelechiidae), Psyllidae (Homoptera), and Tephritidae (Diptera) (Fig. 5). The common occurrence of cecidomyiid galls on Baccharis has already been reported (see FERNANDES et al. 1994, GAGNE 1994). This is the first time that broad patterns in gall distribution are examined within a single host plant taxon (genus) in the tropics (but see DocTERS VAN LEUUWEN REIJNVAAN & DocnRs VAN LEEUWEN 1926, FERNANDES 1992b). Our findings add to and corroborate the latest findings on the ecology of galling insects (see FERNANDES & Downloaded by [University of Sydney] at 13:01 26 August 2013
v.> Table 1...... 00 Insect galls on neotropical Baccharis spp. (Asteraceae).
Gall description Gall Host plant Gall-forming taxa Reference number Organ Shape Colour Pubescence Chambers
B. aphylla (Veil.) DC 1 Psyllidae leaf spherical green glabrous one TAVARES (1917), HouARD (1933) B. artemisioides Hook 1 Neolasioptera heteri- stem elliptical green glabrous one KIEFFER & & Arn thalami' Kieffer & JoRGENSEN (1910), Jorgensen GAGNE (1994) B. bogotensis H.B. & 1 Calopedilla herbsti 1 stem elliptical green - one KIEFFER (1908), K. Kieffer HavARD (1933) 2 Muscidae stem elliptical green - one KIEFFER (1908), HavARD (1933) B. boyacensis Cuatrec. 1 Trioza sp.' apical bud rosette green hairy one AREVALO & ToRRES (1987) B. capitalensis Hering 1 Rhopalomyia sp. ' apical bud rosette green glabrous one GAGNE (1994) B. concinna Barroso 1 C urculionidae stem elliptical brown glabrous one Fig. 1a 2 Lepidoptera stem elliptical green/red glabrous one Fig. 1b 2 Lepidoptera stem elliptical green/red glabrous one Fig. 1b 3 Cecidomyiidae apical bud elliptical green glabrous one Fig. lc 4 Cecidomyiidae stem swelling green glabrous one Fig. 1d 5 Psyllidae leaf legume green glabrous one Fig. 1e 6 Cecidomyiidae apical bud globulous green glabrous various Fig. 1f 7 Cecidomyiidae apical bud rosette red glabrous various Fig. 1g R Lt>rick•rtt>ra stt>m cla\'ifNm grt>t>n glabrnus nne Fig. lh 9 Cecidomyiidae apical bud cylindric brown glabrous various Fig. 1i CJ 10 rung! stem globulous brown glabrous vanous hg. lj ~ 11 Cecidomyiidae stem elliptical green glabrous one Fig. 1k 12 Cecidomyiidae stem globulous green glabrous one Fig. 11 '"I1 n..... 13 Cecidomyiidae flower* swelling yellow glabrous one Fig. 1m ::l ~ 14 Cecidomyiidae leaf discoid yellow glabrous one Fig. ln ::l 15 Coleoptera root elliptical brown glabrous one - 0..n B. confertifolia Colla elliptical green glabrous - KIEFFER & 1 Trioza baccharis leaf "'n Kieffer & Herbst HERBST (1911) ...... (continued) S: Downloaded by [University of Sydney] at 13:01 26 August 2013
Table 1 (continued) CJ ~ Gall description Gall s· Host plant Gall-forming taxa Reference (]"Q number Organ Shape Colour Pubescence Chambers s· "'0 B. coridifolia DC 1 Lasioptera cordoben- apical bud elliptical green hairy one KIEFFER & ...n sis 1 Kieffer & Jorgen- jORGENSEN (1910) "'0 sen ::I 2 Raccharnrn1•ia cnrdn- apical hncl Table 1 (continued) N ""0 Gall description Gall Host plant Gall-forming taxa Reference number Organ Shape Colour Pubescence Chambers B. ef/usa Griseb 1 Percnoptera angusti- - spherical whitish - one KIEFFER & pennis' Philippi JoRGENSEN (1910) B. elaegnoides Steudel 1 Cecidomyiidae apical bud rosette green hairy one TAVARES (1917), HOUARD (1933) B. eupatorioides Hook 1 Scheueria agglomera- apical bud flower like green hairy one KIEFFER & HERBST &Arn ta 1 Kieffer (1909), GAGNE (1994) 2 Scheueria longicornis 1 apical bud rosette green glabrous one KIEFFER & HERBST Kieffer & Herbst (1909), GAGNE (1994) 3 Tephritidae stem globulous white glabrous one KIEFFER & HERBST (1909) B. genistelloides Baker 1 Baccharomyia ramo- leaf elliptical green glabrous various TAVARES sina 1 Tavares (1917, 1922) B. latifolia (R & P) 1 Cecidomyiidae stem globulous green glabrous various AREVALO & Pers. ToRRES (1987) 2 Cecidomyiidae leaf conical reddish glabrous one AREVALO & ToRRES (1987) 3 Orsotricha near veno- stem claviform green glabrous one AREVALO & sa' TORRES (1987) 4 Trioza gibbosa' Craw- flowers flower reddish scales various AREVALO & ford ToRRES (1987) 5 Trioza sp. 2 flowers stellate reddish scales various AREVALO & TORRES (1987) B. linearis (R & Pl I Rhachiptera limbata ' apical bud globulous white hairv one ArlARO f"t al (19~4) Pers. () B 'II.W .. I!:t!.'.- II.B.K. 1 Onutricb.- nc .. u L cnu ;l<.:lll dlipti~al grccnioh glabruuo l.H1C ARL\ALU & ~ sa' ToRREs (1987) 2 Lepidoptera leaf globulous green glabrous - AREVALO & 'Tl ~.... ToRRES (1987) ::I I>) B. microphylla 1 Cecidomyiidae apical bud globulous - - - HIERONYMUS (1884), ::I H.B.K. HouARD (1933) 0.. ~ B. myrsinites (Pers.) 1 Neolasioptera rostra- flower elliptical yellowish glabrous one GAGNE (1994) Cl> ta 1 Gagne ....~ (continued) S: Downloaded by [University of Sydney] at 13:01 26 August 2013 GJ Table 1 (continued) s·~ Gall description (JQ Gall Host plant Gall-forming taxa Reference ..... number ::s Organ Shape Colour Pubescence Chambers "'n ...n B. nitida (R & P) 1 Orsotricha near vena- stem globulous green glabrous one AREVALO & "'0 Pers. sa• ToRRES (1987) ::s green AREVALO & t:l; 2 Insecta leaf glohulous glabrous - 1::. <"> TORRES (1987) <"> ~ B. pauczdentata DC 1 Coleoptera stem globulou~ brown glabruus one lluLARIJ (1933) I::> 2 Cecidomyiidae stem globulous greenish glabrous one HouARD (1933) a. B. poeppigiana DC 1 Insecta stem elliptical - glabrous one KIEFFER & HERBST (1905) B. prunifolia H.B.K. 1 Orsotricha near vena- stem elliptical green glabrous one AREVALO & sa• ToRREs (1987) B. rosmarinifolia 1 Dasineura chilensis 1 apical bud rosette green glabrous one KIEFFER & HERBST Hook & Arn Kieffer & Herbst (1909), GAGNE (1994) 2 Dasineura subnervis 1 apical bud rosette green glabrous one KIEFFER & HERBST Kieffer & Herbst (1909), GAGNE (1994) 3 Calopedilla herbsti 1 apical bud elliptical brown glabrous one KIEFFER (1903), Kieffer GAGNE (1994) 4 Percnoptera angusti- apical bud spheroid green glabrous one HouARD (1933) penn is' Phillipi 5 Dasineura sp. ' stem elliptical - glabrous one KIEFFER & HERBST (1911), HowARD (1933) 6 Cecidomyiidae elliptical green glabrous one KIEFFER & HERBST (1909) 7 Homoptera B. salicifolia Pers 1 Cecidotrio:r.a mendon- flower - - - - KIEFFER & cina' Kieffer & Jar- JoRGENSEN (1910) gensen 2 Aciura baccharidis' stem elliptical green glabrous one KIEFFER & Kieffer & Jorgensen JoRGENSEN (1910) (continued) v. ....N Downloaded by [University of Sydney] at 13:01 26 August 2013 ..... N Table 1 (continued) N Gall description Gall Host plant Gall-forming taxa number Reference Organ Shape Colour Pubescence Chambers 3 Aciura fa/cigera ' stem elliptical green glabrous one KIEFFER & Kieffer & Jorgensen JoRGENSEN (1910) 4 Meunierie/la ornati- stem elliptical green glabrous various KIEFFER & cornis 1 Kieffer & Jor- JoRGENSEN (1910) gensen 5 Trypeta cucu/i ' stem spherical whitish - one KIEFFER & Kieffer & Jorgensen JoRGENSEN (1910) 6 Asphondylia hierony- stem spherical green - one HIERONYMUS (1884), mi 1 Hieronymus GAGNE (1994) 7 Rhoasphondylia eras- stem spherical green hairy one KIEFFER & sipalpis 1 Kieffer & JoRGENSEN (1910), Jorgensen GAGNE (1994) 8 Misospatha globifex 1 flower spherical green glabrous various KIEFFER & Kieffer JoRGENSEN (1910) 9 T rioza alacris ' Flor leaf elliptical green glabrous one KIEFFER & JoRGENSEN (1910) 10 Lepidoptera leaf concave green glabrous one KIEFFER & JoRGENSEN (1910) 11 Cecidomyiidae leaf discoid green glabrous one TAVARES (1915) 12 Baccharomyia ornati- apical bud elliptical green glabrous two TAVARES (1915) cornis 1 Kieffer & Jor- gensen CJ 13 Rhopalomvia t;,lnhi- apical bud flower like green hairy one KIEFFER & fex 1 Kieffer & Jor- JoRGENSEN (1910), ~ gensen GAGNE (1994) 'TJ B. schultzii Baker 1 Lepidoptera stem clavi form green glabrous one TAVARES (1917) (1).... ::I 2 Rhoasphondylia /ri- leaf globulous green glabrous one TAVARES (1917), I)) ::I burguensis ' Tavares GAGNE (1994), 0. ARAUJO et al. (1995) (1) "'(1) (continued) ... S: Downloaded by [University of Sydney] at 13:01 26 August 2013 C) Table 1 (continued) ~ Gall description s· Gall (JQ Host plant Gall-forming taxa Reference .... number Organ Shape Colour Pubescence Chambers ~ I'D n"' B. serru/ata Person 1 Tecia mendozetia• stem elliptical green glabrous one STRAND (1910), ... Strand HouARD (1933) "'0 2 Cecidomyiidae stem elliptical green glabrous one K1E.FFER & ~ ~ jORGENSEN (1910) ...,s::. B. spartzozdes C. Gay 1 Neo/aszoptera rostra- flower elliptical yellowish glabrous one GAGNE (1994) ta' Gagne ~ 2 Neo/asioptera heteri- stem elliptical green glabrous one KIEFFER & il thalami' Kieffer & jORGENSEN (1910), Jorgensen GAGNE (1994) B. subu/ata D. Don 1 Baccharomyia inter- apical bud elliptical green glabrous two TAVARES (1915), rupta' Kieffer & Jor- GAGNE (1994) gensen 2 Fapua a/bineroe//a • stem elliptical green - one STRAND (1910), Strand HouARD (1933) B. tricuneata Gardn. 1 Orsotricha near veno- stem elliptical green/red glabrous one AREVALO & sa• TORRES (1987) B. trimera DC 1 Baccharomyia ramo· stem elliptical green glabrous several TAVARES (1917), sina ' Tavares GAGNE (1994) B. trinerois Persoon 1 Gera/desia cumbren- leaf blister - glabrous one GAGNE (1994) sis' Mohn 2 Rhoasphondy/ia fri- stem globulous green glabrous one TAVARES (1917) burguensis' Tavares GAGNE (1994), ARAU:Jo et al. (1994) Baccharis sp. 1 1 Arphondy/ia bac- flower cylindric black hairy one K1E.FFER & HERBST charis' Kieffer & (1905), GAGNE (1994) Herbst Baccharis sp. 2 1 Cecidomyiidae stem elliptical brown glabrous one Fig. 3a 2 Lepidoptera apical bud elliptical green glabrous one Fig. 3b 3 Cecidomyiidae leaf discoid yelow glabrous one Fig. 3c 4 Cecidomyiidae apical bud globulous red glabrous one Fig. 3d 5 Cecidomyiidae stem elliptical brown glabrous one Fig. 3e 6 Cecidomyiidae apical bud elliptical brown glabrous one Fig. 3£ \.» IV (continued) \.» Downloaded by [University of Sydney] at 13:01 26 August 2013 \,;> tv ""'" Table 1 (continued) Gall description Gall Host plant Gall-forming taxa Reference number Organ Shape Colour Pubescence Chambers 7 Tephritidae flower rosette yellow glabrous one Fig. 3g 8 Psyllidae leaf legume green glabrous one Fig. 3h 9 Cecidomyiidae lateral bud conical yellow glabrous one Fig. 3i 10 Cecidomyiidae flower elliptical yellow glabrous one Fig. 3j 11 Cecidomyiidae leaf discoid green glabrous one Fig. 3k Baccharis sp. 3 1 Cecidomyyidae leaf discoid brownish glabrous one Fig. 4a 2 Cecidomyiidae stem globulous brownish glabrous one Fig. 4b Baccharis sp. 4 1 Cecidomyiidae leaf discoid brownish glabrous one Fig. 4c 2 Cecidomyiidae stem globulous brownish glabrous one Fig. 4d Baccharis p/atypoda 1 Cecidomyiidae buds swelling green glabrous several Fig. 4e DC 2 Cecidomyiidae leaf discoid yellowish glabrous one Fig. 4f Baccharis sp. 5 1 Cecidomyiidae stem elliptical brownish glabrous one Fig. 4g 2 Cecidomyiidae apical bud tortuous brownish glabrous one Fig. 4h Baccharis sp. 6 1 Cecidomyiidae leaf discoid green glabrous one Fig. 4i Baccharis sp. 7 1 CedJumyiiJa~ >lt:m glubuluus brownish glabrous one Fig. 4j Baccharis sp. 8 1 Tephritidae flower elliptical yellowish glabrous one Fig. 4k C) * Male flowers only, **both male and female flower, 1 = Cecidomyiidae, 2 = Psyllidae, '= Tephritidae, • = Gelechiidae. ~ ~... ::3 ~ ...n"' S: Galling insects on Baccharis 325 Downloaded by [University of Sydney] at 13:01 26 August 2013 Fig. 1. -Galling insects on Baccharis concinna from Serra do Cip6, Minas Gerais (see Table 1). 326 G. W. Fernandes et alii \!)A ug Downloaded by [University of Sydney] at 13:01 26 August 2013 Fig. 2. -Galling insects on Baccharis dracunculifolia from Belo Horizonte, Caratinga, and Serra do Cip6, Minas Gerais (see Table 1). Galling insects on Baccharis 327 c g Downloaded by [University of Sydney] at 13:01 26 August 2013 Fig. 3. -Galling insects on Baccharis sp. 2. from Belo Horizonte, Minas Gerais (see Table 1). 328 G. W. Fernandes et alii a c ':'.·.-·~ \ 0 I!: .' \0 '\ f ~ ·. .~·"' '-~,-·~ . ·.. _;,..-- . -~:... . 0 I Downloaded by [University of Sydney] at 13:01 26 August 2013 k Fig. 4. -Galling insects on Baccharis spp. (Baccharis sp. 3 to Baccharis sp. 8, and B. p/atypoda) from Serra do Cip6, Minas Gerais (see Table 1). Galling insects on Baccharis 329 (/) 80 80 ILl 0 CJ Number ILl 0. ~ Percentage (/) ~ 60 60 -0 (!)z -UJ :J (!) ..J ~ <[ 40 40 z (!) UJ 0 0"- 0: UJ 0: 20 20 0. UJ CD :E ~ z 0 0 E ...... c .. c ..,c ..c :!! :5! a ;g iC :5! &... ·;;. ..,0 1: 0:: 0 E ~ D. .2 0 ·a. a. "S :5! .. ~ ~ u ...J :::0 .. (.) (.) Fig. 5. - Proportion of galling insect taxa on neotropical species of Baccharis (Asteraceae). PRICE 1988, 1991; FERNANDES et al. 1994) and add to them. A few important observations are of central interest: first, the most diverse gall communities were on Bacchris species that inhabit harsh environments, such as B. concinna in the rupes trian, high altitudinal fields of Minas Gerais, Brazil, B. salicifolia in the dry and low nutrient chaco vegetation of Argentina (see HuECK 1972), and B. dracunculifolia and Baccharis sp. 1 of the dry and low nutrient cerrado of Lagoa Santa in Minas Gerais. Although many galling species on B. dracunculifolia were found in Argentina and Chile, all of them were also found in the dry cerrado vegetation of Minas Gerais. In these localities, habitat harshness may be produced primarily by nutrient stress as rainfall is generally high (see HARIDASAN 1982). This pattern fits the harsh environ Downloaded by [University of Sydney] at 13:01 26 August 2013 ment hypothesis on gall diversity as proposed by FERNANDES & PRicE (1988, 1991) and PRICE et al. (1996). Second, host plant area (distribution over space) may not affect gall diversity as approximately equal numbers of galls may be found on widely distributed as well as restricted host species. This is in agreement with the findings of FERNANDES & PRicE (1988) for temperate species. Nevertheless, more detailed studies should be performed to confirm this finding. Descriptive knowledge of the diverse community of galling insects on Baccharis spp. is important and is the first step for the development of detailed studies. Baccharis spp. are distributed in tropical and temperate regions of the New World, and thus are appropriate for broad comparisons on the evolutionary ecology of galling insects. The evolutionary questions we are interested in were only briefly mentioned here. Future studies will focus on the diversity patterns and mechanisms that influence gall diversity on species of Baccharis of the cerrado and rupestrian fields of Brazil. 330 G.W. Fernandes et alii ACKNOWLEDGMENTS We thank R.J. Gagne, P.W. Price, and an anonymous reviewer for their comments on the manuscript. We also thank G. Barroso, J. Lombardi and R. Gagne for the identification of several plant and insect species, the Funda~iio Biodiversitas and M.M. Duarte for the illustrations, the Caratinga Biological Station, and IBAMA (Parque Nacional da Serra do Cip6-MG) for logistical support. This work was supported by CNPq grants (52.0970/93-4, 52.1772/95-8), FAPEMIG (821/90, 078/91), PRPqfUFMG, Mestrado em Ecologia, Conserva~iio e Manejo de Vida Silvestre, and IFS (H/2487-1). REFERENCES ALJARO M.E., FRIAS D. & MoNTENEGRO G. 1984. Life cycle of Rhachiptera limbata (Diptera, Tephritidae) and its relationship with Baccharis linearis (Compositae). 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