Diversity of the free-living marine and freshwater Copepoda (Crustacea) in Costa Rica: a review

The studies on marine copepods of Costa Rica started in the 1990’s and focused on the largest coastal-estuarine systems in the country, particularly along the Pacific coast. Diversity is widely variable among these systems: 40 species have been recorded in the Culebra Bay influenced by upwelling, northern Pacific coast, only 12 in the Gulf of Nicoya estuarine system, and 38 in Golfo Dulce, an anoxic basin in the southern Pacific coast of the country. Freshwater environments of Costa Rica are known to harbor a moderate diversity of continental copepods (25 species), which includes 6 calanoids, 17 cyclopoids and only two harpacticoids. Of the +100 freshwater species recorded in Central America, six are known only from Costa Rica, and one appears to be endemic to this country. The freshwater copepod fauna of Costa Rica is clearly the best known in Central America. Overall, six of the 10 orders of Copepoda are reported from Costa Rica. A previous summary by 2001 of the free-living copepod diversity in the country included 80 marine species (67 pelagic, 13 benthic). By 2009, the number of marine species increased to 209: 164 from the Pacific (49% of the copepod fauna from the Eastern Tropical Pacific) and 45 from the Caribbean coast (8% of species known from the Caribbean Basin). Both the Caribbean and Pacific species lists are growing. Additional collections of copepods at Cocos Island, an oceanic island 530 km away of the Pacific coast, have revealed many new records, including five new marine species from Costa Rica. Currently, the known diversity of marine copepods of Costa Rica is still in development and represents up to 52.6% of the total marine microcrustaceans recorded in the country. Future sampling and taxonomic efforts in the marine habitats should emphasize oceanic environments including deep waters but also littoral communities. Several Costa Rican records of freshwater copepods are likely to represent undescribed species. Also, the biogeographic relevance of the inland copepod fauna of Costa Rica requires more detailed surveys.     

A new species of Petasiger (Digenea: Echinostomiformes: Echinostomatidae) in the brown pelican, Pelecanus occidentalis, (Aves: Pelecaniformes: Pelecanidae), from the Area de Conservación Guanacaste, Costa Rica

A new species of Petasiger inhabits Pelecanus occidentalis, from the Area de Conservación Guanacaste, Costa Rica. The new species most closely resembles Petasiger novemdecim Lutz 1929 and Petasiger caribbensis Nassi, 1980 by having 19 circumoral spines and vitelline follicles confluent between the ventral sucker and gonads. The new species differs from both of these species by having a cirrus sac that is anteroposteriorly elongate and that reaches posteriorly to the midventral sucker, an ovary and Mehlis' gland that overlap the anterior testis dorsally, a uterus that lies dorsal to the ventral sucker, and a dextromedial genital pore. In P. novemdecim and P. caribbensis the cirrus sac is round and does not extend posteriorly to the anterior margin of the ventral sucker, both the ovary and Mehlis' gland are anterior to the anterior testis, the uterus runs lateral to and not dorsal to the ventral sucker, and the genital pore opens medially and sinistromedially, respectively.     

Some Entolomataceae (Agaricales) from Costa Rica

A preliminary account of Entolomataceae from Costa Rica is presented. Three new taxa are described, two inClitopilus, one inRhodocybe, and three new combinations are proposed inInocephalus. Five taxa ofClitopilus are reported for the first time from Costa Rica and a key to species is provided. Four species ofRhodocybe are discussed and a key to the six species known from Costa Rica is also provided.Alboleptonia earlei, Inocephalus murraii, Inocephalus quadratum, Rhodocybe incarnata andRhodocybe pseudonitellina are now known to occur in Costa Rica.     

Marine biodiversity of Costa Rica: Crustacea: Infraorder Anomura

The anomuran crabs are among the best known crustacean groups from the Pacific coast. However, this group is poorly known from the Caribbean coast of Costa Rica. In this compilation based on the literature and the collection at the Zoology Museum, Biology School, University of Costa Rica, we report the presence of 114 species of the Infraorder Anomura for Costa Rica, 20 species from the Caribbean, 96 species from the Pacific (two are present on both coasts). Twenty-nine species are new reports for Costa Rica, 15 from the Caribbean coast (74% of the total of species from that coast) and 14 from the Pacific (15% of the total from the Pacific). The range often species is extended to Costa Rica, siete from the Caribbean and three from the Pacific. Six species are reported for the first time from Cocos Island, where there are also four endemic species.     

Tinamutrema canoae n. gen. et n. sp. (Trematoda: Digenea: Strigeiformes: Brachylaimidae) in Crypturellus cinnamomeus (Aves, Passeriformes, Tinamidae) from the Area de Conservación Guanacaste, Costa Rica

We propose Tinamutrema as a new genus for Brachylaima centrodes (Braun, 1901) Dollfus, 1935 and for T. canoae, as a new species inhabiting tinamus in the Area de Conservación Guanacaste, Costa Rica. Specimens from Costa Rica resemble B. centrodes in having an elongate body, pretesticular genital pore and terminal genitalia, intercecal uterine loops occupying all available space between the anterior testis and the intestinal bifurcation, an oral sucker width:pharynx width ratio of approximately 1:0.55, an oral sucker:ventral sucker width ratio of approximately 1:1, and vitelline follicles extending into the forebody closer to the pharynx than to the anterior margin of the ventral sucker and by living in the cloaca. They differ from B. centrodes in having vitelline follicles that do not extend as far anteriorly as those in B. centrodes, which extend anteriorly to the level of the anteriormost extent of the cecal "shoulders," dense tegumental spination as opposed to sparse or no spination, relatively smaller cirrus with fewer spines, longer and more sinous pars prostatica, and forebody averaging 36% of total body length (TBL) as opposed to 42% TBL. Both species differ from other members of the Brachylaimidae in possessing a spinose cirrus and a cirrus sac containing both the cirrus and the pars prostatica. Preliminary phylogenetic assessment suggests that these traits are plesiomorphic, and thus the species are basal to the rest of the Brachylaimidae, whose diagnosis we emend accordingly.     

The classification, evolution and biology of the Costa Rican species of Cryptophion (Hymenoptera: Ichneumonidae)

The six known Costa Rican species of the campoplegine ichneumonid genus Cryptophion Viereck are described and keyed. The distribution of species throughout Costa Rica is detailed based on data gathered by an intensive Malaise trap survey of the ichneumonids of the country. Five new species are recognized: C. espinozai. C. guillermoi, C. manueli. C. moragai and C. tickelli , and a sixth, C. inaequalipes (Cresson) redescribed. The monophyly of the genus is demonstrated and the phylogeny of the Costa Rican species reconstructed. Host relationships have been established for all species in Costa Rica; they develop as koinobiont endoparasitoids of first to third instar larvae of Sphingidae or Saturniidae. Analysis of the host relationships from a phylogenetic perspective suggested that the genus first evolved using macroglossine sphingid larvae feeding on rubiaceous understorey plants as hosts, and subsequently diversified to utilize sphingine sphingids and ceratocampine saturniids feeding on a variety of food-plants. Most species appear to be monophagous and oligophagy is apparently a derived feature of one sister-species pair, C. espinozai and C. manueli . In Santa Rosa National Park, Costa Rica, only a small proportion of the species of Saturniidae and Sphingidae present are used as hosts by Cryptophion species. No one host species is parasitized by more than one Cryptophion species. No species of Cryptophion is known to parasitize more than one host species feeding on any one plant species. The monophagous species C. inaequalipes is only known to develop in its sphingid host when that host is feeding on one of its two alternative food-plants.     

Neohaematotrephus arayae n. sp. (Digenea: Echinostomiformes: Cyclocoelidae) in Jacana spinosa (Aves: Charadriiformes: Jacanidae) from the Area de Conservación Guanacaste, Costa Rica

Specimens of a species of cyclocoelid digenean inhabiting Jacana spinosa from the Area de Conservación Guanacaste, Costa Rica, most closely resemble Haematotrephus facioi (Brenes and Arroyo, 1962) Yamaguti, 1971, in the same host from Aranjuez, Puntarenas Province, Costa Rica, in having confluent vitelline follicles posteriorly, diagnostic of Neohaematotrophus, and in pharynx length, ovary width, and cirrus sac on the sinistral side. The new species is also highly similar in appearance to H. gendrei Dubois, 1959, also inhabiting a jacanid (from West Africa), which has vitelline follicles confluent posteriorly, and extending anteriorly to the intestinal bifurcation and genital pore opening immediately posterior to the anterior margin of the pharynx. Like H. facioi, H. gendrei has a relatively much shorter and broader cirrus sac than does the new species. Examination of the holotype and paratype of H. facioi confirmed that the specimens from Guanacaste differ in having a longer body, a larger ovary and eggs, and smaller testes. They also have the ovary on the sinistral rather than the dextral side of the body, genital pore anterior to the pharynx rather than at or posterior to the level of the posterior margin of the pharynx, longer and thinner cirrus sac, and eggs without eyespotted miracidia. Half the eggs in both specimens of H. facioi have well-developed eyespotted miracidia, whereas the typical condition for cyclocoelids is for virtually all eggs to exhibit eyespotted miracidia. Both H. facioi and H. gendrei are transferred to Neohaematotrophus, along with the new species.     

A new species of the genus Helcogramma (Blenniiformes, Tripterygiidae) from Taiwan

A new species of triplefin fish (Blenniiformes: Tripterygiidae), Helcogramma williamsi, is described from six specimens collected from southern Taiwan. This species is well distinguished from its congeners by possessing 13 second dorsal-fin spines; third dorsal-fin rays modally 11; anal-fin rays modally 19; pored scales in lateral line 22-24; dentary pore pattern modally 5+1+5; lobate supraorbital cirrus; broad, serrated or palmate nasal cirrus; first dorsal fin lower in height than second; males with yellow mark extending from anterior tip of upper lip to anterior margin of eye and a whitish blue line extending from corner of mouth onto preopercle. Comparisons and a diagnostic key are provided for the species of Helcogramma now known from Taiwan: Helcogramma fuscipectoris, Helcogramma inclinata, Helcogramma striata, Helcogramma trigloides, and the newly recorded, Helcogramma rhinoceros.     

A new species of snake of the genus Sibon (Serpentes: Colubridae) of the Caribbean slope of Costa Rica

Sibon lamari, a new species of colubrid snake from northeastern Costa Rica is described on the basis of six specimens. The new form differs from the closely allied S. annulatus in color pattern, head size, subcaudal count, and number of labials bordering eye-orbit. This is the third species of Sibon, together with S. argus and S. longifrenis, possessing a green dorsal ground color in Costa Rica. Sibon lamari coexists sympatrically with S. annulatus, S. argus. S. longifrenis and S. nebulatus at the locality of Guayacán de Siquirres (Limón Province).     

The genera of Chrysomelinae (Coleoptera: Chrysomelidae) in Costa Rica

Keys in Spanish and English are given for the genera of Chrysomelinae known from Costa Rica. For each genus, a list of species compiled from collections in the University of Costa Rica, the National Biodiversity Institute, and the entomological literature is presented. The genus Planagetes Chevrolat 1843 is recorded for the first time from Central America, and the genus Leptinotarsa St?l 1858 is synonymized with Stilodes Chevrolat 1843.     

New records of benthic marine algae and Cyanobacteria for Costa Rica,and a comparison with other Central American countries

We present the results of an intensive sampling program carried out from 2000 to 2007 along both coasts of Costa Rica, Central America. The presence of 44 species of benthic marine algae is reported for the first time for Costa Rica. Most of the new records are Rhodophyta (27 spp.), followed by Chlorophyta (15 spp.), and Heterokontophyta, Phaeophycea (2 spp.). Overall, the currently known marine flora of Costa Rica is comprised of 446 benthic marine algae and 24 Cyanobacteria. This species number is an under estimation, and will increase when species of benthic marine algae from taxonomic groups where only limited information is available (e.g., microfilamentous benthic marine algae, Cyanobacteria) are included. The Caribbean coast harbors considerably more benthic marine algae (318 spp.) than the Pacific coast (190 spp.); such a trend has been observed in all neighboring countries. Compared to other Central American countries, Costa Rica has the highest number of reported benthic marine algae; however, Panama may have a similarly high diversity after unpublished results from a Rhodophyta survey (Wysor, unpublished) are included. Sixty-two species have been found along both the Pacific and Caribbean coasts of Costa Rica; we discuss this result in relation to the emergence of the Central American Isthmus.     

A new species of Dicrotendipes (Diptera: Chironomidae) from Costa Rica

The adult male of Dicrotendipes baru is described from the Dominical area of southwestern Costa Rica. The species is characterized by its elongate club-like superior volsella. This species represents the third member of the genus known from Costa Rica.     

A new species of hognose pitviper, genus Porthidium, from the southwestern Pacific of Costa Rica (Serpentes: Viperidae)

A new species of terrestrial pitviper, Porthidium porrasi, is described from mesophytic forests of the Península de Osa and surrounding area of the Pacific versant of southwestern Costa Rica. It is most similar to P. nasutum and is characterized by a pattern of bands, persistence of the juvenile tail color in adults, and a high number of dorsal scales. Analysis of mtDNA sequences confirms its distinction from P. nasutum. The existence of this species reinforces the notion of elevated herpetofaunal endemism in southwestern Costa Rica.     

Presence of the dinoflagellates Ceratium dens, C. fusus and C. furca (Gonyaulacales: Ceratiaceae) in Golfo de Nicoya, Costa Rica

Harmful Algae Blooms (HAB) are a frequent phenomenon in the Gulf of Nicoya, Costa Rica, as in other parts of the world. The morphology and physiology of these microalgae are important because HAB species have adaptive characteristics. The production of high concentrations of paralytic toxins by Ceratium dinoflagellates has only been documented at the experimental level. However, this genus has been associated with the mortality of aquatic organisms, including oyster and shrimp larva, and fish, and with decreased water quality. Recently, fishermen reported massive mortality of encaged fish near Tortuga Island (Gulf of Nicoya). Samples were taken from an algal bloom that had produced an orange coloration and had a strong foul-smelling odor. Ultrastructural details were examined with scanning electron microscopy. The dinoflagellates Ceratium dens, C. furca and C. fusus were found in samples taken at the surface. The cell count revealed four million cells of this genus per liter. The morphological variability of these species is high; therefore electron microscopy is an useful tool in the ultrastructural study of these organisms. This is the first time that three Ceratium species are reported concurrently producing harmful blooms in Costa Rica.     

Whallwachsia illuminata n. gen., n. sp. (Trematoda: Digenea: Plagiorchiformes: Prosthogonimidae) in the steely-vented hummingbird Amazilia saucerrottei (Aves: Apodiformes: Trochilidae) and the yellow-olive flycatcher Tolmomyias sulphurescens (Aves: Passeriformes: Tyraninidae) from the Area de Conservación Guanacaste, Guanacaste, Costa Rica

A new species of digenean found in the intestines of the steely-vented hummingbird Amazilia saucerrottei and the yellow-olive flycatcher Tolmomyias sulphurescens from the Area de Conservación Guanacaste, Guanacaste, Costa Rica, resembles members of the Prosthogonimidae in having a highly lobate ovary; an elongate cirrus sac containing the cirrus, pars prostatica, and internal seminal vesicle; no external seminal vesicle; 2 fields of extracecal vitelline follicles restricted to the area between the intestinal bifurcation and testes; and uterine loops occupying all available space in the hind body. The new species differs from all other members of the family in having genital pores opening laterally to the cecum, immediately anterior to the acetabular level, and markedly oblique rather than symmetrical testes. Consequently, we propose the new genus Whallwachsia for the species. Preliminary phylogenetic assessment suggests that the species is the sister group of all other prosthogonimids.     

Review and redescription of species in the Oecetis avara group,with the description of 15 new species (Trichoptera,Leptoceridae)

The O. avara group of Oecetis is formally defined to include 4 described species, O. avara (Banks), O. disjuncta (Banks), O. elata Denning & Sykora, and O. metlacenis Bueno-Soria, and 15 new species. Oecetis marquesi Bueno-Soria, previously considered a member of the O. avara group, is treated as incertae sedis to species group, but is also redescribed and treated in the current work. New species described here (with their respective distributions) include: O. acciptrina (Costa Rica, Panama, Ecuador), O. agosta (Mexico), O. angularis (Guatemala to Ecuador), O. apache (SW USA), O. campana (Ecuador), O. constricta (Mexico to Ecuador, Venezuela, and Trinidad), O. houghtoni (North America), O. maritza (Costa Rica), O. mexicana (Mexico to Ecuador), O. patula (Guatemala, Nicaragua), O. protrusa (Mexico to Ecuador), O. sordida (Mexico, USA, Canada), O. tumida (Costa Rica), O. uncata (Costa Rica), and O. verrucula (Mexico to Costa Rica). A key to the species is also provided.     

Ashworthius Patriciapilittae n. sp. (Trichostrongyloidea: Haemonchinae), an abomasal nematode in Odocoileus virginianus from Costa Rica,and a new record for species of the genus in the Western Hemisphere

A species of Ashworthius is reported for the first time in the Western Hemisphere, and A. patriciapilittae n. sp. is described on the basis of specimens in white-tailed deer Odocoileus virginianus from Costa Rica. Among 8 known species, A. patriciapilittae is morphologically similar to A. tuyenquangi in red muntjac Muntjacas muntjak from northern Vietnam. The synlophe in A. patriciapilittae is composed of 26 ridges in the cervical zone and is continuous to the caudal extremity in males and females. Males are characterized by a complex dorsal ray and narrow trifurcate spicules (351-356 microm long) lacking an "eyelet." with dissimilar ventral and dorsal processes; the gubernaculum is 45-48% of the spicule length. Females have a prominent linguiform flap at the vulva and large eggs (108-142 microm long). The presence of A. patriciapilittae in Costa Rica is examined in the context of competing hypotheses for cospeciation or contemporary host-switching in cervids: either A. patriciapilittae is a component of an endemic Central and South American fauna that has diversified through coevolution of Ashworthius and cervid hosts or it has been introduced. Among haemonchines in the Western Hemisphere, specimens of A. patriciapilittae may be confused with 3 species of Haemonchus, including H. contortus, H. placei, and H. similis, that occur in both domestic and wild ruminants. Discovery of A. patriciapilittae emphasizes the continued need for survey and inventory to define the structure and distribution of parasite faunas in wild and domestic ruminants from the Nearctic and Neotropical regions.     

Widespread co‐occurrence of divergent mitochondrial haplotype lineages in a Central American species of poison frog (Oophaga pumilio)

Aim To analyse the phylogeographic structure of the strawberry poison frog, Oophaga pumilio (Dendrobatidae), across a large part of its range using a combination of mitochondrial and nuclear markers. Location Costa Rica and Panama. Methods Sequence analyses of a mitochondrial (cytochrome b) and a nuclear (RAG‐1) gene fragment as well as analyses of seven microsatellite loci were carried out on 269 individuals of O. pumilio sampled from 24 localities and on two individuals of O. vicentei. Results Two main mitochondrial haplotype lineages, corresponding to a northern (north Costa Rica) and a southern (south Costa Rica and eastern Panama) lineage, were identified. They differed by up to 7% uncorrected distance. We observed co‐occurrence of both lineages in seven populations in Costa Rica and Panama, indicating a pattern of extensive admixture. The two main mitochondrial lineages of O. pumilio roughly corresponded to a previously described phylogeographic pattern. Microsatellites indicate admixture spanning over a wide geographic area, but significant variation between the northern and southern groups was also found with microsatellite data. While microsatellite data reconstructed a separation south of an assumed Caribbean valley barrier, mitochondrial haplotypes of the ‘southern lineage’ shifted this barrier towards the north. Main conclusions Despite admixture, all three markers showed significant variation between the northern and southern groups. Phylogeographical breaks known from other anuran species in the study region could not be verified for O. pumilio. The unexpected clustering of the population from Escudo de Veraguas and the individuals of O. vincentei with the northern O. pumilio lineage indicates the need for a fundamental and careful taxonomic revision, including an interspecific phylogeography of the entire genus.