Varying and unchanging whiteness on the wings of dusk-active and shade-inhabiting Carystoides escalantei butterflies.

Whiteness, although frequently apparent on the wings, legs, antennae, or bodies of many species of moths and butterflies, along with other colors and shades, has often escaped our attention. Here, we investigate the nanostructure and microstructure of white spots on the wings of Carystoides escalantei, a dusk-active and shade-inhabiting Costa Rican rain forest butterfly (Hesperiidae). On both males and females, two types of whiteness occur: angle dependent (dull or bright) and angle independent, which differ in the microstructure, orientation, and associated properties of their scales. Some spots on the male wings are absent from the female wings. Whether the angle-dependent whiteness is bright or dull depends on the observation directions. The angle-dependent scales also show enhanced retro-reflection. We speculate that the biological functions and evolution of Carystoides spot patterns, scale structures, and their varying whiteness are adaptations to butterfly's low light habitat and to airflow experienced on the wing base vs. wing tip.

Organismal responses to habitat change: herbivore performance, climate and leaf traits in regenerating tropical dry forests.

The ecological effects of large-scale climate change have received much attention, but the effects of the more acute form of climate change that results from local habitat alteration have been less explored. When forest is fragmented, cut, thinned, cleared or otherwise altered in structure, local climates and microclimates change. Such changes can affect herbivores both directly (e.g. through changes in body temperature) and indirectly (e.g. through changes in host plant traits). We advance an eco-physiological framework to understand the effects of changing forests on herbivorous insects. We hypothesize that if tropical forest caterpillars are climate and resource specialists, then they should have reduced performance outside of mature forest conditions. We tested this hypothesis with a field experiment contrasting the performance of Rothschildia lebeau (Saturniidae) caterpillars feeding on the host plant Casearia nitida (Salicaceae) in two different aged and structured tropical dry forests in Area de Conservación Guanacaste, Costa Rica. Compared to more mature closed-canopy forest, in younger secondary forest we found that: (1) ambient conditions were hotter, drier and more variable; (2) caterpillar growth and development were reduced; and (3) leaves were tougher, thicker and drier. Furthermore, caterpillar growth and survival were negatively correlated with these leaf traits, suggesting indirect host-mediated effects of climate on herbivores. Based on the available evidence, and relative to mature forest, we conclude that reduced herbivore performance in young secondary forest could have been driven by changes in climate, leaf traits (which were likely climate induced) or both. However, additional studies will be needed to provide more direct evidence of cause-and-effect and to disentangle the relative influence of these factors on herbivore performance in this system.

Revision of Belvosia Robineau-Desvoidy (Diptera, Tachinidae) and 33 new species from Area de Conservación Guanacaste in northwestern Costa Rica with a key to known North and Mesoamerican species.

Background: This revision is part of a continuing series of taxonomic work aimed at the description of new taxa and the redescription of known taxa of the Tachinidae of Area de Conservación Guanacaste in northwestern Costa Rica. Here we describe 33 new species in the genus Belvosia Robineau-Desvoidy, 1830 (Diptera: Tachinidae). All species described here were reared from this ongoing inventory of wild-caught caterpillars spanning a variety of families (Lepidoptera: Erebidae, Eupterotidae, Noctuidae, Notodontidae, Saturniidae, and Sphingidae). We provide a morphological description of each species with limited information on life history, molecular data, and photographic documentation. In addition to the new species, the authors provide a redescription of the genus Belvosia, as well as provide a key to the identification of the species present in the Meso- and North-American fauna. New information: The following 33 new species of Belvosia Robineau-Desvoidy, 1830, all authored by Fleming & Woodley, are described: Belvosiaadrianguadamuzi Fleming & Woodley sp. n., Belvosiaanacarballoae Fleming & Woodley sp. n., Belvosiaangelhernandezi Fleming & Woodley sp. n., Belvosiabrigittevilchezae Fleming & Woodley sp. n., Belvosiaalixtomoragai Fleming & Woodley sp. n., Belvosiacarolinacanoae Fleming & Woodley sp. n., Belvosiaciriloumanai Fleming & Woodley sp. n., Belvosiadiniamartinezae Fleming & Woodley sp. n., Belvosiaduniagarciae Fleming & Woodley sp. n., Belvosiaduvalierbricenoi Fleming & Woodley sp. n., Belvosiaeldaarayae Fleming & Woodley sp. n., Belvosiaeliethcantillanoae Fleming & Woodley sp. n., Belvosiafreddyquesadai Fleming & Woodley sp. n., Belvosiagloriasihezarae Fleming & Woodley sp. n., Belvosiaguillermopereirai Fleming & Woodley sp. n., Belvosiaharryramirezi Fleming & Woodley sp. n., Belvosiahazelcambroneroae Fleming & Woodley sp. n., Belvosiajorgehernandezi Fleming & Woodley sp. n., Belvosiajosecortezi Fleming & Woodley sp. n., Belvosiajoseperezi Fleming & Woodley sp. n., Belvosiakeinoraragoni Fleming & Woodley sp. n., Belvosialuciariosae Fleming & Woodley sp. n., Belvosiamanuelpereirai Fleming & Woodley sp. n., Belvosiamanuelriosi Fleming & Woodley sp. n., Belvosiaminorcarmonai Fleming & Woodley sp. n., Belvosiaosvaldoespinozai Fleming & Woodley sp. n., Belvosiapabloumanai Fleming & Woodley sp. n., Belvosiapetronariosae Fleming & Woodley sp. n., Belvosiaricardocaleroi Fleming & Woodley sp. n., Belvosiarobertoespinozai Fleming & Woodley sp. n., Belvosiarostermoragai Fleming & Woodley sp. n., Belvosiaruthfrancoae Fleming & Woodley sp. n., Belvosiasergioriosi Fleming & Woodley sp. n.Belvosiacanalis Aldrich, 1928 is reared and recorded from the inventory; new information relative to host is provided and the species is rediscribed.The following are proposed by Fleming & Woodley as new synonyms of Belvosia Robineau-Desvoidy, 1830: Brachybelvosia Townsend, 1927 syn. n., Belvosiomimops Townsend, 1935 syn. n.The following three new combinations are proposed as a result of the new synonymies: Belvosiabrasilensis (Townsend, 1927), comb. n.; and Belvosiabarbiellinii (Townsend, 1935), comb. n.The authors also propose the following new synonymies: Belvosiabrasilensis (Townsend, 1927) = Belvosiaaurulenta (Bigot, 1888), syn. n.; Belvosiapollinosa Rowe, 1933 = Belvosiaborealis Aldrich, 1928 syn. n.; Belvosiaweyenberghiana (Wulp, 1883) = Belvosiafuliginosa (Walker, 1853) syn. n.; Belvosiabrasiliensis Townsend, 1927 = Belvosiafuliginosa (Walker, 1853) syn. n.; Belvosialuteola Coquillett, 1900 = Belvosiaochriventris (Wulp, 1890) syn. n.; Belvosiasocia (Walker, 1853) = Belvosiaproxima (Walker, 1853) syn. n.; Belvosiachrysopyga (Bigot, 1887) = Belvosiaunifasciata (Robineau-Desvoidy, 1830) syn. n.; Belvosiachrysopygata (Bigot, 1888) = Belvosiaunifasciata (Robineau-Desvoidy, 1830) syn. n.

Cryptic biodiversity of tropical hesperiid caterpillar-attacking parasitoid wasps: three new species of Creagrura Townes (Hymenoptera, Ichneumonidae, Cremastinae) from Costa Rica and Perú.

Background: We describe three new species of the previously monotypic genus Creagrura Townes from Central and South America: C.alejandromasisi sp. n. and C.rogerblancoi sp. n. from Costa Rica and C.allpahuaya sp. n. from Peru, all of which emphasise the unknown parasitoid insect diversity yet to be revealed in the tropics. New information: Host relationships of the two Costa Rican species are described in detail. In addition, it is inferred that the Creagrura wasps find and oviposit in the caterpillar when it is exposed at night, rather than when it is concealed during daylight hours.

Aprica: a new genus and life history for the pteridivore Xanthiapatula Druce, 1898 (Lepidoptera, Noctuidae).

Aprica Goldstein, gen. nov. is described to accommodate Xanthiapatula Druce, 1898. Recent discovery of its larva, which has been recorded eating foliage of species in six families of leptosporangiate ferns, suggest a possible subfamily assignment within the Eriopinae, but this cannot be substantiated based on adult morphology. This species has no obvious close relatives either among the core noctuid pteridivore genera currently recognized in the Eriopinae (e.g., Callopistria Hübner, [1821]), nor among genera more recently discovered to be fern-feeders but which remain incertae sedis with respect to subfamily (e.g., Leucosigma Druce, 1908, Lophomyra Schaus, 1911). The recorded foodplant profile is similar to that of another ambiguously placed Nearctic species Fagitanalittera (Guenée, 1852) (Noctuidae: Noctuinae: Xylenini, incertae sedis) with which it shares no obvious synapomorphies.

Cryptic diversity in Lithobateswarszewitschii (Amphibia, Anura, Ranidae).

Lithobateswarszewitschii is a species of ranid frog distributed from southern Honduras to Panama. This species suffered severe population declines at higher elevations (above 500 m a.s.l.) from the 1980s to early 1990s, but there is more recent evidence of recovery in parts of its range. Here we advocate for the status of Lithobateswarszewitschii as a candidate cryptic species complex based on sequence data from mitochondrial genes CO1 and 16S. Using concatenated phylogenies, nucleotide diversity (K2P-π), net between group mean distance (NBGMD) (πnet) and species delimitation methods, we further elucidate cryptic diversity within this species. All phylogenies display polyphyletic lineages within Costa Rica and Panama. At both loci, observed genetic polymorphism (K2P-π) is also high within and between geographic populations, surpassing proposed species threshold values for amphibians. Additionally, patterns of phylogeographic structure are complicated for this species, and do not appear to be explained by geographic barriers or isolation by distance. These preliminary findings suggest L.warszewitschii is a wide-ranging species complex. Therefore, we propose further research within its wider range, and recommend integrative taxonomic assessment is merited to assess species status.

A revision of Dolichogenidea (Hymenoptera, Braconidae, Microgastrinae) with the second mediotergite broadly rectangular from Area de Conservación Guanacaste, Costa Rica.

The first species of Dolichogenidea (Hymenoptera: Braconidae, Microgastrinae) with the second mediotergite broadly quadrate to rectangular are revised, and eight new species from Area de Conservación Guanacaste (ACG), Costa Rica are described, all authored by Fernandez-Triana & Boudreault: alejandromasisi, angelagonzalezae, carlosmanuelrodriguezi, genuarnunezi, josealfredohernandezi, melaniamunozae, rogerblancoi, and yeimycedenoae. A new species group (carlosmanuelrodriguezi) within the genus is proposed to accommodate those species, as well as additional undescribed species from the Neotropical region found in collections. All new species are found in rainforests (120-900 m) and all are parasitoids of Depressariidae (except for one species parasitizing Choreutidae). The unique shape of the second mediotergite and long ovipositor are features shared with the alejandromorai species group in the genus Apanteles, an example of convergent evolution; both wasp groups also parasitize similar hosts in ACG.

Checklist of invertebrates and conspicuous fishes in rocky reefs and Sargassum beds in the North Pacific of Costa Rica / Listado de invertebrados y peces conspicuos en arrecifes rocosos y mantos de Sargassum en el Pacífico Norte de Costa Rica

Introduction: The marine biodiversity is well known in some areas and for some marine ecosystems of the Pacific coast of Costa Rica. The marine sector of Área de Conservación Guanacaste, in the North Pacific of the country, is a priority area for conservation of marine and coastal biodiversity. Objective: Our aim was to increase the knowledge of marine biodiversity in a conservation priority site in Costa Rica and in poorly or scarcely studied marine ecosystems, such as rocky reefs and Sargassum beds, respectively. Methods: The study was conducted in Bahía Salinas and Golfo de Santa Elena in 2013. In January, four rocky reefs were visited; while in February three rocky reefs and one Sargassum bed were visited. We identified the macroinvertebrates and conspicuous fish species through underwater visual census methods (timed swim and band transects). The mean density of each species was estimated through band transects method. Results: A total of 25 invertebrates and 34 fish species were identified. We found a high number of species of invertebrates (23) and fish species (33) in rocky reefs, but few species on Sargassum beds (eight invertebrates and ten fish species). Species composition differed between marine ecosystems. Conclusions: Our study further evidence the limited knowledge of invertebrates and fish communities in these marine ecosystems in the North Pacific of Costa Rica. Considerably more surveys and field work are required to support management decisions that are backed by sound scientific knowledge.

Introducción: La biodiversidad marina es bien conocida en algunas áreas y para algunos ecosistemas marinos de la costa Pacífica de Costa Rica. El sector marino del Área de Conservación Guanacaste, en el Pacífico norte del país, es un área prioritaria para la conservación de la biodiversidad marina y costera. Objetivo: Nuestro objetivo fue el de incrementar el conocimiento de la biodiversidad marina en un sitio prioritario de conservación de Costa Rica y en ecosistemas marinos poco o nulamente estudiados, como son los arrecifes rocosos y mantos de Sargassum, respectivamente. Métodos: El estudio se realizó en Bahía Salinas y Golfo de Santa Elena en el 2013. En enero, cuatro arrecifes rocosos fueron visitados; mientras que en febrero tres arrecifes rocosos y un manto de Sargassum fueron visitados. Identificamos las especies de invertebrados y peces conspicuos a través métodos de censos visuales submarinos (buceos cronometrados y transectos de banda). La densidad promedio de cada especie se estimó mediante el método de transectos de banda. Resultados: Se registró un total de 25 invertebrados y 34 especies de peces. Encontramos un mayor número de especies de invertebrados (23) y peces (33) en los arrecifes rocosos, pero pocas especies en el manto de Sargassum (ocho especies de invertebrados y diez de peces). La composición de especies difirió entre ecosistemas marinos. Conclusiones: Nuestro estudio evidencia el limitado conocimiento de las comunidades de invertebrados y peces en estos ecosistemas marinos en el Pacífico Norte de Costa Rica. Considerablemente más encuestas y trabajo de campo son requeridos para apoyar decisiones de manejo que sean respaldados por conocimientos científicos solidos.

Diversidad de los hábitats submareales de la Península de Santa Elena e Islas Murciélago, Pacífico Norte, Costa Rica / Subtidal habitats diversity of Santa Elena Peninsula and Murciélago Islands, North Pacific, Costa Rica

Introducción: Los hábitats costeros de todo el mundo están experimentando una presión cada vez mayor debido a la contaminación, el desarrollo costero, la pesca y el cambio climático. Identificar y registrar la biodiversidad costera es esencial para evaluar la salud, los cambios y el alcance de la pérdida de biodiversidad de los ecosistemas. Los hábitats costeros tropicales como los arrecifes de coral y los lechos de pastos marinos han sido el foco de investigación de los científicos durante las últimas décadas; sin embargo, se han descuidado otros ecosistemas, como los bosques de macroalgas, los fondos de sedimentos submareales (20-30 m de profundidad) y los arrecifes rocosos. Objetivo: Este trabajo reporta la biodiversidad marina asociada con arrecifes rocosos, arrecifes de coral, bosques de Sargazo y fondos submareales sedimentarios (20-30 m de profundidad), en un área tropical de afloramiento estacional (Península de Santa Elena, Bahía de Santa Elena e Islas Murciélago, Costa Rica). Métodos: Durante la Expedición Santa Elena (21 de abril-2 de mayo de 2018), se visitaron un total de 28 sitios para registrar la biodiversidad en cuatro hábitats diferentes (arrecifes rocosos y de coral, bosques de sargazo y fondos de sedimentos submareales), utilizando SCUBA, en muestreos sistemáticos, evaluaciones visuales y en algunos fondos de sedimentos se utilizó una draga de arrastre por 20 minutos. Resultados: Se identificaron un total de 254 taxones, siendo los peces óseos el grupo más diverso con 91 especies, seguido de los gasterópodos (25 spp.), las algas rojas (21 spp.) y los antozoos (19 spp.). Se reportan cuatro nuevos registros para el Pacífico continental de Costa Rica, incluida la subclase de una anémona tubícola (Ceriantharia). Conclusiones: Los resultados muestran que la península de Santa Elena, la bahía de Santa Elena y las islas Murciélago albergan hábitats poco comunes y no estudiados, como bosques del alga parda Sargassum, fondos de sedimentos, arrecifes construidos por el coral masivo Pavona gigantea, que es muy inusual, y comunidades de arrecifes rocosos dominadas por colonias calcáreas de poliquetos (Salmacina tribranchiata).

Introduction: Worldwide, coastal habitats are experiencing increasing pressure from pollution, coastal development, fisheries, and climate change. Identifying and recording coastal biodiversity is essential to assess ecosystem health, changes and the extent of biodiversity loss. Coastal tropical habitats such as coral reefs and seagrass beds have been the research focus for scientists during the last decades; however, other ecosystems have been neglected, such as macroalgae forests, subtidal (20-30 m deep) sedimentary bottoms, and rocky reefs. Objective: Our study reports the marine biodiversity associated with rocky reefs, coral reefs, Sargassum forests and sedimentary subtidal bottoms (20-30 m deep), in a tropical seasonal upwelling area (Santa Elena Peninsula, Santa Elena Bay, and Murciélago Islands, Costa Rica). Methods: During the 'Santa Elena Expedition' (April 21-May 2 2018), a total of 28 sites were visited in order to record the biodiversity across four different habitats (rocky and coral reefs, Sargassum forests, subtidal sedimentary bottoms), using SCUBA and both systematic surveys and visual assessments; in some sedimentary bottoms a 20-minute dredge tows were done. Results: A total of 254 taxa were identified, being bony fishes the most diverse group (91 species), followed by gastropods (25 spp.), red algae (21 spp.) and anthozoans (19 spp.). We report four new records for the Pacific mainland of Costa Rica, including the subclass of tube-dwelling sea anemone (Ceriantharia). Conclusions: Our results show that Santa Elena Peninsula, Santa Elena Bay, and Murciélago Islands harbor uncommon and unstudied habitats, such as Sargassum forests, sedimentary bottoms, reefs constructed by the stony coral Pavona gigantea -which is very unusual- and rocky reef communities dominated by calcareous colonies of polychaetes (Salmacina tribranchiata).

New hosts and morphological data for the star pearlfish Carapus mourlani (Ophidiiformes: Carapidae) from collections made in the North Pacific coast of Costa Rica / Nuevos hospederos y datos morfológicos para el Pez perla estrella Carapus mourlani (Ophidiiformes: Carapidae) a partir de recolectas realizadas al norte de la costa Pacífico de Costa Rica

Introduction: The family Carapidae includes about 40 species of marine fishes distributed in coastal habitats worldwide. The family includes some free-living species, however, most of them are found as commensal inquilines or parasites of marine invertebrates, including several echinoderm species. In the Eastern Tropical Pacific, the biology and host use of the representatives of the Carapidae is relatively poorly known. Objective: The present study reports the occurrence of the Star pearlfish Carapus mourlani within three previously unknown hosts in the region: the sea stars Nidorellia armata, Phataria unifascialis, and the sea cucumber Stichopus horrens. Some ecological implications and considerations regarding such symbiotic relationships are raised and discussed. Additional morphometric and meristic data for the fish and the echinoderms are also provided and discussed. Methods: Echinoderms were collected, from 25 localities along the North Pacific coast of Costa Rica, and were carefully examined searching for commensal/parasitic fishes. Echinoderms and fishes were identified and characterized in accordance with the specialized literature. Results: A total of 497 echinoderms, including about 60 species, were collected and examined. Commensal/parasitic fish (a single species represented by 13 specimens) were found in three echinoderm specimens/species. Conclusions: The list of echinoderm hosts for this carapid fish, through its whole distribution range, rises to 12 species (six sea stars and six sea cucumbers) and that could be a consequence of its wide geographic distribution, its generalist feeding habits and opportunistic commensal behavior.

Introducción: La familia Carapidae está compuesta por alrededor de 40 especies de peces marinos, presentes en hábitats costeros alrededor de todo el mundo. Se incluyen dentro de esta familia algunas especies de vida libre, no obstante, la mayoría de carápidos son inquilinos oportunistas o parásitos de algunos grupos de invertebrados marinos, incluyendo varias especies de equinodermos. En el Pacífico Tropical Oriental (PTO), se sabe relativamente poco sobre la biología de estos peces, así como de las diversas asociaciones existentes y de los hospederos utilizados. Objetivo: En este trabajo reportamos la ocurrencia del Pez perla estrella Carapus mourlani en tres nuevos hospederos: las estrellas de mar Nidorellia armata y Phataria unifascialis, y el pepino de mar Stichopus horrens. También se discuten algunas implicaciones y consideraciones ecológicas relacionadas a estas asociaciones simbióticas. Además, se proveen y discuten datos morfométricos y merísticos de los peces y sus hospederos. Métodos: Se realizaron recolectas de equinodermos, entre 2018 y 2019, en un total de 25 localidades distribuidas al norte de la costa Pacífico de Costa Rica, los cuales fueron cuidadosamente revisados en búsqueda de peces comensales/parásitos. Los equinodermos y los peces fueron identificados y caracterizados de acuerdo con la literatura especializada. Resultados: Se recolectaron y examinaron un total de 497 equinodermos, incluyendo alrededor de 60 especies, de los cuales solo tres individuos-especies estuvieron ocupados por peces comensales/parásitos. Conclusiones: La lista de hospederos equinodermos de C. mourlani a lo largo de su ámbito de distribución geográfico llega a 12 especies (seis estrellas de mar y seis pepinos de mar), lo cual podría ser un reflejo de su amplia distribución geográfica, de sus hábitos de alimentación generalistas y de su comportamiento oportunista en lo relativo al uso de hospederos.

Conservation status of fish and marine invertebrate of rocky reefs and sandy substrates in two unprotected bays of the Papagayo Gulf, Costa Rica / Estado de conservación de peces e invertebrados marinos de arrecifes rocosos y zonas arenosas en dos bahías desprotegidas del Golfo de Papagayo, Costa Rica

Introduction: North Pacific Costa Rica is generally considered rich in marine habitats and species. However, some areas are subject to fishing pressures that could degrade ecosystems. The Gulf of Papagayo comprises protected and unprotected areas with various degrees of impact. Objectives: Our aim was to evaluate the conservation status of fish and invertebrate communities in rocky reefs and sandy areas in two unprotected bays of the Gulf of Papagayo. Methods: The study was conducted at Cabuyal and Zapotillal Bays, south of Santa Rosa National Park. From December 2017 to April 2018, a total of thirty-five transects were done parallel to the coast at 3-10 m depth. We identified fish and invertebrate species and estimated biomass by trophic group in fish, and density in invertebrates. Results: We found a high number of species of fish (81) and invertebrates (70) in rocky reefs, which indicates a biodiverse ecosystem, but few species of fish in sandy areas. Species composition differed between the two bays. Density of juvenile fishes was high in Zapotillal Bay, suggesting that the area could be important for recruitment and breeding. Fish biomass was higher in sandy areas than in rocky reefs and all trophic groups had lower biomass than in other unprotected areas of the region. On average, reef fish biomass was 1.57 ± 0.67 (s.e.) t ha-1, similar to previous reports for Costa Rican North Pacific. By trophic group, piscivorous and planktivorous had the highest biomass in sandy areas and piscivorous and carnivorous in rocky reefs. Mean coral cover was low with 4.09 % ± 2.51 (s.e.), similar to previously reported for the region. Conclusions: Cabuyal and Zapotillal Bays in the Gulf of Papagayo are biodiverse in fish and invertebrates. However, the area may be impacted by local fisheries and other human activities, affecting fish and invertebrate populations, and coral formations. Our study fills some gaps in knowledge of marine biodiversity in the Gulf of Papagayo that can contribute to the conservation of marine life in North Pacific Costa Rica.

Introducción: El Pacífico Norte de Costa Rica generalmente se considera rico en hábitats y especies marinas. Sin embargo, algunas zonas están sujetas a presiones de pesca que pueden degradar los ecosistemas. El Golfo de Papagayo comprende zonas protegidas y desprotegidas con diversos grados de impacto. Objetivos: Nuestro objetivo fue evaluar el estado de conservación de las comunidades de peces e invertebrados en los arrecifes rocosos y las zonas arenosas de dos bahías desprotegidas del Golfo de Papagayo. Métodos: El estudio se llevó a cabo en las bahías de Cabuyal y Zapotillal, al sur del Parque Nacional Santa Rosa. Desde diciembre 2017 hasta abril 2018, se realizaron un total de 35 transectos paralelos a la costa entre 3-10 m de profundidad. Se identificaron especies de peces e invertebrados y se estimó la biomasa por grupo trófico en los peces, y la densidad en los invertebrados. Resultados: Encontramos un alto número de especies de peces (81) e invertebrados (70) en arrecifes rocosos, lo que indica un ecosistema biodiverso, pero pocas especies de peces en zonas arenosas. La composición de especies difirió entre las dos bahías. La densidad de peces juveniles fue elevada en la bahía de Zapotillal, lo que sugiere que la zona podría ser importante para el reclutamiento y la reproducción. La biomasa de peces fue mayor en zonas arenosas que en arrecifes rocosos y todos los grupos tróficos tuvieron menores biomasas que en otras zonas desprotegidas de la región. En promedio, la biomasa de peces de arrecifes fue de 1.57 ± 0.67 (s.e.) t ha-1, similar a lo reportado anteriormente en el Pacífico Norte de Costa Rica. Por grupo trófico, los piscívoros y planctívoros tuvieron la biomasa más alta en las zonas arenosas y los piscívoros y carnívoros en los arrecifes rocosos. La cobertura media de coral fue baja con 4.09 % ± 2.51 (s.e.), similar a la reportada anteriormente para la región. Conclusiones: Las bahías Cabuyal y Zapotillal en el Golfo de Papagayo son biodiversas en peces e invertebrados. Sin embargo, la zona puede verse afectada por las pesquerías locales y otras actividades humanas, lo que afecta a las poblaciones de peces e invertebrados y formaciones coralinas. Nuestro estudio llena algunos vacíos de conocimiento de la biodiversidad marina en el Golfo de Papagayo que pueden contribuir a la conservación de la vida marina en el Pacífico Norte de Costa Rica.

A synopsis of the genus Ethmia Hübner in Costa Rica: biology, distribution, and description of 22 new species (Lepidoptera, Gelechioidea, Depressariidae, Ethmiinae), with emphasis on the 42 species known from Área de Conservación Guanacaste.

We discuss 45 Costa Rican species of Ethmia Hübner, 1819, including 23 previously described: Ethmiadelliella (Fernald), Ethmiabittenella (Busck), Ethmiafestiva Busck, Ethmiascythropa Walsingham, Ethmiaperpulchra Walsingham, Ethmiaterpnota Walsingham, Ethmiaelutella Busck, Ethmiajanzeni Powell, Ethmiaungulatella Busck, Ethmiaexornata (Zeller), Ethmiaphylacis Walsingham, Ethmiamnesicosma Meyrick, Ethmiachemsaki Powell, Ethmiabaliostola Walsingham, Ethmiaduckworthi Powell, Ethmiasandra Powell, Ethmianigritaenia Powell, Ethmiacatapeltica Meyrick, Ethmialichyi Powell, Ethmiatransversella Busck, Ethmiasimilatella Busck, Ethmiahammella Busck, Ethmialinda Busck, and 22 new species: Ethmiablaineorum, Ethmiamillerorum, Ethmiadianemillerae, Ethmiaadrianforsythi, Ethmiastephenrumseyi, Ethmiaberndkerni, Ethmiadimauraorum, Ethmiabillalleni, Ethmiaehakernae, Ethmiahelenmillerae, Ethmiajohnpringlei, Ethmialaphamorum, Ethmiapetersterlingi, Ethmialesliesaulae, Ethmiaturnerorum, Ethmianormgershenzi, Ethmianicholsonorum, Ethmiahendersonorum, Ethmiarandyjonesi, Ethmiarandycurtisi, Ethmiamiriamschulmanae and Ethmiatilneyorum. We illustrate all species and their male and female genitalia, along with distribution maps of Costa Rican localities. Immature stages are illustrated for 11 species, and food plants are listed when known. Gesneriaceae is added as a new food plant family record for Ethmia. CO1 nucleotide sequences ("DNA barcodes") were obtained for 41 of the species.

Revision of the genus Pseudapanteles (Hymenoptera, Braconidae, Microgastrinae), with emphasis on the species in Area de Conservación Guanacaste, northwestern Costa Rica.

Pseudapanteles is a moderately diverse genus of Microgastrinae parasitoid wasps (Hymenoptera: Braconidae), endemic to the New World and with the vast majority of its species (including many undescribed) in the Neotropical region. We describe here 25 new species from Area de Conservación Guanacaste (ACG), northwestern Costa Rica, based on 400 studied specimens. A key to all 36 known species of Pseudapanteles is provided (except for Pseudapantelesbrunneus, only known from a single male), and species are placed in three newly created species-groups. Host records are known for only 25% of the species; most are solitary parasitoids of the caterpillars of several families of small Lepidoptera (Crambidae, Elachistidae, Gelechiidae, Incurvariidae, Sesiidae, Tineidae). DNA barcodes (part of the CO1 gene) were obtained for 30 species (83%), and provide a start for future study of the genus beyond ACG. Brief descriptions (generated by Lucid 3.5 software) and extensive illustrations are provided for all species. The following new taxonomic and nomenclatural acts are proposed: Pseudapantelesmoerens (Nixon, 1965), comb. n., Pseudapantelesbrunneus Ashmead, 1900, comb. rev., a lectotype is designated for Pseudapantelesruficollis (Cameron, 1911), and the following 25 species nova of Pseudapanteles (all authored by Fernández-Triana and Whitfield): alfiopivai, alvaroumanai, analorenaguevarae, carlosespinachi, carlosrodriguezi, christianafigueresae, hernanbravoi, jorgerodriguezi, josefigueresi, laurachinchillae, luisguillermosolisi, margaritapenonae, mariobozai, mariocarvajali, maureenballesteroae, munifigueresae, oscarariasi, ottonsolisi, pedroleoni, raulsolorzanoi, renecastroi, rodrigogamezi, rosemarykarpinskiae, soniapicadoae, teofilodelatorrei.

Review of Apanteles sensu stricto (Hymenoptera, Braconidae, Microgastrinae) from Area de Conservación Guanacaste, northwestern Costa Rica, with keys to all described species from Mesoamerica.

More than half a million specimens of wild-caught Lepidoptera caterpillars have been reared for their parasitoids, identified, and DNA barcoded over a period of 34 years (and ongoing) from Area de Conservación de Guanacaste (ACG), northwestern Costa Rica. This provides the world's best location-based dataset for studying the taxonomy and host relationships of caterpillar parasitoids. Among Hymenoptera, Microgastrinae (Braconidae) is the most diverse and commonly encountered parasitoid subfamily, with many hundreds of species delineated to date, almost all undescribed. Here, we reassess the limits of the genus Apanteles sensu stricto, describe 186 new species from 3,200+ parasitized caterpillars of hundreds of ACG Lepidoptera species, and provide keys to all 205 described Apanteles from Mesoamerica - including 19 previously described species in addition to the new species. The Mesoamerican Apanteles are assigned to 32 species-groups, all but two of which are newly defined. Taxonomic keys are presented in two formats: traditional dichotomous print versions and links to electronic interactive versions (software Lucid 3.5). Numerous illustrations, computer-generated descriptions, distributional information, wasp biology, and DNA barcodes (where available) are presented for every species. All morphological terms are detailed and linked to the Hymenoptera Anatomy Ontology website. DNA barcodes (a standard fragment of the cytochrome c oxidase I (COI) mitochondrial gene), information on wasp biology (host records, solitary/gregariousness of wasp larvae), ratios of morphological features, and wasp microecological distributions were used to help clarify boundaries between morphologically cryptic species within species-complexes. Because of the high accuracy of host identification for about 80% of the wasp species studied, it was possible to analyze host relationships at a regional level. The ACG species of Apanteles attack mainly species of Hesperiidae, Elachistidae and Crambidae (Lepidoptera). About 90% of the wasp species with known host records seem to be monophagous or oligophagous at some level, parasitizing just one host family and commonly, just one species of caterpillar. Only 15 species (9%) parasitize species in more than one family, and some of these cases are likely to be found to be species complexes. We have used several information sources and techniques (traditional taxonomy, molecular, software-based, biology, and geography) to accelerate the process of finding and describing these new species in a hyperdiverse group such as Apanteles. The following new taxonomic and nomenclatural acts are proposed. Four species previously considered to be Apanteles are transferred to other microgastrine genera: Dolichogenidea hedyleptae (Muesebeck, 1958), comb. n., Dolichogenidea politiventris (Muesebeck, 1958), comb. n., Rhygoplitis sanctivincenti (Ashmead, 1900), comb. n., and Illidops scutellaris (Muesebeck, 1921), comb. rev. One European species that is a secondary homonym to a Mesoamerican species is removed from Apanteles and transferred to another genus: Iconella albinervis (Tobias, 1964), stat. rev. The name Apanteles albinervican Shenefelt, 1972, is an invalid replacement name for Apanteles albinervis (Cameron, 1904), stat. rev., and thus the later name is reinstated as valid. The following 186 species, all in Apanteles and all authored by Fernández-Triana, are described as species nova: adelinamoralesae, adrianachavarriae, adrianaguilarae, adrianguadamuzi, aichagirardae, aidalopezae, albanjimenezi, alejandromasisi, alejandromorai, minorcarmonai, alvarougaldei, federicomatarritai, anabellecordobae, rostermoragai, anamarencoae, anamartinesae, anapiedrae, anariasae, andreacalvoae, angelsolisi, arielopezi, bernardoespinozai, bernyapui, bettymarchenae, bienvenidachavarriae, calixtomoragai, carloscastilloi, carlosguadamuzi, eliethcantillanoae, carlosrodriguezi, carlosviquezi, carloszunigai, carolinacanoae, christianzunigai, cinthiabarrantesae, ciriloumanai, cristianalemani, cynthiacorderoae, deifiliadavilae, dickyui, didiguadamuzi, diegoalpizari, diegotorresi, diniamartinezae, duniagarciae, duvalierbricenoi, edgarjimenezi, edithlopezae, eduardoramirezi, edwinapui, eldarayae, erickduartei, esthercentenoae, eugeniaphilipsae, eulogiosequeira, felipechavarriai, felixcarmonai, fernandochavarriai, flormoralesae, franciscopizarroi, franciscoramirezi, freddyquesadai, freddysalazari, gabrielagutierrezae, garygibsoni, gerardobandoi, gerardosandovali, gladysrojasae, glenriverai, gloriasihezarae, guadaluperodriguezae, guillermopereirai, juanmatai, harryramirezi, hectorsolisi, humbertolopezi, inesolisae, irenecarrilloae, isaacbermudezi, isidrochaconi, isidrovillegasi, ivonnetranae, jairomoyai, javiercontrerasi, javierobandoi, javiersihezari, jesusbrenesi, jesusugaldei, jimmychevezi, johanvargasi, jorgecortesi, jorgehernandezi, josecalvoi, josecortesi, josediazi, josejaramilloi, josemonteroi, joseperezi, joserasi, juanapui, juancarrilloi, juangazoi, juanhernandezi, juanlopezi, juanvictori, juliodiazi, juniorlopezi, keineraragoni, laurahuberae, laurenmoralesae, leninguadamuzi, leonelgarayi, lilliammenae, lisabearssae, luciariosae, luisbrizuelai, luiscanalesi, luiscantillanoi, luisgarciai, luisgaritai, luishernandezi, luislopezi, luisvargasi, manuelarayai, manuelpereirai, manuelriosi, manuelzumbadoi, marcobustosi, marcogonzalezi, marcovenicioi, mariachavarriae mariaguevarae, marialuisariasae, mariamendezae, marianopereirai, mariatorrentesae, sigifredomarini, marisolarroyoae, marisolnavarroae, marvinmendozai, mauriciogurdiani, milenagutierrezae, monicachavarriae, oscarchavesi, osvaldoespinozai, pablotranai, pabloumanai, pablovasquezi, paulaixcamparijae, luzmariaromeroae, petronariosae, randallgarciai, randallmartinezi, raulacevedoi, raulsolorsanoi, wadyobandoi, ricardocaleroi, robertmontanoi, robertoespinozai, robertovargasi, rodrigogamezi, rogerblancoi, rolandoramosi, rolandovegai, ronaldcastroi, ronaldgutierrezi, ronaldmurilloi, ronaldnavarroi, ronaldquirosi, ronaldzunigai, rosibelelizondoae, ruthfrancoae, sergiocascantei, sergioriosi, tiboshartae, vannesabrenesae, minornavarroi, victorbarrantesi, waldymedinai, wilbertharayai, williamcamposi, yeissonchavesi, yilbertalvaradoi, yolandarojasae, hazelcambroneroae, zeneidabolanosae.

Cryptic species within cryptic moths: new species of Dunama Schaus (Notodontidae, Nystaleinae) in Costa Rica.

Based on almost 1,700 recently reared and wild-collected specimens, the genus Dunama Schaus (Notodontidae, Nystaelinae) in Costa Rica is reviewed. Eight species are recorded of which seven are newly described: Dunama jessiehillae Chacón, Dunama jessiebarronae Chacón, Dunama janewaldronae Chacón, Dunama jessiebancroftae Chacón, Dunama janecoxae Chacón, Dunama biosise Chacón, Dunama indereci Chacón. Dunama angulinea Schaus is redescribed and associated with its correct genitalia. Dunama tuna (Schaus), previously listed as ocurring in Costa Rica, is restricted to Colombia. Most species are described through their distinctive CO1 barcodes, genitalia and life histories. Dunama adults and caterpillars, their foodplants, and their parasites in Area de Conservación Guanacaste (ACG) in northwestern Costa Rica are described where known. Many life history stages are illustrated.

A review of the New World species of the parasitoid wasp Iconella (Hymenoptera, Braconidae, Microgastrinae).

The New World species of Iconella (Hymenoptera: Braconidae, Microgastrinae) are revised. Iconella andydeansi Fernández-Triana, sp. n., Iconella canadensis Fernández-Triana, sp. n., and Iconella jayjayrodriguezae Fernández-Triana, sp. n., are described as new. Iconella isolata (Muesebeck, 1955), stat. r., previously considered as a subspecies of Iconella etiellae (Viereck, 1911), is here elevated to species rank. All species have different, well defined geographic distributions and hosts. Taxonomic keys are presented in two formats: traditional dichotomous hardcopy versions and links to electronic interactive versions (software Lucid 3.5). Numerous illustrations, computer-generated descriptions, distributional information, host records (mostly Lepidoptera: Crambidae and Pyralidae), and DNA barcodes (where available) are presented for every species. Phylogenetic analyses of the barcoding region of COI indicate the possibility that Iconella is not monophyletic and that the New World species may not form a monophyletic group; more data is needed to resolve this issue.

Time partitioning among jaguar Panthera onca, puma Puma concolor and ocelot Leopardus pardalis (Carnivora: Felidae) in Costa Rica's dry and rainforests / Separación temporal del jaguar Panthera onca, puma Puma concolor y ocelote Leopardus pardalis (Carnivora: Felidae) en los bosques tropicales húmedos y secos de Costa Rica

Abstract Segregation of daily activity patterns is considered an important mechanism facilitating the coexistance of competing species. Here, we evaluated if temporal separation existed among jaguar (Panthera onca), puma (Puma concolor) and ocelot (Leopardus pardalis) and if their activity patterns were related to that of a particular prey. We used camera trap records to estimate the activity schedules of these predators and their prey. We used the coefficient of overlapping (Δ; ranging from 0 to 1) to quantify the temporal interactions between predators and prey, and calculated confidence intervals from bootstrap samples. Strong temporal overlap occurred among the three felids (Δ = 0.63 - 0.82) in both dry and rainforests. However, a greater temporal separation was observed between the closest competitors (jaguar and puma, puma and ocelot). Jaguar and puma had a strong temporal overlap with medium and large-sized prey, while ocelots' activity matched that of smallsized prey. High overlapping coefficients among the felids suggest that temporal segregation is not the main mechanism facilitating their coexistence in these areas. However, fine-scale or spatiotemporal differences in their activity patterns might contribute to their coexistence in tropical environments.(AU)

Resumen La segregación de los patrones de actividad diaria es considerado un importante mecanismo para la coexistencia de especies competidoras. En este estudio, evaluamos si existía separación temporal entre el jaguar (Panthera onca), puma (Puma concolor) y ocelote (Leopardus pardalis), y si sus patrones de actividad se relacionaban con los de sus presas potenciales. Utilizamos registros de cámaras trampa para estimar los patrones de actividad de los felinos y sus presas. Usamos el coeficiente de traslape (Δ; de 0 a 1) para cuantificar las interacciones temporales entre depredadores y presas y calculamos los intervalos de confianza mediante bootstrapping. Observamos un fuerte traslape temporal (Δ= 0.63-0.82) entre los felinos tanto en el bosque seco como en el lluvioso. La mayor segregación temporal ocurrió entre los competidores más cercanos (jaguar y puma, puma y ocelote). El jaguar y el puma tuvieron un fuerte traslape con las presas medianas y grandes, mientras que el ocelote se traslapó con presas más pequeñas. Los altos coeficientes de traslape entre los felinos sugieren que la segregación temporal no es el mecanismo principal de coexistencia en estos sitios. Sin embargo, pequeñas diferencias en los patrones de actividad (e.g. picos de actividad desfasados) podrían contribuir a su coexistencia en ambientes tropicales.(AU)

Mariapanteles (Hymenoptera, Braconidae), a new genus of Neotropical microgastrine parasitoid wasp discovered through biodiversity inventory.

A new genus of microgastrine parasitoid wasps, Mariapanteles Whitfield & Fernández-Triana, gen. n., is described from rain forests of the Neotropics. The new genus is related to the common and speciose genus Pseudapanteles, but can be distinguished from the latter by having a complete transverse carina on the propodeum which forks around the spiracles. A molecular analysis based on data from COI from specimens of the proposed new genus plus possibly related genera confirms its generic distinctness. A key to two known species, Mariapanteles felipei Whitfield, sp. n. (Costa Rica) and Mariapanteles dapkeyae Fernández-Triana, sp. n. (Brazil) is provided. Evidence from collections suggests that there are other undescribed Neotropical congenerics. Specimens of Mariapanteles were likely confused in the past with the genus Beyarslania (referred to as Xenogaster until recently) but present information suggests that Beyarslania is restricted to the Afrotropical region while the Neotropical species clearly belong to a different genus, which we propose as new.

Actividades enzimáticas en aislamientos bacterianos de tractos digestivos de larvas y del contenido de pupas de Automeris zugana y Rothschildia lebeau (Lepidoptera: Saturniidae)

Enzymatic activities of bacteria isolated from the digestive tract of caterpillars and the pupal content of Automeris zugana and Rothschildia lebeau (Lepidoptera: Saturniidae). The enzymatic activities of bacteria isolated from the digestive tracts of caterpillars and the pupal contents of Automeris zugana and Rothschildia lebeau was studied. This digestive tract represents an extreme microenvironment due to its high pH and presence of antimicrobial substances secreted by the insect or derived from ingested plant tissue. At the same time, it contains large amounts of nutrient-rich food, for which microbes may compete among themselves and with the caterpillar. There is little information about the microbiota associated with tropical caterpillar guts, although bacteria from different genera have been isolated from gut and pupae samples. The study of the enzymatic activities generated by these organisms constitutes a starting point to understand their metabolic and physiological relationships with their hosts, and to find enzymes that have potential biotechnological applications. In this study we evaluated several enzymatic activities in two collections of bacteria isolated from caterpillar guts and pupae of the tropical lepidopteran species A. zugana and R. lebeau. Bacteria grown under aerobic conditions were tested for an array of enzymes, including gelatinases, caseinases, lipases, esterases, cellulases, xylanases, amylases and chitinases. Both collections displayed similar patterns of enzymatic activity. No isolate showed activity for all enzymatic tests, but as a whole, at least some bacteria in each collection were able to degrade each substrate tested. Isolates with the same taxonomic identification obtained from caterpillar guts and pupae had almost the same enzymatic activities. In both collections, it was possible to group bacterial isolates according to their enzyme activity pattern. In addition to a heterogeneous ensemble of isolates exhibiting two or less enzymatic activities, there were two groups with at least five activities that showed an apparent specialization for the substrates they were able to use. The first consisted exclusively of isolates of the family Enterobacteriaceae, which were positive for lipolytic and chitinolytic activities, but completely lacked amylasic, cellulolytic and xylanolytic activities. The second group, composed mainly of Gram-positive rods, exhibited the opposite pattern: they were positive for amylasic, cellulolytic and xylanolytic activities, lacked chitinolytic activity and had few isolates with lipolytic activity. This work forms the foundation for future research to explore the biotechnological potential of bacterial isolates from caterpillar guts. Rev. Biol. Trop. 55 (2): 401-415. Epub 2007 June, 29.

El tracto digestivo de orugas constituye un microambiente extremo, debido a su elevado pH y presencia de sustancias antimicrobianas secretadas por el insecto o derivadas del tejido vegetal ingerido. Al mismo tiempo, el intestino alberga gran cantidad de alimento, por el cual los microorganismos presentes podrían competir entre sí y con su hospedero. Existe poca información sobre la microbiota asociada con el intestino de orugas tropicales, aunque se ha demostrado la presencia de bacterias de diversos géneros tanto en el intestino como en el interior de pupas. El estudio de las actividades enzimáticas de estos microorganismos constituye un punto de partida en la comprensión de la posible relación metabólica y fisiológica que establecen con sus hospederos, a la vez que permite investigar enzimas con potenciales aplicaciones biotecnológicas. En este trabajo se evaluó la presencia de actividades gelatinolítica, caseinolítica, esterásica, lipolítica, quitinolítica, amilásica, celulolítica y xilanolítica en dos colecciones de aislamientos bacterianos provenientes de tractos digestivos de orugas y de pupas de los lepidópteros Automeris zugana y Rothschildia lebeau. Se utilizaron ensayos bioquímicos tradicionales para detectar enzimas secretadas en condiciones aerobias, en las que ambas colecciones exhibieron un comportamiento enzimático similar. Ningún aislamiento produjo un resultado positivo en todas las pruebas, pero como conjunto ambas colecciones fueron capaces de utilizar todos los sustratos evaluados. Los aislamientos obtenidos de pupas presentaron prácticamente las mismas actividades que sus homólogos provenientes de intestinos. En ambas colecciones fue posible agrupar los aislamientos de acuerdo con su patrón de producción de enzimas. Además de un conjunto heterogéneo de aislamientos poco activos (dos o menos actividades), se destacan dos grupos muy activos (al menos cinco actividades), que manifiestan una aparente especialización en los sustratos que utilizan. El primero de ellos está constituido exclusivamente por miembros de la familia Enterobacteriaceae, los cuales exhibieron un alto porcentaje de positividad en actividades lipolítica y quitinolítica, pero no demostraron la expresión de las actividades amilásica, celulolítica ni xilanolítica. El segundo grupo, formado en su gran mayoría por bacilos Gram-positivos, presenta la situación opuesta: alta positividad en actividades amilásica, celulolítica y xilanolítica, no detección de actividad quitinolítica y pocos aislamientos con actividad lipolítica. Este trabajo pretende ser la base de futuras investigaciones que exploren el potencial biotecnológico de aislamientos bacterianos provenientes del tracto digestivo de orugas.