Where host plant goes , galls go too : new records of the Neotropical galling Cecidomyiidae ( Diptera ) associated with Calophyllum brasiliense Cambess . ( Calophyllaceae )

Calophyllum brasiliense, a Neotropical species, hosts seven galling Cecidomyiidae reported only for Brazil. Our hypothesis is the distribution of the galling insects associated with C. brasiliense is similar to that of the host plant. We checked gall occurrence by field sampling and by searching data in literature and herbaria. Current results indicate that the occurrence of galling Cecidomyiidae associated with C. brasiliense extends to 13 Brazilian states, as well as to 11 countries of Neotropical America.

Galling insects are specialized herbivores, capable of inducing the redifferentiation of specialized plant tissues (Oliveira and Isaias 2010), mainly by hyperplasia and/or cell hypertrophy (Mani 1964).Redifferentiated tissues altogether constitute the galls, abnormal structures that guarantee nutrition and a safe site for development of galling insects and their offspring (Shorthouse et al. 2005).The Cecidomyiidae (Diptera) are the most diverse taxa among the galling insects, with 6,203 described species grouped in 736 genera (Gagné and Jaschhof 2014) with global distribution.In Brazil, there are about 200 species of Cecidomyiidae, with 92% of them monophagous (Carneiro et al. 2009).
The interaction between host plant and galling insect usually results in a gall morphotype with peculiar shape, size, color and indumentum (Isaias et al. 2013(Isaias et al. , 2014)), which reflects the extended phenotype of the gallinducing insect (Stone and Schönrogge 2003).Actually, galls are believed to be intimate representations of galling insects' diversity (Carneiro et al. 2009).This specificity is well expressed in plants capable of hosting several gall morphotypes, the super host plants (Isaias et al. 2013(Isaias et al. , 2014)).Distinct morphotypes are irrefutable proof suggesting each plant species has a different set of responses to different insect species' stimuli, which increases diversity of morphotypes (Araújo et al. 2013).
Calophyllum brasiliense is a superhost of galling cecidomyiids, with five gall morphotypes related to five distinct galling species: two fusiform gall morphotypes, the first induced by Contarinia gemmae Maia, 2003

Notes oN GeoGraphic DistributioN
fusiform gall induced by L. linearis Maia, 2003(Madeira et al. 2002); and a rolling gall induced by an undescribed species of Cecidomyiidae (Maia 2013;Proença and Maia 2015).
Information on the distribution of galling insects is scarce and mostly limited to the localities where species were originally sampled (Gagné and Jaschhof 2014).This scarcity limits the knowledge on dispersion, evolution and population dynamics of those specialized herbivores.However, information on distribution of C. brasiliense has been better documented due to major efforts on plant sampling, studying and systematically adequate records in herbaria.A recent study has documented at four Brazilian sites the distribution of galling cecidomyiids associated with C. brasiliense: Rio de Janeiro, São Paulo, Minas Gerais and Goiás states (Proença and Maia 2015), in the Atlantic Forest and Cerrado biomes.We have chosen C. brasiliense as a model species to check the viability of using virtual collections of plant material to improve the estimation of geographical distributions of galling insects.Considering the specificity and the life cycles of the involved galling organisms (Mani 1964), our hypothesis is that the galling insects accompany C. brasiliense throughout its geographic distribution.Nevertheless, some constraints to the irradiation of galling herbivores should occur.Our objective is to contribute to the knowledge on geographic distribution of the Cecidomyiidae associated with Calophyllum brasiliense, taking for granted the Neotropical occurrence of this host plant.Current objectives are: i) verify matching between distribution of Cecidomyiidae related to C. brasiliense and host plants; and ii) check and describe the existence of new records for association of C. brasiliense and its related galling cecidomyiids throughout virtual and classical herbaria.
Current data were obtained both from primary and secondary sources.The primary sources were field sampling of host plants at Parque Estadual da Serra dos Montes Altos, municipality of Sebastião Laranjeiras (14°23ʹ41ʺ S, 043°0ʹ35ʺ W), Bahia state, from April to March, 2015, and at Parque Estadual Acaraí, municipality of São Francisco do Sul (26°14ʹ36ʺ S, 048°38ʹ17ʺ W), Santa Catarina state, from June to August, 2015.Stem and leaf galls were sampled, photographed and described according to Isaias et al. (2013).Vouchers are deposited in Herbarium Joinvillea, of the Universidade da Região de Joinville (JOI) and in the Herbarium of Universidade do Estado da Bahia (HUNEBA), under registration numbers JOI 16568 and HUNEBA 24983, respectively.
The secondary sources were a review of literature data in which C. brasiliense and its cecidomyiid-induced galls are reported (Madeira et al. 2003;Maia et al. 2008;Luz et al. 2012;Maia 2013;Julião et al. 2014aJulião et al. , 2014b;;Proença and Maia 2015), and direct observation of herbaria The knowledge on distribution of the cecidomyiids associated with C. brasiliense up to 2015 was restricted to Rio de Janeiro, São Paulo, Minas Gerais and Goiás states, in Brazil, with six gall morphotypes recorded (Maia 2013;Proença and Maia 2015).A seventh morphotype (clavate leaf gall), also induced by a Cecidomyiidae, was reported for Minas Gerais and Mato Grosso do Sul states (Luz et al. 2012;Julião et al. 2014a).This distribution is widened to the Amazonas state (Julião et al. 2014a), which was not included in  the last report of cecidomyiid distribution (Proença and Maia 2015).
The distribution of cecidomyiids associated with C. brasiliense reflects this family's overlapping distribution with the host plant.As we are dealing with species-specific interactions, we can confidently assume that host plant geographical distribution influences the geographical distribution of its associated galling herbivores.The wide geographical distribution of C. brasiliense reflects in a wide geographical distribution for its associated herbivores.Geographical distribution analysis of the seven cecidomyiid species associated with C. brasiliense in the Neotropical region (Gagné and Jaschhof 2014) provides evidence that the clavate gall morphotype has the most restricted distribution (Minas Gerais and Mato Grosso do Sul states); C. gemmae (from São Paulo to Amazonas state), L. conspicua (Paraná and Amazon states) and the leaf intralaminar gall (Santa Catarina and Amazon states) extends being the southern tropical region.L. elliptica and L. linearis have a wide distribution (Santa Catarina to México and the Caribean), and L. caulinaris has the widest distribution of all (from Santa Catarina state in Brazil to Costa Rica).Consequently, there are four species restricted to Brazil, two previously described by Gagné and Jaschhof (2014), C. gemmae and L. conspicua, and two herein recorded for the first time, inducers of leaf rolling gall and of clavate gall morphotypes.
Latitudinal distribution of cecidomyiida associated with C. brasiliense provides evidence that five species occur in the Subtropical region (Santa Catarina and Paraná states), seven in the south tropical region (from São Paulo to Amazonas state), and three species in the north region (Amapá and Roraima states up to Mexico).Geographical distribution of Cecidomyiidae overlaps to the distribution of C. brasiliense.A reduction and a turnover of cecidomyiid species are observed towards the northern hemisphere, probably due to constraint of phenotypic plasticity (Valladares et al. 2007) imposed both on the host plant and galling herbivores.This hypothesis is corroborated by the report of Trioza sp.(Tryozidae: Hemiptera) as the inducer of the marginal leaf rolling gall morphotype in C. brasiliense, exclusively in Costa Rica (Martin and Hollis 1992).In addition, formation of ecotypes of host plants due to their wide geographical distribution may also be a start point for the irradiation of galling organisms.Anyway, knowledge on the biodiversity of south hemisphere increased due to efforts in sampling along Neotropical countries, which is also true for the Cecidomyiidae.
The analyses of the model species C. brasiliense and its associated galling cecidomyiids have proved the usefulness of virtual herbaria not only to document plant distribution, but also to document host plantgalling herbivore associations.The better the collection is, the better the estimative is made, which is herein observed for New York Botanical Gardens digital collection.Current estimation widens the geographical distribution of Cecidomyiidae-C.brasiliense systems throughout Brazilian biomes.