Checklist and identification key of Anomalini (Coleoptera, Scarabaeidae, Rutelinae) of Costa Rica

Abstract A checklist and identification key for the species of the tribe Anomalini in Costa Rica are presented. The Anomalini species are important economically, as they have larvae that are or can become agricultural pests, as well as ecologically, having potential as bioindicators. In spite of their importance and richness, identification tools for the group in the Neotropics remain scarce. The Costa Rican fauna comprises six genera (Anomala, Anomalorhina, Callistethus, Epectinaspis, Moroniella, and Strigoderma) and a total of 120 species. Anomala contusa Filippini, Micó, Galante, 2015 is proposed as a synonym of Anomala inbio (Ramírez-Ponce, Bitar, Curoe 2014); Anomala limon nom. n. is proposed as a new name for Anomala inbio Filippini, Galante, Micó, 2015, a homonym of Anomala inbio (Ramírez-Ponce, Bitar, Curoe, 2014); Anomala cinaedias nom. n. is proposed as a new name for Anomala chloropyga Ohaus, 1897, a homonym of Anomala chloropyga Burmeister, 1844; and Anomala chrysomelina is moved to the genus Callistethus.


Introduction
One reason for the "taxonomic neglect" (Jameson et al. 2003) of the genus Anomala over the past centuries is due to the inverse relationship between the number of species in that genus and the available taxonomic information about them. Most descriptions date to the early 20 th century and earlier, and the brevity of these descriptions makes reliable identification impossible without consulting the type material. The largest contribution to the study of the Anomalini in Central America was made by H. W. Bates, with his collaboration on the volumes of Biologia Centrali-Americana (1888 for the volume on Anomalini).
Adults of most Anomalini species are nocturnal and are easily captured by light traps. Although they may serve as bioindicators for ecosystem conservation, the lack of proper identification tools makes such a role difficult (Morón 1997). The larvae of Anomalini are subterranean feeders, consuming roots and organic material (Ritcher 1966), and are considered ecologically important for their role in the airing of soil and decomposition of organic material (Morón and Aragón 2003). Some species are known to cause damage to crops, and some have become invasive agricultural pests in countries where they are adventive. While no invasive species of Anomalini have been recorded in Costa Rica (I3N Costa Rica http://invasoras.acebio.org), a few species of Anomala have been found to be associated with different crops, together with the scarab beetles Phyllophaga (Melolonthinae) and Cyclocephala species (Dynastinae). The lack of knowledge about the species' larval morphology, however, makes it difficult to identify which species are associated with crop damage, and identification is usually done on adults collected nearby (Abarca and Quesada 1997). Larval descriptions are available for only four of the species recorded in Costa Rica (Anomala discoidalis, A. undulata, A. ludoviciana, A. cupricollis;Micó et al. 2003).
In this paper, a checklist for the Anomalini of Costa Rica is presented, which comprise 120 species, including photographs of the habitus and drawings of male genitalia for nearly all species, and a comprehensive key for use in identification. This work is the final part of the of a three year taxonomic study on Costa Rican Anomalini performed by the authors, which has resulted in the description of more than 50 new species of Anomala and Callistethus (Filippini et al. 2013(Filippini et al. , 2014(Filippini et al. , 2015a Anomala contusa Filippini, Micó Galante 2015 is proposed as a synonym of A. inbio (Ramírez-Ponce, Bitar, Curoe, 2014); Anomala limon is proposed as a new name for A. inbio Filippini, Galante, Micó, 2015, a homonym of A. inbio (Ramírez-Ponce, Bitar, Curoe, 2014; Anomala cinaedias is proposed as a new name for A. chloropyga Ohaus, 1897, homonym of A. chloropyga Burmeister, 1844 and Anomala chrysomelina is moved to the genus Callistethus.

Methods
Specimens cited in this publication are deposited in the following collections: Procedures for the preparation of endophalli, measurements, definitions, and morphological terminology follow Filippini et al. (2013Filippini et al. ( , 2014. In contrast to Ramírez-Ponce and Morón (2009), who group New World Anomala species into a new genus (Paranomala) we follow the traditional inclusion in the genus Anomala (Jameson et al. 2003) as a more conservative classification, while waiting for a more extensive study at global scale. We use the phylogenetic species concept described by Wheeler and Platnick (2000), which defines species as the smallest aggregation of sexual populations that are diagnosable by a unique combination of character states.

BMNH
Line drawings were traced using a GNU image manipulation program (GIMP version 2.8, www.gimp.org). Original drawings were produced with the aid of a camera lucida attached to a stereo microscope (Leica M80) for endophalli, or from photographs for aedeagi (taken with a Leica DFC450 camera mounted on a Leica M205C stereo microscope).
For each species in the checklist, the provinces where it is located are given in Figure 1. The distribution data were gathered using labels from identified specimens in the MNCR and CEUA collections, and the Atta database (http:// http://atta.inbio. ac.cr/) of INBio, Costa Rica.
Species identification was undertaken by consulting original species descriptions and, when possible, by type comparison. For a list of the types consulted, see Filippini et al. (2015b). Nomenclatural changes are suggested in accordance with the International Code of Zoological Nomenclature.

Data resources
The data from specimens deposited at CEUA are deposited at GBIF, the Global Bio diversity Information Facility, http://www.gbif.es/ic_colecciones.php?ID_Coleccion=9709

Results
In the last three years, new species descriptions have led to a 49% increase in the known Costa Rican fauna of the tribe (Filippini et al. 2013(Filippini et al. , 2014(Filippini et al. , 2015a. The lack of taxonomical and faunistic studies in other Neotropical countries, however, makes it difficult to make comparisons with similar regions, or even to determine which species are endemic. For example, only 42 Anomalini species are registered for Panama (Ratcliffe 2002), and only 79 for Ecuador (which has a surface area five times that of Costa Rica) (Paucar-Cabrera 2005). It is safe to say that most of the Anomalini diversity in the Neotropical region is yet to be described.
The present work does not exhaust the actual diversity of Costa Rican Anomalini; there were at least a few species that, for reasons such as the scarcity of specimens, were not included in the descriptive work.
Based on the data gathered from the studied specimens, the richest habitats for the species were various types of evergreen tropical forests located on the slopes of the country's main mountain ranges (unpublished material).
The comparison between specimens of A. contusa and the description and illustrations of A. inbio lead to the conclusion that they are the same species. In particular, the diagnostic characteristics that include the species in the trapezifera species group (pronotum bronze with an irregular shaped macula, elytra with ochre background covered by small numerous flecks, tridentate protibia, tectum shorter than or similar in size to the basal piece) (Filippini et al. 2015), the particular texture of the pronotum of this species (not shared by other Costa Rican species), and the peculiar shape of aedeagus, are coincident.
A. inbio (Ramírez-Ponce, Bitar, Curoe, 2014) was originally described in the genus Paranomala. For the reasons explained in Methods it is here placed in the genus Anomala.
Etymology. this species is specific to the Costa Rican Province of Limón, where most of the type material was collected. To be used as a name in apposition.
Anomala chloropyga Ohaus, 1897 is a homonym for A. chloropyga Burmeister, 1844, a species from the Philippines, so a replacement name is necessary.
Etymology. from the Greek κιναιδίας, a precious stone, for the smooth and shiny appearance of this species. To be used as a noun in apposition. (Bates, 1888), comb. n. Bates, 1888 is moved to the genus Callistethus, as it presents the following diagnostic characteristics of this genus, as described in Filippini et al. (2015): wide interocular space and small elongated eyes; posterior margin of the pronotum is smooth, without bead, and directly opposite to the scutellum; presence of a mesosternal process produced slightly beyond the apex of the mesocoxae.