Chrysomelinae species (Coleoptera, Chrysomelidae) and new biological data from Rio de Janeiro, Brazil.

Chrysomelinae is one of the largest subfamilies in Chrysomelidae, yet much basic information remains unknown for Neotropical species. The present study aims to compile the first regional list of Chrysomelinae for the State of Rio de Janeiro, Brazil, and assemble natural history traits obtained from our fieldwork from 2005 to 2010 in Serra dos Órgãos National Park, a mountainous area of Atlantic forest. The species list was compiled from data from field work, collections, and literature, and recorded a total of 100 species, belonging to 21 genera in one tribe (Chrysomelini) and three subtribes: Chrysolinina (91 species), Chrysomelina (eight species) and Entomoscelina (one species). Of these, 91 species are new records for the state. Serra dos Órgaõs National Park holds records of 43 species, with Platyphora being the most species-rich genus, and Solanaceae the most common host plant family. Some new records of reproductive mode (larviparous vs. oviparous) and larval behavior are also given. These Brazil Chrysomelinae species exhibited a clear seasonal pattern, with more species recorded in the hot and rainy season from October to January, and considerably fewer species from June to August, during the drier and colder months. The fraction of new records in comparison with published species and natural history information illustrates how little we know of Chrysomelinae in the state and in the country.

Introduction

is the fifth largest subfamily of , after , , and (Reid 2006), with 3,000 species and 132 genera (Daccordi 1994, 1996, Riley et al. 2002), but these numbers vary among authors (see Seeno and Wilcox 1982, Reid 1995). Two tribes are generally recognized: , which is monogeneric with Latreille (ca. 100 species); and containing the remainder (Seeno and Wilcox 1982, Daccordi 1994). However, there are still many problems concerning taxonomy. Daccordi (1996) listed 38 genera for the Neotropical region, out of which 31 are exclusive to the area. Some of the main contributions for Neotropical taxonomy and cataloguing are those by Jan Bechyně (e.g. 1954, 1958, 1980), which include many species descriptions and some regional lists, and, more recently, a key to the genera in Costa Rica by Wills Flowers (2004), modified from Bechyně and Springlova de Bechyně (1965). Both larvae and adults normally feed on leaves of the same host plant species and species tend to be monophagous or to feed on a narrow group of related plant species (Jolivet 1988). The same author pointed out that host plants are known for nearly 40% of genera, and data are largely lacking for tropical species. In the Neotropical area, are frequently associated with , , , and (Jolivet and Hawkeswood 1995). However, knowledge of biology is rare in this region, remaining so two decades after being underlined by Jolivet (1997).

Except for some ecological studies and species records confined to entomological collections, no list of species exists for the subfamily in the State of Rio de Janeiro or even in Brazil. Since we have conducted extensive research in a protected area in the state and have accumulated considerable biological information on species, our aim here is to compile the first regional list of in Brazil, and to assemble natural history traits for the species found in Serra dos Órgãos National Park, State of Rio de Janeiro.

Materials and methods

For species list compilation for Brazil, four national collections were examined: Coleção entomológica do Laboratório de Ecologia de Insetos / Universidade Federal do Rio de Janeiro, Rio de Janeiro (); Museu Nacional / Universidade Federal do Rio de Janeiro, Rio de Janeiro (), except species from Itatiaia; Coleção entomológica da Fundação Instituto Oswaldo Cruz, Rio de Janeiro (); and Museu Paraense Emílio Goeldi, Belém (). The digital collection of the Museo del Instituto de Zoologia Agricola, Universidad Central de Venezuela (MIZA) was also consulted. Finally, the literature was searched for additional records. These records are indicated in Table 1. Location is given by the municipality within the State of Rio de Janeiro, which comprises 43,696 km² and represents less than 1% of the country’s area. Taxonomy follows Daccordi (1994) and Seeno and Wilcox (1982).

Table 1.

List of species. species from the State of Rio de Janeiro, indicating the municipality of the record and specific location, when available. Numbers indicate the source of information (see footnote below table). SONP = Serra dos Órgãos National Park; INP = Itatiaia National Park.

Species	Location
Chrysomelini: Chrysolinina – 15 genera and 91 species
Calligrapha polyspila (Germar, 1821) (Fig. 1A)	Angra dos Reis3, Itatiaia3, Paraty (Pedra Branca)1, Resende3, Teresópolis3 (SONP1)
Cosmogramma decora Stål, 1859	Itatiaia (INP)1
Cosmogramma fulvocincta Stål, 1859	Itatiaia3
Cosmogramma wygodzinskyi Bechyně, 1948	Itatiaia4
Cryptostetha hieroglyphica Lucas, 1857	Itatiaia3 (INP1)
Cryptostetha notatifrons Stål, 1863	Itatiaia3
Deuterocampta achardi Bechyně, 1944	Mendes4
Deuterocampta cruxnigra Stål, 1859	Angra dos Reis3
Deuterocampta fallax Bechyně, 1950	Itaboraí2, Rio de Janeiro (Gávea4, Tijuca2)
Deuterocampta humeralis Bechyně, 1944	Petrópolis (SONP)3
Deuterocampta leucomelaena (Perty, 1832)	Itatiaia3,4 (INP1)
Deuterocampta pustulicollis Stål, 1859	Macaé2,4
Deuterocampta sedula Stål, 1859 (Fig. 1B)	Teresópolis2 (SONP1)
Deuterocampta semistriata (Fabricius, 1775)	Petrópolis (SONP)4, Rio de Janeiro (Rio de Janeiro4, Corcovado3)
Deuterocampta stauroptera (Wiedmann, 1821)	Rio de Janeiro (Botafogo4, Corcovado3, Gávea4, Rio de Janeiro3, Tijuca4)
Deuterocampta undulata Bechyně, 1950	Rio de Janeiro4
Deuterocampta vittulosa Bechyně,1944	Rio de Janeiro (Engenho de Dentro4)
Dorysterna cruentata (Baly, 1858)	Cambuci (Funil)3
Dorysterna dorsosignata (Stål, 1857)	Itatiaia (INP)1, Rio de Janeiro (Corcovado2, Rio de Janeiro2)
Dorysterna riopardensis Bechyně, 1948	Nova Friburgo2
Dorysterna salvatori Bechyně, 1948	Teresópolis (SONP)1
Elytrosphaera breviuscula Stål, 1858	Grande Rio (Baixada fluminense4)
Elytrosphaera lahtivirtai Bechyně, 1951	Itatiaia (INP1)
Elytrosphaera noverca Stål, 1858	Teresópolis (SONP)1
Elytrosphaera xanthopyga Stål, 1858 (Fig. 1C)	Itatiaia1,3, Resende3, Teresópolis2,3 (SONP1)
Eugonycha bryanti Bechyně, 1946	Rio de Janeiro4
Gavirga subaenea Bechyně, 1946	Itatiaia4
Grammodesma elongata Bechyně, 1952	Itatiaia (INP)8
Grammodesma luridipennis (Baly, 1859)	Itatiaia (INP)8
Grammodesma obliqua (Stål, 1859)	Itatiaia3,4 (PNI)8
Grammodesma rubroaenea (Stål, 1859) (Fig. 1D)	Teresópolis (SONP)1
Grammodesma stulta (Stål, 1859)	Rio de Janeiro (Corcovado2, Rio de Janeiro4, Tijuca2)
Metastyla insignis Achard, 1923	Rio de Janeiro (Corcovado2,4, Rio de Janeiro3, Tijuca2)
Monocampta crucigera (Sahlberg, 1823)	Angra dos Reis3, Itatiaia (Itatiaia3, Penedo3), Rio de Janeiro (Alto da Boa Vista3, Corcovado3, Tijuca3,4), Teresópolis2 (SONP1)
Platyphora acuminata (Olivier, 1790)	Itatiaia3
Platyphora angulata Stål, 1858	Rio de Janeiro5
Platyphora axillaris (Germar, 1824) (Fig. 1E)	Angra dos Reis3, Itatiaia3, Nova Friburgo3, Rio de Janeiro (Gávea3, Tijuca3), Silva Jardim1, Teresópolis2,3 (SONP1,10), Guapimirim (SONP)1, Três Rios3
Platyphora biforis (Germar, 1824)	Itatiaia3, Laje do Muriaé3, Rio de Janeiro2
Platyphora bigata (Germar, 1824) (Fig. 1F)	Teresópolis3 (SONP)1
Platyphora bullata (Stål, 1858)	Nova Friburgo2
Platyphora cincta (Germar, 1821)	Itatiaia3, Teresópolis (SONP)3
Platyphora congener (Stål, 1858) (Fig. 1G)	Nova Iguaçu (ReBio do Tinguá3), Rio de Janeiro (Tijuca3), Teresópolis (SONP)1
Platyphora conviva (Stål, 1858)	Itatiaia3 (INP1)
Platyphora curticollis (Stål, 1857) (Fig. 1H)	Teresópolis (SONP)1
Platyphora dejeani (Germar, 1824) (Fig. 1I)	Casimiro de Abreu (ReBio União)1, Itatiaia3, Nova Iguaçu (ReBio do Tinguá)1, Petrópolis1, Rio de Janeiro (Corcovado3, Tijuca3), Teresópolis2,3 (SONP)1
Platyphora difficilis (Stål, 1859) (Fig. 1J)	Teresópolis (SONP)1
Platyphora dilaticollis (Stål, 1858)	Cambuci (Funil)3, Itatiaia3, Teresópolis (SONP)1
Platyphora fasciatomaculata (Stål, 1857) (Fig. 1K)	Itatiaia (INP)1, Teresópolis (SONP)1
Platyphora fervida (Fabricius, 1775) (Fig. 1L)	Itatiaia3, Teresópolis2 (SONP1,9)
Platyphora figurata (Germar, 1824)	Angra dos Reis3, Rio de Janeiro3
Platyphora flavovittata (Stål, 1858) (Fig. 1M)	Itatiaia3 (INP1), Teresópolis (SONP)1
Platyphora fraterna (Stål, 1857) (Fig. 1N)	Teresópolis (SONP)1
Platyphora histrio (Olivier, 1807)	Angra dos Reis3, Itatiaia3, Rio de Janeiro (Rio de Janeiro2, Corcovado3),
Platyphora irrorata (Stål, 1857)	Itatiaia3, Rio de Janeiro (Corcovado3, Rio de Janeiro3)
Platyphora itatiayensis (Bechyně, 1950) (Fig. 1O)	Itatiaia3, Teresópolis (SONP)1
Platyphora jucunda (Stål, 1857) (Fig. 1P)	Itatiaia3, Teresópolis (SONP)1
Platyphora langsdorfi (Germar, 1824) (Fig. 1Q)	Teresópolis (SONP)1
Platyphora pardalina (Stål, 1858)	Itatiaia3
Platyphora pastica (Germar, 1824) (Fig. 1R)	Angra dos Reis3, Itatiaia3, Rio de Janeiro (Alto da Boa Vista3), Teresópolis (SONP)1
Platyphora pervicax (Stål, 1859)	Itatiaia3
Platyphora princeps Gray, 1832	Itatiaia3
Platyphora reticulata (Fabricius, 1787)	Itatiaia3, Teresópolis (SONP)3
Platyphora semiviridis Jacoby, 1903	Itatiaia3, Resende6
Platyphora signiceps (Stål, 1857)	Itatiaia3, Petrópolis (SONP)3
Platyphora sp.	Itatiaia (INP)1
Platyphora strigilata (Stål, 1859)	Itatiaia3 (INP1)
Platyphora tesselata (Olivier, 1807)	Teresópolis (SONP)3
Platyphora variolaris (Stål, 1859)	Nova Friburgo2
Platyphora vidanoi Daccordi,1993 (Fig. 1S)	Itatiaia3 (INP1), Teresópolis (SONP)1
Platyphora vigintiunopunctata (Chevrolat, 1831)	Itatiaia3, Teresópolis (SONP)2
Platyphora zikani (Bechyně,1950) (Fig. 1T)	Teresópolis (SONP)1
Platyphora zonata (Germar, 1824) (Fig. 1U)	Macaé (Parque Nacional da Restinga de Jurubatiba)1, Itatiaia3, Teresópolis (SONP)1
Stilodes flavosignata (Stål, 1859)	Nova Friburgo2, Rio de Janeiro (Rio de Janeiro2, Corcovado3), Teresópolis (SONP)1
Stilodes jocosa (Stål, 1859)	Rio de Janeiro (Corcovado2,4)
Stilodes nigriventris (Germar, 1824)	Itaguaí2, Macaé (Restinga de Jurubatiba)1, Rio de Janeiro (Corcovado2,4)
Stilodes peltasta (Stål, 1865)	Rio de Janeiro (Corcovado2)
Stilodes sp. 1	Teresópolis (SONP)1,9
Stilodes sp. 2	Teresópolis (SONP)1
Stilodes thetis Stål, 1860 (Fig. 1V)	Itatiaia (INP)1, Teresópolis (SONP)1
Stilodes trimaculicollis Stål, 1859	Rio de Janeiro (Rio de Janeiro3, Corcovado2), Teresópolis (SONP)1
Stilodes (Eustilodes) cordata Achard, 1923	Rio de Janeiro4, Teresópolis (SONP)1
Stilodes (Eustilodes) cornuta (Bechyně, 1947)	Itatiaia3
Stilodes (Eustilodes) denticeps (Stål, 1860)	Macaé4
Stilodes (Grammomades) impuncticollis (Stål, 1859) (Fig. 1W)	Itatiaia3, Laje do Muriaé3, Teresópolis2 (SONP1,9)
Stilodes (Isostilodes) bisbilineata Stål, 1859	Itatiaia3
Trichomela notaticollis (Stål, 1858)	Itatiaia3, Teresópolis (SONP)3
Trichomela xantholoma (Stål, 1857) (Fig. 1X)	Teresópolis (SONP)1
Zygogramma appendiculata Stål, 1859 (Fig. 1Y)	Teresópolis (SONP)1
Zygogramma novemstriata Stål, 1859	Angra dos Reis3
Zygogramma (Tritaenia) mendesi Bechyně, 1948	Itatiaia3,4, Resende3
Zygogramma (Tritaenia) virgata (Stål, 1859)	Rio de Janeiro (Tijuca2)
Chrysomelini: Chrysomelina – 5 genera and 8 species
Lioplacis meridionalis Bechyně, 1948	Itatiaia (INP)1
Phaedon confinis Klug, 1829	Angra dos Reis3, Itatiaia3
Phaedon consimilis Stål, 1860	Rio de Janeiro (Manguinhos3)
Phaedon pertinax Stål, 1860	Nova Friburgo4, Itatiaia3, Resende3, Rio de Janeiro (Manguinhos3)
Pixis clavigera Stål, 1860	Rio de Janeiro (Corcovado2)
Pixis columbina Stål, 1860	Itatiaia7, Teresópolis (SONP)1
Plagiodera gounelli Achard, 1925	Rio de Janeiro (Corcovado2, Tijuca2)
Trochalonota badia (Germar, 1824)	Rio de Janeiro (Anil2, Corcovado2, Tijuca3)
Chrysomelini: Entomoscelina – 1 genus and 1 species
Microtheca ochroloma Stål, 1860	Rio de Janeiro (Deodoro2, Rio de Janeiro2)

1 CLEI; 2 MNRJ; 3 CEIOC; 4 MIZA, 5 MPEG; 6 Olckers 1998; 7 Bechyně 1958; 8 Sampaio and Monné 2016; 9 Flinte et al. 2009b; 10 Flinte et al. 2015.

For documentation of species’ natural history and host plants, data assembled from field expeditions during different research projects conducted at Serra dos Órgãos National Park (), State of Rio de Janeiro, between 2005 and 2010 was used. The duration, months and number of participants of field expeditions per year are as follows: 2005 (1 or 2 days every month, 3 to 5 collectors); 2006 (2 to 4 days every month, 3 to 5 collectors); 2007 (2 to 4 days every month, 3 to 5 collectors); 2008 (1 or 2 days every month, 2 or 3 collectors); 2009 (1 or 2 days every month, 2 or 3 collectors); 2010 (1 or 2 days every month, 1 or 2 collectors).

Study Site

The park covers an area of 20,024 ha of well-preserved Atlantic Rain Forest (see Veloso et al. 1991 for more on local vegetation) and is located ca. 100 km from Rio de Janeiro, in a mountainous area ranging from 80 m to 2263 m elevation. The climate is tropical, with a colder drier season from May to August, and a rainy warmer period from November to February (Flinte et al. 2009b). Mean annual temperature is around 18 °C, maximum of 38 °C and minimum of 0 °C. Annual precipitation varies between 1250 and 1500 mm (Flinte et al. 2008).

Species study and collection

Species were sampled by a combination of manual collecting, sweep nets and malaise traps, during the conduction of other projects with in the park. When a species was initially found in the field, individuals in as many different developmental stages as possible were brought to the laboratory and reared in plastic containers for host plant confirmation and observations on behavior and biology. In an attempt to describe species seasonal distribution in the area, considering data on labels of specimens from all collections, we recorded the different months on which they were collected and summed the number of species per month (independent of year).

Identification and vouchers

species were identified by Mauro Daccordi. host plants were identified by Lucia d’Ávila Freire de Carvalho (Jardim Botânico do Rio de Janeiro) and Luciano Bianchetti (Embrapa/Brasília), by Roberto Lourenço Esteves (Universidade do Estado do Rio de Janeiro), by Rosângela Simão-Bianchini (Herbário SP - Instituto de Botânica) and by Massimo Bovini (Jardim Botânico do Rio de Janeiro). Thiago Marinho Alvarenga (Universidade de Campinas) identified parasitoids. Species collected at Serra dos Órgãos National Park are deposited at CLEI-UFRJ, Rio de Janeiro, Brazil.

Results and discussion

General patterns of richness and distribution

The Neotropical fauna is thought to comprise ca. 38 genera (Daccordi 1996) and 1,020 species (Blackwelder 1944), but these are outdated numbers and no such information could be found specifically for Brazil. One hundred species occurring in the State of Rio de Janeiro were recorded, belonging to 21 genera in one tribe () and three subtribes: , and (Table 1). was represented by 91 species, followed by with eight and with only one species (Table 1). According to Daccordi (1996), there are many endemic and taxa in the Neotropical region, where they reach their maximum diversity. Only nine species are from previously published sources, the other 91 species we found are new records for the state. The genus with most species records was (n = 39) representing 42.4% of found and 40% of total species records, followed by (n = 13, 12.9% of all species recorded) and (n = 11, 10.9%), genera restricted to the Neotropical region (Daccordi 1996). Indeed, is the most species-rich genus in South America (Daccordi 1994), with approximately 500 species (Chaboo et al. 2014 and references therein).

The findings presented here also revealed a high diversity of species and genera, typical for the Atlantic rain forest, in comparison to other studies in South and Central America. Flowers (2004) documented 67 species in 11 genera for Costa Rica and, similar to our work, and were the most species-rich genera. During a six-year field study in a Mexican state, 47 species and eight genera were found; , and were the genera with most species records (Burgos-Solorio and Anaya-Rosales 2004). Chaboo and Flowers (2015) found 158 species and 18 genera for Peru, based on species catalogues.

Species were recorded from only 17 (18.5%) of the state’s municipalities, and 62 species were known from only one location (Table 1). We found a similar pattern in an inventory of for the same state (Flinte et al. 2009a), with most records concentrated near the city of Rio de Janeiro and in large protected areas, such as Petrópolis and Teresópolis (Serra dos Órgãos National Park) and Itatiaia (Itatiaia National Park). The high number of single locality records is probably due more to sampling effort than to endemism, considering that species normally are not very abundant and are more easily collected manually than with traps.

Biology and ecology of at Parque Nacional da Serra dos Órgãos

A total of 43 species were recorded from Serra dos Órgaõs National Park (Table 1, under SONP; Figure 1), all , 42 occurring within the subtribe and only one from (). Within , was the genus with most species records (23 species) out of the 10 genera found, PageBreakPageBreakPageBreakfollowed by (7) and (3), much like the pattern found generally over the state (Table 1). Species showed an enormous variation in color. Adult polymorphism expressed by variation in pronotum color was observed in , (Fig. 1–L1, L2), while the degree of fusion in stripes on the elytra varied greatly among individuals in (Fig. 1–Y1). Other species, such as (Fig. 1–E1), (Fig. 1–I1) and (Fig. 1–N1), displayed strikingly similar coloration to the leaves of their host plant, while other species including (Fig. 1G), (Fig. 1A) and (Fig. 1C) were highly conspicuous to the human eye.

Figure 1.

species in Rio de Janeiro. Some species occurring in Serra dos Órgãos National Park, State of Rio de Janeiro, Brazil. (A); adult (B1) and larva (B2); (C); adult (D1) and larva (D2); adult (E1) and larva (E2); (F); (G); adult (H1), larval cannibalism (H2) and larval aggregation (H3); adult (I1) and larva (I2) () (J); adult (K1) and larva (K2); yellow-pronotum adult and larva (L1) and red-pronotum female ovipositing (L2); (M); adult (N1) and larval aggregation (N2); adult (O1) and larvae (O2); adult (P1) and larval aggregation (P2); adult (Q1) and larva (Q2); (R) (S); adult (T1) and young larvae (T2); (U); (V); adult (W1), eggs (W2) and larva (W3); (X); polymorphic adults in copula (Y1), larvae feeding (Y2), larval cycloalexy (Y3), adult aggregation (Y4), egg mass (Y5), larva attacked by hemipteran nymph (Y6).

The subfamily in SONP exhibited a clear seasonal pattern (Fig. 2), with more species recorded in the hot rainy season, from October to January, than during the drier and colder months, between June and August. This seasonal pattern is well-established for the family in the area, with annual variation in temperature and precipitation and effects on host plant phenology being likely the main drivers of the temporal dynamics in these beetles (Flinte et al. 2009b, 2011, 2015). This is particularly so because many of the records were made at altitudes above 1000 m, where the pattern normally more closely resembles that found in the subtropical zone (e.g. Medeiros and Vasconcellos-Neto 1994, Nogueira-de-Sá et al. 2004). However, the present results are, to our knowledge, the first to examine the seasonal pattern for such a large number of species in a single area. Ideally, a standardized collecting effort across the year would better describe the seasonal differences we observe here. However, as we have conducted research in the area over many years, doing the same surveys at least once a month every year, we are confident that this represents the seasonal pattern of chrysomeline species occurrence in the area. Moreover, the species which were intensively studied over the year, (Flinte et al. 2015), and (Flinte et al. 2009b) exhibited the same low densities during the drier and colder months.

Figure 2.

Seasonal distribution of . Number of species recorded on each month, obtained for 40 species from collections and fieldwork, in Serra dos Órgãos National Park, southeast Brazil.

Host plant and/or biological information were found for almost half of the species (n = 16) (Table 2) that we (VF, AA, MVM, RFM) collected in the park (n = 35). was the most common host plant family, followed by , , and . As expected, this is a pattern that reflects preference for (Jolivet and Hawkeswood 1995, Chaboo et al. 2014 and references therein). Intense host plant defoliation was observed in (Fig. 1–W1, W2, W3) on (), (Fig. 1–N1, N2) on () and (Fig. 1–Y1, Y2, Y3) on ().

Table 2.

Ecological data on species. Species at Serra dos Órgãos National Park with host plant record and/or biological data obtained from our research at the area. Published records are indicated by numbers (see footnote for references).

Species	Host plant family	Host plant species	Reproduction	Larvae
Calligrapha polyspila	Malvaceae 1	?	oviparous1	?
Deuterocampta sedula	?	?	?	solitary
Grammodesma rubroaenea	Asteraceae	?	oviparous	solitary
Platyphora axillaris	Solanaceae 2	Solanum scuticum 2	larviparous2	solitary
Platyphora curticollis	Solanaceae	Solanum swartzianum	larviparous	aggregated
Platyphora dejeani	Convolvulaceae	Ipomoea philomega	oviparous	solitary
Platyphora fervida	Solanaceae 3	Solanum lhotskyanum 3	larviparous	solitary
Platyphora flavovittata	Apocynaceae	?	oviparous	?
Platyphora fraterna	Solanaceae	Solanum swartzianum	larviparous	aggregated
Platyphora itatiayensis	Solanaceae	Solanum megalochiton	larviparous	aggregated
Platyphora jucunda	Solanaceae 4	Solanum swartzianum	larviparous	aggregated
Platyphora langsdorfi	Convolvulaceae 5	Ipomoea philomega	oviparous	solitary
Platyphora zikani	Solanaceae	Solanum swartzianum	larviparous	aggregated
Stilodes (Grammomades) impuncticollis	Solanaceae 3	Capsicum mirabile 3	oviparous	solitary
Stilodes sp. 1	Asteraceae	Baccharis stylosa	?	?
Zygogramma appendiculata	Malvaceae	Callianthe regnelli, Callianthe rufinerva	oviparous	aggregated

1 Grissell et al. 1987; 2 Flinte et al. 2015; 3 Flinte et al. 2009b; 4 Olckers 2000; 5 Jolivet and Hawkeswood 1995.

Maternal care was not recorded for any species in this study, although subsocial behavior is known in the subfamily for several species, including some and species (Windsor et al. 2013, Chaboo et al. 2014). Other interesting behavioral defenses were recorded, including larval cycloalexy in (Fig. 1–Y3) and (Fig. 1–H3), a defensive behavior of gregarious circular formation at rest (Jolivet et al. 1990, Vanconcellos-Neto and Jolivet 1994, Dury et al. 2014). Additionally, we recorded larvae of (Fig. 1–N2) attaching trichomes from host plant leaves to hairs on their backs, a behavior already described in (Bernardi and Scivittaro 1991), which may contribute to larval camouflage. Larval PageBreakaggregations were observed in many species (Table 2), but also for adults of on young folded leaves in the field (Fig. 1–Y4). Larval gregarious behavior may serve to reduce individual risk against small invertebrate parasitoids and predators, and promote defense against larger predators through the cumulative effect of individuals’ toxins (Grégoire 1988). Thanatosis (“feigning death”) was observed in adults of (Fig. 1–E1), (Fig. 1–L1, L2), and (Fig. 1–N1), and both in adults and larvae of (Fig. 1–W1, W3).

Seven oviparous and seven larviparous species were found, most being new records of reproductive biology (Table 2). is the subfamily of leaf beetles with the most diversity in reproductive biology, containing oviparous, ovoviviparous and viviparous species (Bontems 1988), sometimes in the same genus, and also different levels of social behavior (Chaboo et al. 2014). The last two types of development may be more costly to the mothers, but ensure a quicker development of the vulnerable larval stage, among other advantages, as proposed by Jolivet and Hawkeswood (1995) and Chaboo et al. (2014 and references therein), which is why it is sometimes considered to be a parental care preceding birth (Hinton 1981). Interestingly, viviparous species may result in solitary larvae, as in (Fig. 1–E2), or larval aggregations, as in (Fig. 1–P2). Oviparous species may also have solitary or gregarious larvae, as in (Fig. 1–I2) and (Fig. 1–Y2), respectively, although larval aggregations seem rarer in this type of reproduction. In their work on subsocial PageBreakneotropical , Windsor et al. (2013) found, among species, two with solitary larvae and nine which formed larval aggregations, but all eleven species were larviparous. We observed a single case of larval cannibalism in the viviparous during laboratory rearing (Fig. 1–H2), a behavior already described for some genera (Wade 1994, Mafra-Neto and Jolivet 1996, Windsor et al. 2013) that grants nutritional benefits.

Except for the eggs of , which are laid in masses on the underside of its host plant leaves (Fig. 1–Y5), no other eggs of oviparous were found in the field. This is probably because chrysomelids often lay their eggs in the soil or in secluded parts of plants (Selman 1994). All oviparous species reared in laboratory laid chorion-covered yellowish eggs on the bottom of the vials or on leaves, normally grouped in clutches (Fig. 1–W2). In the field, the number of eggs of varied from 80 to 100 per group (90.4 ± 8.3 SD; n=7 egg masses), and larval aggregations comprised between 10 and 233 individuals per group (49.2 ± 38.7; n=50 groups). Larvae of different egg masses may cohabit the same aggregation of this species, since differently sized larvae were observed in the same aggregation. larvae (Fig. 1–N2) were grouped in aggregations of 24.9 ± 13.1 SD individuals (n=14 groups), with a minimum of seven and maximum of 44 larvae per group. No pupa has yet been found in the field, but in the laboratory, prepupae always buried themselves in earth layer at the bottom of the vials. While pupation in may be arboreal or underground (Takizawa 1976), it seems that underground pupation is most common in our taxa as indicated by laboratory rearing.

Only a few observations on natural enemies of were made. Phoretic wasps of () were found on adults of (Fig. 1–D1) and (Fig. 1–B1). are well known parasitoids of chrysomeline larvae (Cox 1994). On one occasion, a () nymph was seen preying on a larva of (Fig. 1–Y6). Many chrysomeline species presented unprotected larvae without any apparent behavioral defense, but several gain chemical defenses by the sequestration of host plants toxins or by synthesizing defensive compounds from plant precursors, especially in (Pasteels et al. 2001, Termonia et al. 2002).

Conclusions

The high proportion of new host, biological data and occurrence records in Rio de Janeiro reflects the limited knowledge we have about this subfamily in this immediate area. In Brazil, the picture is not very different, as no inventory for the subfamily has been compiled and the relatively few published records come from ecological studies such as Medeiros and Vasconcellos-Neto (1994), Medeiros et al. (1996), Macedo et al. (1998), Vasconcellos-Neto and Jolivet (1998), and Flinte et al. (2009b, 2015). However, is known to be very species-rich in Brazil, including known enPageBreakdemic species, such as (Macedo et al. 1998). Because of their high host specificity (Jolivet 1988, Jolivet and Hawkeswood 1995) and low dispersal ability (Freijeiro and Baselga 2016) the chrysomelines are expected to have many narrowly distributed species, especially in mountainous areas, as has already been found for other tropical species (García-Robledo et al. 2016, Macedo et al. 2016). These traits then would make these Brazil species especially vulnerable to extinction as the mountains within the Atlantic forest biome are largely degraded and threatened (Martinelli 2007).