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Revista de la Sociedad Entomológica Argentina

versão impressa ISSN 0373-5680versão On-line ISSN 1851-7471

Rev. Soc. Entomol. Argent. vol.78 no.1 La Plata mar. 2019



Aspects of the natural history of Phrynus barbadensis (Pocock, 1893) (Amblypygi: Phrynidae)

Aspectos de la historia natural de Phrynus barbadensis (Pocock, 1893) (Amblypygi: Phrynidae)


TORRES, Richard A.1,3*, de ARMAS, Luis F.2 & TOVAR-MÁRQUEZ, José1


1 Laboratório de Aracnologia, Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais. Belo Horizonte, MG, Brazil. * E-mail:
2 P. O. Box 4327. San Antonio de los Baños, Artemisa 38100, Cuba.
3 Grupo de Investigación en Zoología y Ecología, Universidad de Sucre. Colombia.


RESUMEN. La información sobre las presas y los depredadores de los amblipígidos es escasa, en gran parte dispersa y, a veces, olvidada por el aracnólogo. Phrynus barbadensis es una araña látigo neotropical ampliamente distribuida, pero su historia natural es poco conocida. El objetivo principal de esta contribución se refere a las presas, los depredadores, el esfuerzo reproductivo y las preferencias de microhábitats de P. barbadensis en una localidad colombiana. Dos investigadores llevaron a cabo el trabajo de campo durante cuatro expediciones de seis días cada una, entre junio y noviembre de 2017. Dos casos observados de depredación intragremial se relacionaron con la araña Scytodes longipes Lucas (Araneae: Scytodidae) y la araña látigo gigante Heterophrynus caribensis Armas, Torres-Contreras & Álvarez (Phrynidae: Heterophryninae). También se registra un nuevo caso de un molusco como presa de las arañas látigo. El número de embriones por lote en P. barbadensis fue 14-79 (n = 27, promedio = 40, error estándar = 3,83), teniendo una correlación positiva con el tamaño de la hembra. Esta especie estuvo altamente asociada a troncos caídos y rocas, pero pobremente a la hojarasca y las paredes rocosas (χ2 = 16,26; P < 0,001); con respecto a la temperatura del sustrato, mostró una asociación positiva. El canibalismo y la necrofagia entre las arañas látigo son brevemente comentadas, y también se proporcionan listas actualizadas de las presas y depredadores conocidos de los amblipígidos.

PALABRAS CLAVE. Ecología. Heterophrynus. Necrofagia. Scytodes. Sudamérica.

ABSTRACT. Information on preys and predators of the amblypygids is scarce, largely disperse and sometimes overlooked by the arachnologist. Phrynus barbadensis is a Neotropical whip spider widely distributed, but its natural history is poorly known. The main purpose of this contribution deals with preys, predators, reproductive effort and microhabitat preferences of P. barbadensis in a Colombian locality. Field work was carried out by two researchers during four expeditions of six days each, between June and November 2017. Two observed cases of intragremial predation concerned to the spitting spider Scytodes longipes Lucas (Araneae: Scytodidae) and the giant whip spider Heterophrynus caribensis Armas, Torres-Contreras & Álvarez (Phrynidae: Heterophryninae). A new case of a mollusk as prey of whip spiders is also recorded. The number of embryos per batch in P. barbadensis was 14-79 (n = 27, mean = 40, standard error = 3.83), having a positive correlation with the female size. This species was highly associated to fallen trunks and rocks, but poorly to the litter and rocky walls (χ2 = 16.26, P < 0.001); with respect to the substrate temperature, it showed a positive association. The cannibalism and necrophagy among the whip spiders are briefy commented, and updated lists of the known preys and predators of the amblypygids are also provided.

KEYWORDS. Ecology.Heterophrynus. Necrophagy.Scytodes. South America.




Amblypygi is a moderately diverse order of arachnids distributed in most tropical and subtropical terrestrial ecosystems. They have dorso-ventrally fattened bodies, raptorial pedipalps, and frst pair of elongate legs, mainly of sensorial function (Weygoldt, 2000; Chapin & Hebets, 2016). T o date, there are approximately 200 extant named species, which are included in 17 genera and 4 families (Maquart & Réveillion, 2016). Nevertheless, data on its natural enemies and preys is poorly documented (Weygoldt, 2000, Armas et al., 2013, Gonçalves-Souza et al., 2014; Torres-Contreras et al., 2015; Chapin & Hebets, 2016; Prous et al., 2017). Most of its known predators are scorpions and other whip spiders (intraguild competition and cannibalism), whereas the only true spider (Araneae) recorded as its predator is an undetermined Lycosidae (Chapin, 2011).

Under particular conditions, intraguild predation, including cannibalism, might play an important role in the trophic structure of those ecosystems in which the whip spiders are abundant (Chapin & Hebets, 2016). Although feld studies on this ecological aspect have not been conducted, sporadic observations seem to reinforce this hypothesis, but this does not constitute a high criterion.

Amblypygids, also known as whip spiders or tailless whipscorpions, are generalist that include several insects in their diet, mainly Orthoptera, Lepidoptera, and Dictyoptera (Chapin & Hebets, 2016), but also other preys, as freshwater shrimps, terrestrial mollusks, schizomids, scorpions, harvestmen, spiders, millipedes, centipedes, frogs and lizards, have been recorded. Owen & Cokendolpher (2006) reported a hummingbird (Apodiformes: Trochilidae) predated by the West Indian Phrynus longipes Pocock, but as it is unknown how the whip spider obtained that prey; necrophagy cannot be disregarded, because previous cases of this behavior have been observed in the group (Peck, 1974; Armas & Abreu-Collado, 1999; García-Rivera et al., 2009; Prous et al., 2017).

Phrynus barbadensis is a phrynid species well represented in the Colombian Caribbean (Chiriví-Joya & Armas, 2012). In recent years, some aspects of its natural history have been documented (Chiriví-Joya & Armas, 2012; Torres-Contreras et al., 2015). In the present contribution, additional preys and predators of this whip spider are documented, as well as data on its habitat preferences and reproductive biology.

Study site

The feld work was conducted in Montes de María (09° 31' N; 75° 20' W - 250 m a.s.l.), Colosó municipality, Sucre department, Colombia. This is a tropical dry forest, in a mountainous landscape with primary forest and several springs. The annual mean temperature is 26.8 °C, the precipitation varies between 1000 and 1200 mm and the relative humidity is 77% (Aguilera, 2005).


The feld work was carried out between 19:00 and 23:00 h by two researchers, during four expeditions of six days each, from June to November, 2017. To determine the reproductive effort, 27 ovigerous females o f P. barbadensis were collected. Four microhabitats (rocks, fallen trunks, crevices in rocky walls, and litter) were manually examined randomly searching for whip spiders. The sampling effort were distributed evenly with 20 minutes for each microhabitat. Temperature of the substratum was assessed by means a Thermoworks 24-inch waterproof digital thermometer.

All the specimens were preserved in ethanol 75%. Voucher specimens are deposited in the Zoological Museum of the University of Sucre, Colombia (MZUSU) with the collection number MZUSU-I00001, MZUSU-I00002, MZUSU-I00003, MZUSU-I00004, MZUSU-I00005. The specimens were collected within the framework permission for the collection of specimens of wild species of biological diversity for the purpose of scientific research granted by the National Environmental License Authority (ANLA) to the University of Sucre.

Statistical analyses

Carapace length was used as a measure of size of specimens and, a correlation analysis of Spearman was applied by means of the software PAST version 3.14 (Hammer et al., 2001).

To determine microhabitat preference, an analysis of binary logistic regression was applied in function of the substratum temperature, by means of the software SPSS version 22 for Windows (IBM, Armonk, NY, USA). A test of homogeneity (χ2) was applied to reject the null hypothesis of similar preference of microhabitat by this species, assuming that all the microhabitats had the same probability of being occupied.

Predation by Scytodes longipes (Fig. 1A, Table I)

On Oct 18th, 2017, a spitting spider Scytodes longipes (Araneae: Scytodidae) was found underneath a rock feeding on the whip spider P. barbadensis. The spider was an adult female (total length: 9.95 mm, prosoma length: 4.15 mm). The whip spider was an adult female (total length: 9.42 mm, carapace length: 3.64 mm). At the moment of the observation, the whip spider was entangled in the web and wrapped by silk on the pedipalps and prosoma, and the spider begun to feed on it.

Predation by Heterophrynus caribensis (Fig. 1B, Table I)

On Aug 26th, 2017, an adult female of H. caribensis (total length: 21.8 mm, carapace median length: 4.8 mm) was found on a rock eating a female P. barbadensis (total length: 15 mm, carapace median length: 2.6 mm). At the time of the observation, the predator strongly held the whole body of the prey while consuming its opisthosoma. The individual of P. barbadensis was still fresh from an ecdysis.

Predation on a gastropod mollusk (Fig. 1C, Table II)

On Sept 11th, 2017, an adult female of P. barbadensis (total length 15.59 mm, carapace median length 5.21 mm) was found under a stone eating a gastropod mollusk. At that moment, the amblypygid grasped with its pedipalps and chelicerae the soft corporal mass of the mollusk. The shell was empty, with a small rupture in its aperture. Measurements (in millimeters) of the shell were as follows: total length: 6.83, width: 4.64, aperture: 2.59 x 1.94.

Reproductive effort

The number of embryos per batch varied between 14 and 79 (n= 27, mean =40, standard error= 3.83), having a positive correlation with the female size: R2 = 0.881, P < 0.001 (Fig. 2A).

Microhabitat preference

Phrynus barbadensis was highly associated to fallen trunks (n = 17) and rocks (n = 15), but poorly represented in litter (n = 2) and rocky walls (n = 1) (χ2 = 16.26, df: 3, P < 0.001). With respect to the substrate temperature, P. barbadensis showed a positive association (Fig. 2B) (substratum temperature: coefficient B = -0.528, standard error = 0.166, P < 0.001; Constant: coefficient B = 15.077, standard error = 4.803, P < 0.001).


Of the recorded cases of whip spider predation (Table

I), three concern to cannibalism (Morales-Álvarez & González, 1986; Torres-Contreras et al., 2015), two events involve other whip spider species (Armas & Ramirez, 1989; this paper), and six concern predation by spiders and scorpions (Armas, 1995; Gering in Hebets, 2002; Chapin, 2011; Armas et al., 2013; Teruel & Toledo, 2014; this paper). Among vertebrates, Weygoldt (2000) indicated that large lizards, shrews, hedgehogs and mongooses are potential predators of whip spiders, but actually few cases have been recorded, mainly involving mammals and lizards (Dammerman, 1948; Armas, 1987; Stewart & Woolbright, 1996; Reid, 1997). As a whole, those cases refer to nine whip spider species (approximately 5% of the total), mostly belonging to Neotropical taxa of the genera Phrynus Lamarck (six species), Heterophrynus Pocock (two species) and Paraphrynus Moreno (one species), showing a clear panorama of the scarcity of the studies on this matter. Chapin & Hebets (2016: Table 3) erroneously included the small Puerto Rican frogs Eleutherodactylus coquiand E. richmondias predating o n P. longipes, but actually, the frogs were predated by the whip spiders (Formanowicz et al., 1981; Stewart & Woolbright, 1996).

Two cases of cannibalism inHeterophrynus guacharo Armas, were reported by Morales-Álvarez & González (1986), but they did not provide additional details. A similar situation occurs with the intraguild predations recorded by Armas (1995), Gering (in Hebets, 2002) and Chapin (2011). Based on the exiguous available data (Armas & Ramirez, 1989; Armas et al., 2013; Teruel & Toledo, 2014; this paper), only two preliminary conclusions may be addressed: (1) size differences determine amblypygid-amblypygid and other arachnid-amblypygid predation, being the larger individuals the predators; (2) as in other arthropods, the ecdysis and its immediately posterior process of sclerotization represent a hard phase in which the whip spiders are dramatically exposed to predators. Those conclusions are congruent with some results obtained by Polis (1981) and Polis & McCormick (1981) in several North American scorpions.

Certainly, intraguild predation and cannibalism offer remarkable opportunities to understand the dynamic relationships within an ecosystem. Nevertheless, as already pointed out, most existing information on whip spider's predators result of opportunistic feld observations, and quantitative assessments of their role as protagonist in the ecosystem trophic structures are lacking (Chapin & Hebets, 2016).


Amblypygids are generalist pantropical arachnids, seemingly opportunistic (Chapin & Hebets, 2016), but it is obvious that our present knowledge on its preys is meager. Remarkable or curious feld observations on the natural history are sometimes totally or partially omitted, mainly because the principal interest had been focused on the taxonomy. In the future, as a result of more detailed researches, perhaps some food items of the amblypygid diet might be not as uncommon as considered at this moment.


Fig. 1. Feed interaction in Phrynus barbadensis. A, predatation by Scytodes longipes and B, by Heterophrynus caribensis. C, eating a gastropod mollusc (Ampullaridae). Photos by Richard A. Torres


A single observation has been recorded for the following whip spider preys: centipedes, scorpions, short-tailed whip scorpions, termites and dipterans (Table II); nevertheless, several amblypygid species utilize termite nests as shelters and potentially as food source too (Weygoldt, 2000; Carvalho et al., 2011, 2012; Réveillion & Maquart, 2015; Chapin & Hebets, 2016). Shrimps, millipedes, harvestmen and Psocoptera are

known as regularly predated by one species each [according with Gil Wizen (pers. com., Nov 27th, 2017), Psocoptera are 1-2 mm long and preyed upon by the juvenile Charinus israelensis Miranda, Aharons, Gavish-Regev, Giupponi & Wizen, 2016, wandering in the bat guano, which they utilize as a hiding spot].

Crickets and cockroaches seem to be frequently predated by several amblypygid species, largely in caves, whereas moths have been reported as recurrent preys of two Neotropical Phrynidae (Beck & Görke, 1974; Hebets, 2002).


Table I. Known predators of the amblypygids Modifed from Chapin & Hebets (2016). Field observations only.





Varanus salvator (Laurenti) (Reptilia: Varanidae)

Catageus dammermani Roewer (Charontidae)

Dammerman (1948)

Solenodon cubanus Peters, 1861 (Mammalia: Soricomorpha)

Paraphrynus robustus (Phrynidae)

Armas (1987: 1)

Lophostoma silvicolum d'Orbigny, 1836 (Chiroptcra: Phyllostomidac)

Unidentified Phryninae [possibly P. barbadensis and/or Paraphrynus laevifrons (Pocock, 1894)1

Humphrey er a/. (1983:287)


Scytodes longipes Lucas, 1844 (Araneae: Scytodidae)

Phrynus barbadensis

This paper

Unidentified Lycosidae (Araneae)

Heterophrynus batesii (Butler, 1873) (Phrynidae)

Chapín (2011:3)

Heterophrynus guácharo Armas, 2015

Heterophrynus guácharo (cited as H. cervinus Pocock, 1894)

Morales-Álvarez & González (1986:73)

Heterophrynus caribensis Armas, Torres-Contreras & Álvarez, 2015

Phrynus barbadensis

This paper

Phrynus barbadensis

Phrynus barbadensis

Torres-Contreras et al. (2015)

Phrynus longipes (Pocock, 1893)

Phrynus hispaniolae Armas & Pérez, 2001 (cited as Ph. leviil)

Armas & Ramírez (1989: 3)

Alayotityus sierramaestrae Armas, 1973 (Scorpiones: Buthidae)

Phrynus damonidaensis Quintero, 1981

Armas et al. (2013)

Heteroctenus Junceus (Herbst, 1800) (Scorpiones: Buthidae)

Phrynus pinarensis FranganiLlo, 1930

Teruel & Toledo (2014)

Centruroides edwardsii (Gervais, 1843) [citedas C. margaritatus (Gervais, 1841)] (Scorpiones: Buthidae)

Phrynus whitei Gervais, 1842

Armas (1995: 2)

Unidentified scorpion (Arachnida: Scorpiones)

Phrynus pseudoparvulus Armas & Víquez, 2001 (cited as Phrynus parvulus Pocock, 1902)

E. Gering in Hebets (2002: 289)

Table II. Known preys of the amblypygids. Field observations only. Amblypygid-amblypygid predation is excluded (see Table I).





Eleutherodactylus coqui Thomas (Anura: Eleutherodactylidae)

Phrynus longipes (Pocock)

Formanowicz et al. (1981); Pfeiffe (1996: 266); Stewart & Woolbright (1996:308)

Eleutherodactylus richmondi Stejneger

Phrynus longipes

Stewart & Woolbright (1996: table 8.7)

Eleutherodactylus sp.

Phrynus sp. (seemingly Ph. pinarensis Franganillo)


Eleutherodactylus sp.

Phrynus sp.

Thomas in Stewart &. Woolbright (1996:308)

Pristimantis achatinus (Boulenger) (Anura: Craugastoridae)

Heierophrynus armiger Pocock, 1902


Anolis crysolepis Duméril & Bibron (Rcptilia: Dactyloidae)

Heterophrynus longicornis (Butler)


Anolis stratulus Cope (Reptilia: Dactyloidae)

Phrynus longipes

Reagan (1996:343, fig. 14.3); Thomas & Kessler (1996:355)

Anolis sp.

Phrynus longipes

Armas & Abreu-Collado (1999)

(Apodiformes: Trochilidae)

Phrynus longipes

Owen & Cokendolpher (2006), although does not determined whether it was a predation event or necrophagy.


Undetermined mollusks (Mollusca)

Heierophrynus guácharo (cited as //. cervinus)

Morales-Álvarez & González (1986:


Undetermined Arapullaridae (Mollusca: Gastropoda)

Phrynus harbadensis

This paper

Macrobrachium sp. (Decapoda: Palaemonidae)

Heterophrynus cheiracanthus (Gervais)

Ladle & Velander (2003)

Undetermined isopods (Isopoda: Oniscidea)

Heierophrynus guácharo (cited as H. cervinus); Charinus israelensis; Charinus ioanniticus (Kritscher)

Morales-Álvarez & González (1986: 73); Miranda et al. (2016:12); G. Wizen (pers. com., December 13* 2017)

Undetermined centipede (Chilopoda: Scolopendromorpha)

Phrynus longipes

Armas & Ramírez (1989: 3)

Undetermined millipedes (Diplopoda)

Phrynus pseudoparvulus

Hebets (2002: 289)

Stenochrus portoricensis Chamberlin (Schizomida: Hubbardiidae)

Phrynus marginemaculatus C. L. Koch

Armas (1989)

Centruroides gracilis (Latreille) (Scorpiones: Buthidae)

Paraphrynus cubensis Quintero

Forcelledo & Armas (2014)

Undetermined opilions (Arachnida: Opiliones)

P. pseudoparvulus

Hebets (2002: 289)


Table II (cont.). Known preys of the amblypygids Field observations only. Amblypygid-amblypygid predation is excluded (see Table I)



Referen ees

Nephila sp. (Araneae: Nephilidae)

Heterophrynus batesii

Chapín (2011: 3, as orbweaving spider); Chapin & Hebets (2016: Table 2, fig. 3b) identified it as Nephila sp.

Loxosceles sp. (Araneae: Sicariidae)

Charinus israelensis

Miranda et al. (2016: 12)

Undetermined spiders (Arachnida: Araneae)

P. pseudoparvulus

Hebets (2002: 289)

Undetermined Psocoptera (Insecta)

Charinus israelensis

Miranda et al. (2016: 12)

Diestrammena sp. (Orthoptera: Rhaphidophoridae)

Sarax yayukae Rahmadi, Harvey & Kojima, 2010 (Charinidae)

Rahmadi et al. (2010:9)

Amphiacusta sp. (cited as a cricket) (Orthoptera: Gryllidae)

Phrynus ¡ongipes

Armas (2010: 59, fig. 3 C)

Phalangopsis sp. (Orthoptera: Phalangopsidae)

Heterophrynus longicornis

Prous et al. (2017: 366-367, fig. 2A)

Undetermined crickets and katydids (Insecta: Orthoptera)

Phrynus pseudoparvulus; P. longipes; Heterophrynus guácharo

Hebets (2002: 289), Armas & Víquez (2001: 14); Peck (1974: 19,

as Tarántulafuscimana); Morales-Álvarez & González (1986: 73)

Aspiduchus cavernícola (Rehn) (Dictyoptera: Blaberidae)

Phrynus longipes

Moyá-Guzmán (2009: 74)

Undetermined cockroaches (Inserta: Dictyoptera)

P. pseudoparvulus; Heterophrynus guácharo

Hebets (2002: 289); Morales-Álvarez & González (1986: 73)

Undetermined termites (Inserta: Isoptera)

Charinus tomasmicheli Armas, (Charinidae)

Rodríguez-Cabrera & Teruel (2016: 69)

Undetermined moths, noctuids, sphyngids (Inserta: Lepidoptera)

Heterophrynus batesii (Butler); H. longicornis; P. pseudoparvulus

Beck & Gorke (1974); Carvalho et al. (2012: 1267); Hebets (2002: 289)

Undetermined Dolichopodidae (Insecta: Díptera)

Phrynus ¡ongipes

Moyá-Guzmán (2009: 74)


Spiders are mentioned as predated by three amblypygid species, but in the case of C. israelensis, only the immature individuals of Loxosceles spp. were consumed (G. Wizen, pers. com., Nov 27th, 2017).

Terrestrial isopods are recorded as eaten by three species (Table II), but C. israelensis was observed that predated on a small-sized isopod species (up to 7 mm in length) in all parts of the cave, mainly on the vertical walls; also, a single adult C. israelensis eaten on a larger isopod (15 mm in length), that seemingly entered the cave from the outside (G. Wizen, pers. com., Nov 27th, 2017).


Fig. 2. A) Positive correlation between the carapace median length (X axis) and the clutch size (Y axis). B) Model of microhabitat selection of Phrynus barbadensis in a Colombian locality, based on substrate temperature.


Mollusks have been recorded as predated by vinegaroons (Armas & Milera, 1989; Armas et al., 1989), harvestmen (Andre & Lamy, 1941), and spiders (Andre & Lamy, 1941; Fernández & Berovides, 1996), but rarely by scorpions (Lamoral, 1971; Alayón-García & Armas, 2010), and whip spiders. This is the second record of a mollusk as part of the diet of a whip spider (Table II)

Frogs and lizards seem to be trapped by largest amblypygids only (Table II). An interesting attempt of predation and active shunning of a dendrobatid poison frog by the Costa Rican Phrynus pseudoparvulus was recorded by Hovey et al. (2016).

With respect to the necrophagy, Peck (1974) reported it for P. longipes (that he identifed as Tarantula fuscimana) from Aguas Buenas Cave, Puerto Rico, but this interesting record has been overlooked by posterior researchers, excepting Moyá-Guzmán (2009). Armas & Abreu-Collado (1999) mentioned for the same species a similar case observed in a Hispaniolan cave [it was mentioned with some doubt by Armas & Pérez-Gonzalez (2001) and Armas (2006)]. A few years later, García-Rivera et al. (2009) recorded three specimens of the Cuban large amblypygid Paraphrynus robustus Franganillo, eating on two dead bats, and recently, Prous et al. (2017) reported bat necrophagy by the large South American amblypygid Heterophrynus longicornis Butler. Now, it is out of any doubt that necrophagy might be a frequent behavior in natural populations of some whip spiders.

Reproductive effort

Chiriví-Joya & Armas (2012) mentioned a Colombian female of P. barbadensis (carapace median length 7.3 mm) that carried 27 embryos, similar to what was found in this work. For this species, Quintero (1981) recorded 9 to 24 eggs or embryos per batch, but its correlation with any morphometric variable was not explored. Positive correlation of the clutch size and the female length was also found by Weygoldt (2000) and Armas & Pérez-González (2001) in three Caribbean Phrynus species, although a linear relationship is not always present (Weygoldt, 2000).

Microhabitat preference

High association of P. barbadensis with decaying trunks and rocks seems to be in correspondence with the usually humid and temperate condition of these microhabitats, excellent food source, availability of daytime shelters and appropriate places for reproduction. Although cannibalism occurs (Torres-Contreras et al., 2015), it is common to find several individuals in the same decaying trunk, usually interacting with each other.

Microhabitats as vertical structures (trees and rocky walls) are infrequently used by P. barbadensis, mainly because these sites are mostly occupied by H. caribensis, a potential predator.

Some species of the genus Phrynus are usually associated with large trees bearing buttresses (Hebets, 2002; Bloch & Weiss, 2002), but they are the biggest amblypygids in those places. Large South American species of the genus Heterophrynus also occur in similar microhabitats (Weygoldt, 1977; Dias & Machado, 2006; Chapin, 2014).



The authors are thankful to Gil Wizen (University of Toronto, Ontario, Canada) for bibliography, data on natural history of C. israelensis and C. ioanniticus, and his permission for publishing them. Thanks to Pedro Luis Atencia (Universidade Federal do Paraná, Brazil) and Jorge Arnaldo Díaz (Universidade Federal de Mato Grosso do Sul, Brazil) for his logistical support. Kenneth Chapin (University of California, Los Angeles), Abel Pérez González (National Museum of Natural History, Buenos Aires), Manuel Iturriaga Monsisbay (Instituto de Ecología y Sistemática, La Habana), Gabriel de los Santos (Museo Nacional de Historia Natural, Dominican Republic) and Carlos Víquez (INBio, Costa Rica), kindly provided some bibliography. T wo anonymous reviewers contributed with opportune suggestions to improve the manuscript.



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