Cyphoedma n. gen. now described after a century of use, with the addition of a new species from Central America (Geometridae, Ennominae)

ABSTRACT Cyphoedma n. gen., is validly described following a century of use as an unpublished manuscript name. Cyphoedma mirafloresa (Dognin, 1892) rev. stat. n. comb. is elevated to full species from subspecific status with Cyphoedma transvolutata (Walker, 1860) n. comb., and a third species in the genus is described from Central America: Cyphoedma ashleyorum n. sp. The adult habitus and male and female genitalia are illustrated for each of the three species and available COI (DNA) barcode data are reviewed.

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.

Patterns of vertebrate biodiversity in a tropical dry and mangrove forest matrix / Patrones de biodiversidad de vertebrados en una matriz de bosque seco tropical y manglar

Introduction: Tropical dry forests and mangroves, two of the world's most endangered ecosystems, each host a different set of environmental conditions which may support unique assemblages of species. However, few studies have looked at the unique vertebrate biodiversity in regions where both habitats occur side-by-side. Objective: To assess the vertebrate diversity and patterns of habitat usage in a mangrove and tropical dry forest matrix in an unprotected region of Northwestern Costa Rica. Methods: The study was conducted in a 7 km2 matrix of mangrove and tropical dry forests between Cabuyal and Zapotillal bays in Northwestern Costa Rica, south of Santa Rosa National Park. From September 2017 to March 2018, we used 13 automatic camera traps over 1 498 trap days to capture species utilizing the region and assess their patterns of habitat usage both spatially and temporally. Results: Seventy vertebrate species from 42 families in 27 orders were detected, including several globally threatened species. Over half of all species were detected in only one habitat, particularly amongst avian (78 %) and mammalian (42 %) species. Tropical dry forests hosted the greatest number of unique species and supported a greater percentage of herbivores than mangrove or edge habitats, which were dominated by carnivorous and omnivorous species. Mean detections per camera trap of all species increased significantly from the coldest and wettest month (Oct) to the hottest and driest months (Jan & Feb) in tropical dry forests. Sample-based rarefaction analysis revealed that survey length was sufficient to sample the tropical dry forest and edge habitats, though mangroves require further sampling. Conclusions: Taxa found to utilize different forest types may utilize each for different stages of their life cycle, moving between areas as environmental conditions change throughout the year. General patterns of global biodiversity favoring carnivore and omnivore usage of mangrove forests was confirmed in our study.

Introducción: Los bosques secos tropicales y los manglares, dos de los ecosistemas más amenazados del mundo, albergan cada uno un grupo de condiciones ambientales que pueden albergar conjuntos únicos de especies. Sin embargo, pocos estudios han analizado la biodiversidad única de vertebrados en regiones donde ambos hábitats se encuentran uno al lado del otro. Objetivo: Evaluar la diversidad de vertebrados y los patrones de uso del hábitat en una matriz de manglar y bosque seco tropical en una región no protegida del noroeste de Costa Rica. Métodos: El estudio se realizó en una matriz de 7 km2 de manglares y bosques secos tropicales en las bahías de Cabuyal y Zapotillal en el noroeste de Costa Rica, al sur del Parque Nacional Santa Rosa. De septiembre 2017 a marzo 2018, utilizamos 13 cámaras trampa automáticas durante 1 498 días trampa para capturar especies que utilizan la región y evaluar sus patrones de uso espacial y temporal del hábitat. Resultados: Se detectaron 70 especies de vertebrados de 42 familias y 27 órdenes, incluidas varias especies amenazadas a nivel mundial. Más de la mitad de todas las especies se encontraron en un solo hábitat, particularmente aves (78 %) y mamíferos (42 %). Los bosques secos tropicales albergan el mayor número de especies únicas y sustentan un mayor porcentaje de herbívoros que los hábitats de borde de manglares, que estaban dominados u hospedados por especies carnívoras y omnívoras. Las detecciones promedio por cámara trampa de todas las especies aumentaron significativamente desde el mes más frío y húmedo (octubre) hasta los meses más cálidos y secos (enero y febrero) en los bosques secos tropicales. El análisis de rarefacción basado en muestras reveló que la duración del estudio fue suficiente para muestrear los hábitats de bosque seco tropical y de borde, aunque los manglares requieren más muestreo. Conclusiones: Se encontró que los taxones pueden usar varios tipos de bosque en las diferentes etapas de su ciclo de vida, moviéndose entre áreas a medida que las condiciones ambientales cambian a lo largo del año. En nuestro estudio se confirmaron patrones generales de la biodiversidad global que favorecen el uso de los bosques de manglar por parte de carnívoros y omnívoros.

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.

Notas sobre los factores naturales y culturales en el desarrollo sociopolítico prehispánico en el extremo noroeste de Guanacaste, Costa Rica

Introducción: El estudio interdisciplinario de los patrones de ocupación y desarrollo de la sociedad humana en tiempos prehispánicos es tenue en la zona del volcán Orosí. El contexto ambiental es usualmente descrito para ilustrar patrones de subsistencia, sin mayor propósito de establecer formas de relación con los cambios que se vislumbran en la composición sociopolítica de los pobladores. Objetivo: En este trabajo, se explora la posibilidad de ilustrar instancias de relación entre los cambios que detecta el registro arqueológico, con potenciales factores naturales y culturales, según se destacan en los aportes de otras disciplinas científicas. Métodos: Una muestra de 111 componentes culturales identificados en sitios arqueológicos fue examinada en cuanto a su localización geográfica en diversos ecosistemas y su posición cronológica, respecto de los períodos culturales definidos en la región de Guanacaste. Resultados: Los componentes arqueológicos se confrontan con diversos factores naturales y culturales que habría incidido como agentes de cambio. Se analizaron las instancias de relación y se valoraron sus efectos por períodos, cubriendo tiempos precerámicos y cerámicos. Conclusiones: Se identifica como agentes naturales de cambio a la calidad de suelos, el tipo de clima, pero, ante todo, destacan los procesos culturales como el intercambio de productos, desarrollo de la agricultura y los movimientos migratorios de la población.

Introduction: Interdisciplinary studies on ancient occupation patterns and the development of human society in the Orosí Volcano region are few. Environmental data are included in archaeological studies of specific sites, mainly to illustrate the likely availability of local subsistence resources, with little regard for biodiversity linked to altitude and an array of different geomorphic areas. Effects surging from volcanic activity and tectonics are largely dispensed. Incidence of all these natural phenomena is rarely considered as agents of sociopolitical changes. Objective: Herein a search is performed in order to detect and illustrate instances of the relationship between sociopolitical change detected by archaeology and diverse natural and cultural factors as described by other scientific disciplines. Methods: A sample of 111 cultural components in archaeological sites were examined in terms of their geographical location, ecological context and chronological position along the periods of the cultural sequence in the Guanacaste region. Results: As settlement models, the archaeological data are confronted with diverse natural and cultural factors looking for their potential incidence as agents for sociopolitical change. Instances of relationship are examined and their apparent effects are evaluated for each period, both in preceramic and ceramic times. Conclusions: Both soil quality and climate are identified as important factors for change and development. Yet, largely salient are distinct cultural factors, namely the exchange of products, agriculture and migration processes.

New hosts and morphological data for the star pearlfish Carapus mourlani (Ophidiiformes: Carapidae) from collections made in the North Pacific coast of 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.

Naturaleza y estructura geológica del borde costero del noroeste de Costa Rica

Introducción: El noroeste de Costa Rica representa un área con una alta riqueza geológica que evidencia rocas generadas en diferentes ambientes, como plano abisal, talud continental, plataforma continental, volcánicos continentales efusivos y explosivos, volcánicos submarinos, costeros, erosivos fluviales, erosivos gravitatorios, de intrusión y el manto terrestre. Objetivo: Describir el contexto geológico actual del noroeste de Costa Rica, mediante el análisis y recopilación de datos de campo, geoquímicos, petrológicos, estructurales, paleontológicos y radiométricos con el fin de reconstruir los eventos y etapas de deformación desde el Jurásico a la actualidad. Métodos: Se hizo una recopilación bibliográfica de estudios en diversas ramas geológicas para establecer un estado del arte del extremo noroeste de Costa Rica. Resultados: La historia geológica representa el registro en rocas desde el Jurásico al Holoceno - actualidad con once etapas definidas a partir de interpretaciones petrológicas, estructurales, geocronológicas, estratigráficas y paleontológicas agrupadas en tres etapas de depositación y afectadas por cuatro fases tectónicas compresivas. Conclusiones: Las etapas de depositación corresponden con una primera etapa magmática con afinidad oceánica entre el Jurásico y el Cretácico Superior, una segunda etapa predominantemente sedimentaria con un rango de edad entre el Cretácico Superior y el Oligoceno, y por último una etapa volcánica efusiva - explosiva ubicada desde el Plioceno hasta la actualidad. Las fases tectónicas se asocian con diferentes eventos entre placas. La primera se dio en el Cretácico Inferior y su principal resultado visible fue la emersión de la Ofiolita de Santa Elena; la segunda sucedió del Cretácico Superior temprano al Campaniano y su reconocimiento está ligado a la posición actual de la Ofiolita de Santa Elena y sus estructuras circundantes; la tercera fase aconteció desde el Eoceno Superior hasta el Mioceno y su principal producto fue la generación del tren de pliegues sedimentarios del Golfo de Santa Elena a bahía de Salinas; y por último se presenta una fase Cuaternaria que originó el eje de basculamiento de los productos piroclásticos frente a la cordillera volcánica de Guanacaste.

Introduction: The Costa Rican northwestern coastline has a wide geological uniqueness that exhibits rocks from different environments such as abyssal plane, continental slope, continental platform, volcanic eruptions (effusive, explosive, and submarine), coastal, erosional (fluvial and gravitatory), intrusions, and upper mantle rocks. Objective: To present the geologic state of the art of the northwestern coastline of Costa Rica through the bibliographic review of geochemical, petrologic, structural, paleontological and geochronological data. Methods: A bibliographic revision was done to propose a state of the art of northwestern coastline of Costa Rica. Results: The geologic record shows eleven stages from the Jurassic to the Holocene. These stages were regrouped from petrologic, structural, geochronologic, stratigraphic and paleontological interpretations in three depositional stages and four compressive tectonic phases. Conclusions: The first depositional stage is volcanic with oceanic affinity between the Jurassic and Upper Cretaceous. The second has a sedimentary predominance with an age range between the Upper Cretaceous and the Miocene. The last depositional stage is a volcanic (effusive and explosive) from the Pliocene until the present. The tectonic phases are associate with different interactions between tectonic plates. The first phase triggered the Santa Elena Ophiolite obduction during the Early Cretaceous. The second phase occurred in the Early Upper Cretaceous - Campanian and is recognized by its current position and surrounding structures. The third phase lasted from the Upper Eocene until the Miocene and its main result was the folding of the Bahía de Salinas sedimentary rocks. Finally, the Quaternary phase created a tilt axis and the aperture for the deposition of pyroclastic density currents in front of the Guanacaste volcanic ridge.

Residency of bull sharks, Carcharhinus leucas (Carcharhiniformes: Carcharhinidae), at San Pedrillo Islet in Islas Murciélago, North Pacific of Costa Rica

Introduction: The bull shark, Carcharhinus leucas, is particularly vulnerable to anthropogenic actions because of its permanence in coastal ecosystems; populations depletion is registered in different places around the world. Aggregations of bull sharks have been reported in the North Pacific of Costa Rica, at Islas Murciélago, within the Guanacaste Conservation Area. Objective: To study the residency of bull sharks at San Pedrillo islet, Islas Murciélago. Methods: During the study period (June 2013 to February 2015) we used passive telemetry to tag 10 bull sharks. Results: All the sharks tagged were females, they were detected on 59 798 occasions by the acoustic receiver deployed in San Pedrillo. Acoustic signals from tagged sharks were received for a total period of 1 to 229 days (mean = 73.9 ± 71.3 days), with the last detections occurring on 9 January 2015. The Residency Index for each tagged shark across the entire monitoring period ranged from 0.41 to 1.00. The bull shark activity showed a significant pattern throughout the day at the receiver that specifically corresponded with the daily light cycle. Conclusions: This study concludes that San Pedrillo is an aggregation site (cleaning station) for bull sharks (C. leucas), possibly related to reproduction and not feeding behaviors.

Introducción: El tiburón toro (Carcharhinus leucas) es una especie tropical y subtropical que habita ríos, estuarios y aguas costeras. Objetivo: Estudiar la residencia del tiburón toro en San Perdillo, Islas Murciélago, Área de Conservación Guanacaste. Métodos: Durante el período de estudio (junio de 2013 a febrero de 2015) se marcaron 10 tiburones toro con telemetría acústica en San Pedrillo. Resultados: Los tiburones marcados fueron todos hembras y fueron detectados en 59 798 ocasiones por el receptor acústico desplegado en San Pedrillo. Las señales acústicas de los tiburones marcados se recibieron durante un período total de 1 a 229 días (media = 73.9 ± 71.3 días), y las últimas detecciones ocurrieron el 9 de enero de 2015. El índice de residencia de cada tiburón marcado durante todo el período de seguimiento osciló entre 0.41 y 1.00. La actividad del tiburón toro mostró un patrón significativo a lo largo del día en el receptor que se corresponde específicamente con el ciclo de luz diario. Conclusiones: Este estudio concluyó que San Pedrillo es un sitio de agregación (estación de limpieza) para el tiburón toro, posiblemente relacionado con comportamientos de reproducción y no de alimentación; y es un sitio crítico para la especie. Se necesitan más estudios para identificar áreas de alimentación y cría en el Área de Conservación de Guanacaste y áreas circundantes.

Diversidad de los hábitats submareales de la Península de Santa Elena e Islas Murciélago, Pacífico Norte, 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).

Checklist of invertebrates and conspicuous fishes in rocky reefs and Sargassum beds in the North Pacific of 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.

New species of Trichuris (Nematoda: Trichuridae) parasitizing Heteromys salvini (Rodentia: Heteromyidae) from Costa Rica, with a key to Trichuris species described from Heteromyidae / Nova espécie de Trichuris (Nematoda: Trichuridae) de Heteromys salvini (Rodentia: Heteromyidae) na Costa Rica, com uma chave para as espécies de Trichuris descritas em Heteromyidae

Abstract Trichuris guanacastei n sp., a parasite isolated from the Salvin' spiny mouse Heteromys salvini, collected from the Guanacaste Conservation Area, Costa Rica, during February 1996 is described. The new species was compared morphologically with the 29 known species that parasitize rodents distributed in 12 families in North and South America; T. guanacastei n. sp. it is characterized by the following set of traits: presence of a spicular tube (measuring 0.72-0.99); thick proximal cloacal tube and a short distal cloacal tube with a total length of 0.72-1.36; eggs 0.03-0.05 long and a semi-protrusible vulva. The new species represents the first one described in the genus in Costa Rica and the fifth one described in the Americas that parasite Heteromyidae.

Resumo O objetivo deste estudo foi descrever Trichuris guanacastei n. sp., um parasita isolado do rato espinhoso de Salvin, Heteromys salvini, coletado na Área de Conservação de Guanacaste, Costa Rica, durante fevereiro de 1996. A nova espécie foi comparada morfologicamente a 29 espécies conhecidas que parasitam roedores, distribuídas em 12 famílias na América do Norte e do Sul. T. guanacastei n. sp. é caracterizado pelo seguinte conjunto de características: presença de um tubo espicular (que mede 0.72-0.99); tubo proximal cloacal e um tubo distal cloacal com uma longitude total de 0.72-1.36; ovos de 0.03-0.05 de largura e uma vulva semi-protrusible. A nova espécie representa a primeira descrita do gênero na Costa Rica e a quinta descrita nas Américas que parasita Heteromyidae.

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.

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 new species of Trismegistomya Reinhard (Diptera: Tachinidae) from Area de Conservación Guanacaste in northwestern Costa Rica.

BACKGROUND: The New World genus Trismegistomya Reinhard, 1967b (Diptera: Tachinidae) previously included only the type species Trismegistomya pumilis (Reinhard, 1967a) from Arizona, U.S.A. NEW INFORMATION: We describe a new species of Trismegistomya, Trismegistomya jimoharai Fleming & Wood sp. n., from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica, reared from wild-caught caterpillars of Melipotis januaris (Guenée, 1852) (Lepidoptera: Erebidae). Our study provides a concise description of the new species using morphology, life history, molecular data and photographic documentation. In addition to the new species description, we provide a redescription of the genus, as well as of its type species Trismegistomya pumilis.

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.

Assessing regional drought impacts on vegetation and evapotranspiration: a case study in Guanacaste, Costa Rica.

This research investigates ecological responses to drought by developing a conceptual framework of vegetation response and investigating how multiple measures of drought can improve regional drought monitoring. We apply this approach to a case study of a recent drought in Guanacaste, Costa Rica. First, we assess drought severity with the Standard Precipitation Index (SPI) based on a 64-yr precipitation record derived from a combination of Global Precipitation Climatology Center data and satellite observations from Tropical Rainfall Measuring Mission and Global Precipitation Measurement. Then, we examine spatial patterns of precipitation, vegetation greenness, evapotranspiration (ET), potential evapotranspiration (PET), and evaporative stress index (ESI) during the drought years of 2013, 2014, and 2015 relative to a baseline period (2002-2012). We compute wet season (May-October) anomalies for precipitation at 0.25° spatial resolution, normalized difference vegetation index (NDVI) at 30-m spatial resolution, and ET, PET and ESI derived with the Priestley-Taylor Jet Propulsion Laboratory (PT-JPL) model at 1-km spatial resolution. We assess patterns of landscape response across years and land cover types including three kinds of forest (deciduous, old growth, and secondary), grassland, and cropland. Results show that rainfall in Guanacaste reached an all-time low in 2015 over a 64-yr record (wet season SPI = -3.46), resulting in NDVI declines. However, ET and ESI did not show significant anomalies relative to a baseline, drought-free period. Forests in the region exhibited lower water stress compared to grasslands and had smaller declines, and even some increases, in NDVI and ET during the drought period. This work highlights the value of using multiple measures to assess ecosystem responses to drought. It also suggests that agricultural land management has an opportunity to integrate these findings by emulating some of the characteristics of drought-resilient ecosystems in managed systems.

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)

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.

New species of Mutillid Mimicking Enoclerus Gahan (Coleoptera: Cleridae: Clerinae) from Mexico and Central America.

Six new species of Enoclerus are described: E. delusus n. sp. from Costa Rica and Panamá, E. incanus n. sp. from México, E. zip n. sp. from México and El Salvador, E. reductesignatus n. sp. from Panamá, and E. citrinifrons n. sp. and E. philogenes n. sp. from Costa Rica. The new species are hypothesized to participate in a Batesian mimicry complex with boldly marked diurnal mutillid wasps as their model. Enoclerus crabronarius var. deletus Wolcott is elevated to species rank as E. deletus Wolcott new status. The identity of E. crabronarius is discussed.