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Braz. J. Biol. vol.68 no.4 suppl.0 São Carlos Nov. 2008
Diversidade e distribuição dos Cyclopoida (Copepoda: Crustacea) de vida livre de água doce nos Neotrópicos
Departamento Ciências do Ambiente, Campus Pantanal, Universidade Federal de Mato Grosso do Sul UFMS, Av. Rio Branco, 1270, CEP 79304-020, Corumbá, MS, Brazil
Cyclopoida species from the Neotropics are listed and their distributions are commented. The results showed 148 species in the Neotropics, where 83 species were recorded in the northern region (above upon Equator) and 110 species in the southern region (below the Equator). Species richness and endemism are related more to the number of specialists than to environmental complexity. New researcher should be made on to the Copepod taxonomy and the and new skills utilized to solve the main questions on the true distributions and Cyclopoida diversity patterns in the Neotropics.
Keywords: Cyclopoida diversity, Copepoda, Neotropics, Americas, latitudinal distribution.
Foram listadas as espécies de Cyclopoida dos Neotrópicos e sua distribuição comentada. Os resultados mostram um número de 148 espécies, sendo que 83 espécies registradas na Região Norte (acima da linha do Equador) e 110 na Região Sul (abaixo da linha do Equador). A riqueza de espécies e o endemismo estiveram relacionados mais com o número de especialistas do que com a complexidade ambiental. Novos especialistas devem ser formados em taxonomia de Copepoda e utilizar novas ferramentas para resolver as questões sobre a real distribuição e os padrões de diversidade dos Copepoda Cyclopoida nos Neotrópicos.
Palavras-chave: diversidade dos Cyclopoida, Copepoda, neotrópicos, américas, distribuição latitudinal.
Cyclopoids are the most abundant and successful Copepoda in freshwater and they inhabit all kinds of freshwater environments such as rivers, streams, pools, lakes, reservoirs, wetlands and temporary pools. The Cyclopoida group commonly represents the main component of the zooplanktonic biomass in tropical water bodies (Rocha, et al., 1995; Santos-Wisniewski and Rocha, 2007).
The freshwater cyclopoids are distributed in two sub-families, Eucyclopinae and Cyclopinae. There are some genera with high diversity, some are distributed mainly in temperate zones such as Cyclops and Diacyclops and others in tropical zones represented principally Mesocyclops and Thermocyclops. In the Neotropical region, the tropical genera prevail, with some species different from those found in Africa and Australia, and there are also cosmopolitan while introduced species (Reid and Pinto-Coelho, 1994; Suárez-Morales et al., 1999).
Cyclopoida studies in the neotropical region are scarce and in all revisions genera new species have been added to the region lists (Gutierrez-Aguirre and Suárez-Morales, 2001; Suárez-Morales, et al., 2004; Silva and Matsumura-Tundisi, 2005). Revisions on the distribution of the Cyclopoida group in the neotropics have considered few genera, such as Thermocyclops (Reid, 1989) and Mesocyclops (Suárez-Morales and Gutierrez-Aguirre, 2001; Gutierrez-Aguirre and Suárez-Morales, 2001), and have been some local efforts such as Silva and Matsumura-Tundisi (2005) for Themocyclops in São Paulo State in Brazil and Suárez-Morales (2004) for Eucyclops in Mexico.
The aim of this work is to show the distribution of the free-living freshwater Cyclopoida group and its diversity in the neotropics, including South America, Central America and the Caribbean.
2. Material and Methods
The records of the Cyclopoida were obtained from a literature review on species in the world, in the neotropics and in some American countries and smaller localities, and from the Brazilian collections at UFSCar (Federal University of São Carlos, São Paulo State), IIE (International Institute of Ecology, São Carlos, São Paulo State), UFMS (Federal University of Mato Grosso do Sul, Mato Grosso do Sul State) and the Biota data bank (www.biota.org.br).
3. Diversity and Distribution
Table 1 shows the species and their distributions based on Figure 1 of the South and North neotropical regions. These results showed 148 species in the neotropics, where 83 species were recorded in the North region and 110 species in the South, these species being distributed in six genera of the sub-family Eucyclopinae and 14 genera of the sub-family Cyclopinae.
3.1. Genus recorded in the Neotropics
Sub-Family Eucyclopinae Kiefer, 1927
Macrocyclops Claus, 1903
Genus with three species in the neotropics, two cosmopolitan M. albidus and M. fuscus, and one neotropical M. albidus principalis. M. albidus is the most common and widely distributed species, from Argentina to Mexico (Dussart and Dafaye, 1985). It inhabits littoral zones of lakes and reservoir.
Eucyclops Claus, 1893
Genus with high diversity and with few studies on its taxonomy. E. serrulatus is the most common species being is described as a cosmopolitan and in the neotropics is distributed from Argentina to Mexico (Dussart and Dafaye, 1985; Rocha and Botelho, 1998; Suárez-Morales, 2004). Another 22 species present in the neotropical region are E. ensifer, E. leptacanthus, E. neotropicus, E. neumani neumani, E. neumani titicae, E. solii, E. solitarius, E. subciliatus, E. sp.¸ E. speratus, E. delachauxi, E. silvestrii, E. bondi, E. ariguanabensis, E. festivus, E. demacedoi, E. pseudoensifer, E. elegans, E. torresphilipi, E. conrowae, E. ariguanabensis, E. breviramatus.
Tropocyclops Kiefer, 1927
Genus that includes small organisms and the most common planktonic Cyclopoida species in the neotropics, with T. prasinus including T. prasinus prasinus (cosmopolitan), T. prasinus meridionalis and T. prasinus mexicanus (Reid, 1989; Rocha and Botelho, 1998). This species complex is distributed from Argentina to Mexico, T. prasinus meridionalis restricted to South America and T. prasinus mexicanus is restricted to Amazon, Central and North American sites. South America presents a high diversity with 10 endemic species, T.prasinus aztequei, T. prasinus peruviana, T. extensus, T.extensus longispina,T. schubarti, T. schubart dispar, T. rarus, T. federensis, T. nanae and T. piscinalis.
Paracyclops Claus, 1893
The genus Paracyclops has 28 species recorded in the world with eleven occurring in the neotropical region, and around 9 are American (Karaytug, 1999). In the neotropics the species P. poppei while P. chiltoni are considered cosmopolitan species, the first one being restricted to North America and the Caribbean zone and the second is restricted to South America and commonly misidentified as P. fimbriatus which does not exist in the Americas (Karaytug, 1999).
The other species recorded in the neotropics are P. pilosus, P. novenarius, P. andinus, P. carectum, P. rochai, P. reidae, P. hardingi, P. uenoi, P. puncatus.
Ectocyclops Brady, 1904
The genus Ectocyclops presents five species, one cosmopolitan, E. pharelatus and four species from South America and the Caribbean region, E. bromelicula, E. herbsti, E. rubescens and E. strenkzei (Dussart and Dafaye, 1985 and Rocha and Botelho, 1998).
Homocyclops (Herrick, 1882)
This genus is monospecific, with only one species, Homocyclops ater, which is an American species and its distribution in the neotropics is from Argentina to Mexico (Rocha and Botelho, 1998).
Sub-Family Cyclopinae Kiefer, 1927
Acanthocyclops G.O. Sars, 1863
This genus in the neotropics is represented by two cosmopolitan species and two endemic ones. A. robustus and A. vernalis are cosmopolitan and they are so close that they include a complex robustus-vernalis (Dodson, 1984). These species have been recorded in Eurasia, Australia and America. In the neotropics, they have been recorded from Argentina to Mexico (Reid, 1985; Dussart and Dafaye, 1985; Rocha and Botelho, 1998, Silva, 2003; Silva and Matsumura-Tundisi, 2002), and A. robustus is the most common form recorded in the neotropics. A. michaelseni and A. scottsbergi are both endemic to the south of Argentina and Chile.
Allocyclops Kiefer, 1932
Genus represented by two species in the neotropical region, one in the Cuba, A. botosaneanui (Dussart and Dafeyae, 1985) and another in Brazil, A. silvaticus (Rocha and Botelho, 1998).
Apocyclops Lindberg, 1942
Genus represented by four species in the neotropics (Dussart and Dafaye, 1985), A. panamensis (Central America and Caribbean), A. viduus (Aruba), A. procerus and A. distans (South America).
Bryocyclops Kiefer, 1927
The species from this genus are essentially Afro-Asian (Dussart and Dafaye, 1985), with two species in the neotropics restricted to South America, the species B. campaneri and B. Caroli.
Diacyclops Kiefer, 1927
The species of this genus is absent in most of the neotropical region, and are found mainly in the southern region of the Atlantic coast, in spite of this genus being the largest of the Cyclopidae group (Stoch, 2001). In the neotropics there are nine species, six of which are restricted to the north region: D. bernardi, D. hispidus, D. chakan, D. pilosus, D. ecabensis, D. puuc and three species restricted to the south above 40° S, D. andinus, D. bicuspidatus and D. urugaiensis.
Hesperocyclops Herbst, 1984
Hesperocyclops is an American genus, with two species, H. herbsti and H. improvisus (Rocha and Botelho, 1998).
Mesocyclops G. O. Sars, 1914
Most of the species from this genus have been recorded in the tropical region and it is one of the most studied groups, including revisions of African (Van de Velde, 1984), American (Suárez-Morales and Gutierrez-Aguire, 2001) and Australian (Holyska, 2000) species. Suárez-Morales and Gutierrez-Aguirre (2001) listed the 21 species and sub-species of Mesocyclops recorded in the neotropics, where one is pan-tropical, M. aspericornis, two neotropical M., M. longisetus longisetus and M. longisetus curvatus, two introduced species from the Afro-Asian region, M. ogunnus (the M. kieferi recorded in Brazil was a misidentification corrected by Matsumura-Tundisi and Silva, 2002 as M. ogunnus) and M. thermocyclopoides, and other species endemic or restricted to some regions of the neotropics such as M. venezuelanus, M. reidae, M. longisetus longisetus, M. longisetus curvatus, M. meridianus, M. pseudomeridianus, M. meridionalis, M. ellipticus, M. brasilianus (note: In the revision of the Mesocyclops species in São Paulo State and in Samuel Reservoir in Rondônia State in Brazil, M. meridianus was identified as M. brasilianus), M. edax, M. evadomingoi with six species are endemic to islands, M. cahci, M. yutsil, M. intermedius, M. pescei, M. annulatus diversus, M. longisetus araucanus and species from the South above 40° S M. annulatus annulatus and M. paranaensis.
Metacyclops Kiefer, 1927
The species from this genus present a high endemism and all species recorded in the neotropics are American species. In the neotropics 18 species were recorded including the endemic species M. subequalis, M. curtipinosus, M. hartmanni, M. paludolicola, M. paludolicola dentatus, M. oremaris, M. grandis, M. leptopus mucubaijensis, M. brauni, M. campestri, M. cushae, M. hirsutus, M. tredecimus and M. rudis. The most widely distributed Metacyclops species in the the neotropics are M. leptopus, M. mendocinus, M. laticornis, M. mendocinus venozuelanus, the latter one having been recorded in Venezuela and Brazil although in Brazil it is na incertae species (Rocha and Botelho, 1998).
Microcyclops Claus, 1893
The species of this genus present a low degree of endemism with many species recorded in more than one hydrographic basin. There are 12 recorded species in the neotropics (Dussart and Dafaye, 1985; Rocha, 1998; Rocha and Botelho, 1998): M. alius, M. anceps anceps, M. anceps pauxensis, M. anceps minor, M. ceibaensis, M. diversus, M. dubitailis, M. elongates, M. finitimus, M. furcatus, M. varicans subaequlis, M. varicans varicans.
Muscocyclops Kiefer, 1937
The species from this genus are all restricted to South America and are composed of three species: M. bidentatus, M. operculatus, M. therasiae. These species inhabit wet fields and organic soil (Rocha and Botelho, 1998).
Neutrocyclops (Lowndes, 1934)
This genus is monospecific, including only one species, N. brevifurca, distributed in the whole neotropical region.
Ponticyclops Reid, 1987
This genus is monospecific, including only one species, P. boscoi, recorded in the central Brazilian region.
This genus, one of the most common in the planktonic environment, in the neotropics consists of eleven species (Silva and Matsumura-Tundisi, 2005). Thermocyclops tenuis is the most widely distributed species in the neotropics, but many records are probably erroneous (Reid, 1989). Thermocyclops decipiens, a pan-tropical species, is the most abundant in the neotropics followed by T. minutus a South American species. T. inversus is a species distributed between tropics in the Americas. The other Thermocyclops species are more geographically restricted, being: T. brehmi, T. crassus, T. hastatus antilhensis, T. iguapensis, T. tenuis longifurcatus, T. parvus.
Yancyclops Reid, 1988
This genus is monospecific, including only one species, Y. ferrarii, recorded in the northern region of Brazil, in the Amazonian region.
The diversity and distribution of the Neotropical species is very likely linked to the pilocene events. Suárez-Morales (2004) observed that in the colonization of the Yucatán Peninsula in Mexico, South American species predominate. This way, the freshwater cyclopoid dispersion in the Neotropics should be starting from South American equatorial and tropical to the temperate zones and North America.
The distribution showed a high number of species of Cyclopoida in the Southern region. In fact, these results were expected, because the south is bigger than the north in area and environmental diversity. Seventeen species are present in the whole neotropical region, which represents 10.8% of the total species. The high number of endemic versus the low number of widely distributed species for Cyclopoid a has been found in all recent taxonomic studies such as Kiefer (1981); Van-de-Velde (1984); Reid (1989) and Karaytug (1999) and Suárez-Morales (2002); Suárez-Morales et al. (2004). This fact has high evolutionary significance, because it shows that the speciation of Cyclopida is bigger than supposed some time ago. Endemism in the neotropics should be much higher than recorded; nowadays endemism is directly linked to the amount of research carried out at a given site. Reid, (1987; 1989; 1991) for central Brazil, Holynska, 2000 for Australia and Suárez-Morales et al. (2004) for the Yucatán Peninsula (Mexico), Silva and Mastsumura-Tundisi (2005) for São Paulo State (Brazil) give examples of endemism and discuss its relationship with research.
Amazonian sites are examples of less-understood environments, with thousands of different freshwater environments, in a giant area certainly having dozens (a conservative estimate) of endemic and new species of freshwater Cyclopoida.
Moreover, even with the well-studied Cyclopoida groups, such as Mesocyclops, there are many doubts about distribution, because of imprecise descriptions or morphological confusion (Suárez-Morales, 2001). Nowadays, new skills have been useful in taxonomy; Silva and Matsumura-Tundisi, 2004, found differences between Mesocylops longisetus longisetus from the United States and M. longisetus longisetus from Brazil using cytogenetic techiniques. This kind of study helps to understand distribution patterns and the speciation process has occurred in the neotropics. In the latter there are three variations for M. longisetus: M. longisetus longisetus var. north, M. longisetus longisetus var. south and M. longisetus curvatus.
Silva and Matsumura-Tundisi (2005), using molecular biology, observed that the genera Themocyclops and Mesocyclops are closely related genera, and that the morphological differences are not supported. This result alerts evolutionary researchers that the study of these genera might not have to be done alone, but together, and their distribution might be interrelated.
On the other hand, human activities usually represent a strong forcing function driving distribution and diversity. Tundisi et al. (1995) discuss the formation of hundreds of hydropower dams in Brazil, which is a country poor in natural lakes, and that before hydropower, the Cyclopoida species was adapted in marginal lakes with strong interannual pulses. Nowadays, there are hundreds of cascades of interlinked reservoirs multiplying environments for plankton. Silva and Matsumura-Tundisi (2002) and Nogueira et al. (2004), observed that in the cascade systems, the Cyclopoida species are usually similar, differing in abundance among reservoirs, and that abundance is often related to the trophic state of the system and the retention time. This anthropic process might be homogenizing the systems and promoting extinction.
Concluding, in the last twenty years has been great improvement in the taxonomical knowledge of freshwater Cyclopoida with an emphasis on the impact of their distribution and diversity. Moreover, greater effort should be made to solve taxonomical doubts including the use of morphological, molecular and cytogenetic skills and in the geographical areas with a high deficit of studies.
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Received March 26, 2008
Accepted March 26, 2008
Distributed November 30, 2008