Comunicación breve
First report of an arboreal scorpion (Bothriuridae: Urophonius) from the temperate forests of southern Chile
Primer reporte de un escorpión arbóreo (Bothriuridae: Urophonius) en los bosques templados del sur de Chile
Jaime Pizarro-Araya1*, Andrés A. Ojanguren-Affilastro2 & Lorenzo Prendini3
1Laboratorio de Entomología Ecológica, Departamento de Biología, Facultad de Ciencias, Universidad de La Serena, Casilla 599, La Serena, Chile.
2División Aracnología, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Avenida Ángel Gallardo 470, CP: 1405DJR, CABA, Buenos Aires, Argentina.
3Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024-5192, USA. *E-mail: japizarro@userena.cl
We present the first observations of an arboreal bothriurid scorpion and the first report of an arboreal scorpion from the temperate forests of southern Chile. Urophonius tumbensis Cekalovic 1981 was observed and captured with ultraviolet (UV) light at a height of up to 6 meters on trunks of Cryptocarya alba (Molina) Looser trees in the Hualpén Botanical Reserve (Bío Bío Region, Chile).
RESUMEN
Presentamos las primeras observaciones de un escorpión bothriúridos arborícola y el primer registro de un escorpión arbóreo de los bosques templados del sur de Chile. Urophonius tumbensis Cekalovic 1981 fue observado y capturado mediante luz ultravioleta (UV) en troncos de árboles Cryptocarya alba (Molina) Looser a una altura de hasta seis metros en la Reserva Botánica de Hualpén (Región del Bío Bío, Chile).
The scorpion family Bothriuridae Simon 1880 has a Gondwana distribution pattern. Most bothriurid species occur in southern and central South America, from Ecuador to southern Patagonia, but there are also two genera in southern Africa and one in Australia (Prendini 2003). The distribution of Bothriuridae in South America is concentrated in arid areas, especially deserts, steppes and savannas (Prendini 2003; Agusto et al. 2006). In the tropical and subtropical forests north of 34°S, bothriurid scorpions are mostly replaced by scorpions of the families Buthidae C.L. Koch 1837 and Chactidae Pocock 1893. The few bothriurid species occurring in these forested habitats are obligate ground dwellers; the arboreal habitats are occupied by chactids and buthids (González-Sponga 1978). Maury (1982) suggested that the absence of bothriurid species in the seasonally flooded tropical rainforests of Amazonia is related to their dependence on the ground for excavating burrows and gestation chambers.
All species of Bothriuridae, for which data have previously been reported, are ground-dwelling scorpions. Most bothriurid species are fossorial, constructing burrows in the substratum, varying in depth from 30 cm to more than 1 m below the surface (Maury 1968, 1969; San Martin & Gambardella 1974; Ojanguren-Affilastro 2005). Many other bothriurids are opportunistic and will shelter under rocks or fallen trees, if available (lapidicolous sensu Prendini 2001).
Several species of different bothriurid genera have been observed at night, climbing on shrubs, bamboo and other low vegetation, usually no more than 30 to 50 cm above the ground. Bothriurids climb on low vegetation only to forage, and when disturbed, they drop immediately to the ground as an escape response. No bothriurid species was previously reported living or foraging in trees (arboreal or corticolous sensu Prendini 2001). Only Bothriurus pampa Ojanguren-Affilastro 2002 (cited as Bothriurus prospicuus Mello-Leitão 1934 by Maury [1973]) and Urophonius iheringi Pocock 1893 were mentioned to shelter under the bark of indigenous trees and introduced Eucalyptus trees (Maury 1968, 1973; Ojanguren-Affilastro 2002). However, we consider this behaviour to be opportunistic, because much of the area where these species occur is a treeless steppe. In the only native forests of the area, the dry "caldén" (Prosopis caldenia Burk) forests, where B. pampa also occurs, this species was not observed on trees at night with ultraviolet (UV) light detection, nor under tree bark during the day. We presume this is because the bark of P. caldenia is not as loose as that of Eucalyptus, making it unsuitable as a shelter.
The scorpion fauna of the humid temperate forests of southern Chile differs from that of the humid forests of Amazonia, in that only family Bothriuridae is present. Bothriurids are highly diversified in this area, which contains several endemic genera, e.g. Tehuankea Cekalovic 1973, Centromachetes Lonnberg 1897 and Phoniocercus Pocock 1893, and endemic species of more widespread genera, e.g. Bothriurus Peters 1861, Brachistosternus Pocock 1893 and Urophonius Pocock 1893 (Cekalovic 1973, 1974, 1981; Lourenço 2001; Mattoni 2002; Ojanguren-Affilastro et al. 2007). Despite the abundance of endemic bothriurid genera and species in the temperate forests of southern Chile, our knowledge of their biology remains fragmentary. Most species are known from only a few specimens and, in most cases, little or nothing is known of their distribution, ecology and phenology (Cekalovic 1966, 1968, 1976). No arboreal scorpions were previously reported from this area.
We recently conducted extensive fieldwork in southern Chile to survey its scorpion fauna. During two consecutive austral winters, August 2009 and July 2010, we visited the Hualpén Botanical Reserve (36°47״51.2׳S, 73°09״29.3׳W) on the Tumbes Peninsula near Concepción in the Bío Bío Region of southern Chile (Fig. 1), searching for a little known bothriurid, Urophonius tumbensis Cekalovic 1981. This species is closely related to the brachycentrus group of Urophonius, all species of which are active on the surface in winter (A.A. Ojanguren-Affilastro, J. Pizarro-Araya & L. Prendini, unpublished data). The Hualpén Botanical Reserve belongs to the "Bosque Caducifolio" botanical region, and its environment corresponds to a temperate to cold deciduous humid forest (Gajardo 1993).
]]>|
Figure 1. Map showing the known distribution of Urophonius tumbensis Cekalovic 1981 in southern Chile (white circles). Figura 1. Mapa que muestra la distribución conocida de Urophonius tumbensis Cekalovic 1981 en el sur de Chile (círculos blancos). |
During these expeditions, we confirmed that U. tumbensis has a winter activity period, and observed and collected several specimens active on trees at night, using UV light detection (Fig. 2a). All were collected from "peumo" trees, Cryptocarya alba (Molina) Looser (Fig. 2b), walking on the trunk and major branches at heights ranging from 2 to 6 m above ground. We cannot dismiss the possibility that specimens were present at even greater heights, because we could not see these scorpions (adult length ca. 3 cm) fluorescing above 5 to6 m. Several specimens of U. tumbensis were observed feeding on spiders, but they were too damaged to identify the species. One specimen was attempting to capture a spider (probably Ariadna sp.) in a small crevice covered by silk, by introducing its chelae and telson inside the spider retreat. Some of the scorpions observed with UV light, were partially hidden under loose bark or small epiphytes and the mud accumulated below them. When we attempted to capture the scorpions, they escaped by retreating into crevices in the bark or under the epiphytes, rather than dropping immediately to the ground as typically observed with ground-dwelling Urophonius species (Ojanguren-Affilastro et al. 2010). We collected several adult males and females of U. tumbensis, easily recognized by their sexual secondary characters typical of the genus (Ojanguren-Affilastro 2005); however no specimens were observed in courtship. Some juveniles were collected on the ground, but they represent only 10% of all specimens collected. It is not uncommon to occasionally find specimens of otherwise obligate arboreal buthid, chactid and liochelid scorpion species on the ground, elsewhere in the world (L. Prendini, unpublished data).
| Figure 2. a). Urophonius tumbensis Cekalovic 1981 sitting on "peumo" tree bark, taken at night under ultraviolet light. b) "Peumo" tree, Cryptocarya alba (Molina) Looser, in the Hualpén Botanical Reserve (Bío Bío region, Chile). ]]> Figura 2. a) Fotografía tomada con luz ultravioleta, de un ejemplar de Urophonius tumbensis Cekalovic 1981 sobre la corteza de un peumo. b) Árbol de peumo, Cryptocarya alba (Molina) Looser, en la Reserva Botánica Hualpén (región del Bío Bío, Chile). |
During daytime, we searched the peumo trees from which U. tumbensis were collected at night, but failed to find any specimens under the bark or epiphytes, or under stones, logs, or other possible retreats at ground level, in their immediate vicinity. As we were unable to find any specimens in their diurnal retreats, we cannot assume that U. tumbensis is not fossorial or lapidicolous like other species of the genus. We consider that unlikely, however, as the observation of some U. tumbensis retreating into bark crevices at night suggests that this microhabitat forms the diurnal retreat of some individuals.
The ground in the Hualpén Botanical Reserve is densely covered by low vegetation, but the broad trunks of peumo, lightly covered with moss and epiphytes, may offer a more suitable substrate on which to forage for prey. Other species of Urophonius have also been collected in forests: Urophonius transandinus Acosta 1998 and U. tregualemuensis Cekalovic 1981 in central Chile (Acosta 1998; Ojanguren-Affilastro et al. 2010; Ojanguren-Affilastro, Pizarro-Araya & Prendini, unpublished data), and U. brachycentrus (Thorell 1876) in central Argentina (Ojanguren-Affilastro 2005). However, the ground in these forests is less densely covered with vegetation, making it more suitable for ground-dwelling scorpions.
In the same area where we observed and collected U. tumbensis, we collected several specimens of two other bothriurid species in the genera Centromachetes and Phoniocercus under rocks and logs. We also collected some Phoniocercus specimens under the bark of a peumo and a willow (Salix sp.), about 2 m above the ground, one feeding on an Ariadna araucana Grismado 2008 spider. In addition, we collected some specimens of Centromachetes and Phoniocercus on the ground surface at night, with UV light detection. However, Centromachetes and Phoniocercus species do not appear to reach the peak of their surface activity in winter, so we prefer not to draw inferences about phenology based on these data alone.
This is the first time a bothriurid scorpion was observed using the microhabitats provided by trees. Prior to the present report, arboreal scorpion species were known only in the families Buthidae, Chactidae and Liochelidae Fet & Bechly 2001 (González-Sponga 1978; Prendini 2001, 2005). Future expeditions to the study area should identify the diurnal and summer retreats of U. tumbensis, and determine whether other scorpion species occupy the arboreal habitats during summer, when most Urophonius species are not active.
ACKNOWLEDGMENTS
We are grateful to Luis Piacentini, Eduardo Soto, Daniel Valdivia, Cristian Grismado, Fermín Alfaro, and Olivia Vergara for their help during fieldwork. We are indebted to Olivia Vergara, María Cecilia Jiménez, and Mauricio Cisternas for logistical support in Concepción; Cristian Grismado for identifying the spiders and spider retreats of the Hualpén Reserve; the authorities of the Universidad de Concepción for permits to collect specimens in the Hualpén Reserve; Luis Letelier for GIS assistance; an anonymous reviewer for comments on the manuscript. Fieldwork was funded by PICT 20071393 to Martín Ramírez, NSF DEB-0228699 to L. Prendini, NSF DEB-0613754 to N.I. Platnick, and DIULS-PF07101 of the Universidad de La Serena to J. Pizarro-Araya.
REFERENCES
Acosta, L.E. 1998. Urophonius transandinus sp. nov. (Bothriuridae), a scorpion from Central Chile. Studies on Neotropical Fauna and Environment 33: 157-164. [ Links ]
]]>Agusto, P., Mattoni, C.I., Pizarro-Araya, J., Cepeda-Pizarro, J. & Lopez-Cortes, F. 2006. Comunidades de escorpiones (Arachnida: Scorpiones) del desierto costero transicional de Chile. Revista Chilena de Historia Natural 79: 407-421. [ Links ]
Cekalovic, T. 1966. Contribución al conocimiento de los escorpiones chilenos. Noticiario Mensual del Museo Nacional de Historia Natural 10: 1-8. [ Links ]
Cekalovic, T. 1968. Alimentación y hábitat de Centromachetes pococki (Kraepelin, 1894) (Scorpionida, Bothriuridae). Boletín de la Sociedad de Biología de Concepción 40: 27-32. [ Links ]
Cekalovic, T. 1973. Tehuankea moyanoi n. gen. y n. sp. de escorpión Chileno, (Scorpiones, Bothriuridae). Boletín de la Sociedad de Biología de Concepción 46: 41-51. [ Links ]
Cekalovic, T. 1974. Dos nuevas especies del género Brachistosternus (Scorpiones: Bothriuridae). (Scorpionida, Bothriuridae). Boletín de la Sociedad de Biología de Concepción 47: 247-257. [ Links ]
]]>Cekalovic, T. 1976. Escorpiofauna del Parque Botánico Hualpén (Concepción, Chile) con la descripción de Bothriurus wilhelmi n. sp. (Arachnida, Scorpiones, Bothriuridae). Boletín de la Sociedad de Biología de Concepción 50: 173-181. [ Links ]
Cekalovic, T. 1981. Dos nuevas especies y un nuevo registro del género Urophonius para Chile (Scorpiones, Bothriuridae). Boletín de la Sociedad de Biología de Concepción 52: 195-201. [ Links ]
Gajardo, R. 1993. La vegetación natural de Chile, clasificación y distribución geográfica. Santiago, Chile. Editorial Universitaria. 165 pp. [ Links ]
González-Sponga, M.A. 1978. Chactas oxfordi (Scorpiones: Chactidae) nueva especie de la Sierra Nevada de Santa Marta, Colombia. Monografías Científicas Augusto Pi Suñer 9: 1-21. [ Links ]
Lourenço, W.R. 2001. On the taxonomy and geographic distribution of the genus Centromachetes Lonnberg, 1897, with a redescription of Centromachetespococki (Kraepelin, 1894) (Scorpiones, Bothriuridae). Entomologische Mitteilungen aus dem zoologischen Museum Hamburg 13: 305-313. [ Links ]
]]>Mattoni, C.I. 2002. La verdadera identidad de Bothriurus vittatus (Guérin-Méneville, 1838) (Scorpiones, Bothriuridae). Revue Arachnologique 14: 59-72. [ Links ]
Maury, E.A. 1968. Aportes al conocimiento de los escorpiones de la República Argentina. I. Observaciones biológicas sobre Urophonius brachycentrus (Thorell, 1877) Bothriuridae. Physis, Sección C 27: 407-418. [ Links ]
Maury, E.A. 1969. Observaciones sobre el ciclo reproductivo de Urophonius brachycentrus (Thorell 1877) (Scorpiones, Bothriuridae). Physis, Sección C 29: 131-139. [ Links ]
Maury, E.A. 1973. Los escorpiones de los sistemas serranos de la Provincia de Buenos Aires. Physis, Sección C 32: 351-371. [ Links ]
Maury, E.A. 1982. Dos Bothriurus del nordeste brasileño (Scorpiones, Bothriuridae). Revista de la Sociedad Entomológica Argentina 41: 253-265. [ Links ]
]]>Ojanguren-Affilastro, A.A. 2002. Descripción de Bothriurus pampa sp. n., con nuevas localidades para el grupo prospicuus (Scorpiones, Bothriuridae). Revista Ibérica de Aracnología 6: 95-102. [ Links ]
Ojanguren-Affilastro, A.A. 2005. Estudio monográfico de los escorpiones de la República Argentina. Revista Ibérica de Aracnología 11: 75-241. [ Links ]
Ojanguren-Affilastro, A.A., C.I. Mattoni & L. Prendini. 2007. The genus Brachistosternus in Chile (Scorpiones. Bothriuridae), with the description of two new species. American Museum Novitates 3564: 1-44. [ Links ]
Ojanguren-Affilastro, A.A., J.A. Ochoa, C.I. Mattoni & L. Prendini . 2010. Systematic revision ofthe granulatus group of Urophonius Pocock 1893 (Scorpiones, Bothriuridae) with description of a new species from central Chile. American Museum Novitates 3695: 1-40. [ Links ]
Prendini, L. 2001. Substratum specialization and speciation in southern African scorpions: the Effect Hypothesis revisited. In: Fet, V. & Selden, P.A. (eds.) Scorpions 2001. In Memoriam Gary A. Polis. British Arachnological Society, Burnham Beeches, Bucks, UK. pp. 113-138. [ Links ]
]]>Prendini, L. 2003. A new genus and species of bothriurid scorpion from the Brandberg Massif, Namibia, with a reanalysis of bothriurid phylogeny and a discussion of the phylogenetic position of Lisposoma Lawrence. Systematic Entomology 28: 149-172. [ Links ]
Prendini, L. 2005. Scorpion diversity and distribution in southern Africa: Pattern and process. In: Huber, B.A., Sinclair, B.J. & Lampe, K.-H. (eds.) African Biodiversity: Molecules, Organisms, Ecosystems. Proceedings ofthe 5th International Symposium on Tropical Biology, Museum Alexander Koenig, Bonn. Springer Verlag, New York. pp. 25-68. [ Links ]
San Martín, P.R. & Gambardella, L.A. 1974. Redescripción de Urophonius iheringi Pocock 1893 y consideraciones sobre morfología, bioecología y distribución. Boletín de la Sociedad de Biología de Concepción 47: 93-119. [ Links ]
Recibido: 05.10.10
Aceptado: 20.05.11
]]>