Two new species of Heterophrynus Pocock, 1894 from Colombia with distribution notes and a new ...

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Zootaxa 3647 (2): 329–342 ISSN 1175-5326 (print edition)
www.mapress.com/zootaxa/ Article ZOOTAXA
Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3647.2.5
http://zoobank.org/urn:lsid:zoobank.org:pub:43984C9D-C80D-4D1C-9737-4CB8879640C9
Two new species of Heterophrynus Pocock, 1894 from Colombia with distribution
notes and a new synonymy (Arachnida: Amblypygi: Phrynidae)
ALESSANDRO P. L. GIUPPONI1& 2 & ADRIANO B. KURY1
1-Departamento de Invertebrados, Museu Nacional/UFRJ. Quinta da Boa Vista, São Cristóvão, 20.940-040, Rio de Janeiro - RJ –
Brazil 2-Laboratório de Referência Nacional em Vetores das Riquetsioses, LIRN-FIOCRUZ. Manguinhos, 21040-360, Rio de Janeiro
–RJ – Brasil. (E-mail: [email protected])
Abstract
The genus Heterophrynus is for the first time recorded from Transandean areas. Heterophrynus boterorum sp. nov. and
Heterophrynus silviae sp. nov. are described respectively from Tolima and Valle del Cauca departments, Colombia, based
on material from the 2006 Arachnological Expedition of Museu Nacional to Colombia. Heterophrynus nicefori Amado &
Morales, 1986, from Meta department is newly considered a junior subjective synonym of Phrynus batesii Butler, 1873
(currently in Heterophrynus). Heterophrynus is currently known from Amazon forest, Brazilian Cerrado, Littoral Ridge
of Venezuela and Andean forests. A revised terminology is proposed for the constituent parts of male and female gonopods
of Heterophrynus.
Resumo
Este é o primeiro registro transandino do gênero Heterophrynus. Heterophrynus boterorum sp. nov. e Heterophrynus sil-
viae sp. nov. são descritos respectivamente dos departamentos de Tolima e Valle del Cauca, baseado em material da Ex-
pedição Aracnológica do Museu Nacional à Colômbia. Heterophrynus nicefori, do departamento de Meta é considerado
sinônimo júnior subjetivo de Phrynus batesii Butler, 1873 (atualmente em Heterophrynus). Heterophrynus é conhecido
atualmente de floresta amazônica, cerrado brasileiro, Cordilheira Costeira da Venezuela e florestas andinas. Uma nova
terminologia é proposta para as partes constituintes dos gonópodos masculino e feminino de Heterophrynus.
Introduction
Heterophrynus is a Neotropical genus of Phrynidae whip-spiders, with 11 species (Harvey 2002; Giupponi 2002)
not including one nomen dubium. Most species occur in the Amazon basin, and the genus has been said to be
“typical” of this biome (Weygoldt 2002), in spite of some literature records from Brazilian Cerrado (Mello-Leitão
1931; 1940; Lourenço & Hertault 1978); the Andean Ridge (Butler 1873; Pocock 1894; Pocock 1902; Amado &
Morales 1986) and Littoral Ridge of Venezuela (Harvey 2002).
The species of Heterophrynus are large-bodied with elongate appendages and occur mostly on large tree
trunks, gullies and boulders. The species of Heterophrynus present a retro-ventral apophysis on pedipalp trochanter
which is a synapomorphy for the genus (Weygoldt 1996), although no systematic value for use on species
relationships has been yet discovered. Their habits are mostly nocturnal (Weygoldt 2000). Mello-Leitão (1931)
provided a list of the species then known, describing two new ones and gave a key to the species. Two Brazilian
species have been redescribed by Weygoldt (1974). More recently, only one species has been described (Amado &
Morales 1986). A checklist of the species is given in Harvey (2002). Finally, two species H. longimanus Piza, 1939
and H. brevimanus Mello-Leitão, 1931 have been synonymized respectively with H. batesii (Butler, 1973) and H.
longicornis (Butler, 1973) (Giupponi 2002).
A revisionary project on Heterophrynus is currently being carried out in the Museu Nacional, including field
work in South America focusing in these animals. In the Colombia Expedition 2006, 15 specimens have been
secured, which are here described as two new species.
Accepted by L. Prendini: 11 Mar. 2013; published: 9 May 2013 329
Methods
The pedipalp article here called “tibia” is also sometimes referred to as “patella” in the literature. The former is
used in many recent works such as Giupponi & Miranda (2012) and Weygoldt (2000; 2002) while the latter is used
in Ramahdi et al. (2010; 2011).
The most widely used terminology for male and female genitalia in Phrynidae is that of Weygoldt (1974, 2000)
and Charinidae Weygoldt (1972) We here propose a new scheme, based on study of genitalia of several species, as
exemplified by the two new species and Brazilian species H. vesanicus (Figs 1–3). The male gonopod consists in a
set of more or less soft fleshy bodies which shape the spermatophore. Weygoldt recognized four structures, the
medial lobe = lm and dorsal lobe = ld, the lateral lobe = ll and the inner ventral medial lobe = lvl. Most material of
the gonopod is made by a paramedian pair of thick fleshy tubes with quite smooth surface, and which are slightly
depressed dorso-ventrally, not always straight, sometimes a little angular. These tubes are each here called fistula =
Fi. The median-dorsal part of each tube is projected forming a pair of triangular lobes (each here called lobus
dorsalis = LoD), while the median-ventral part is truncated. The fistulae are tipped by two telescoping softer
bodies, the basalmost, spongy and always covered with small hooks and denticles, is here called lobus lateralis
primus (= LoL1) and the distalmost (here called lobus lateralis secundus = LoL2) may be a spiny fleshy tongue (as
in H. boterorum) or a saccular set of concentric folds (as in H. silviae). More internally, each half of the fistula has
a stiff blade arising from its median-dorsal part, and here called processus internus (= PI) and partially surrounding
it, a leaf-like lamina (here called lamina medialis = LaM).
Abbreviations of the repositories used are:
BMNH—The Natural History Museum, London, England;
CAS—California Academy of Science, California, United States;
IBSP—Instituto Buntantan, São Paulo, Brazil;
ICN—Universidad Nacional, Instituto de Ciencias Naturales, Bogotá, Colombia;
MLS—Museo La Salle, Bogotá, Colombia;
MNRJ—Museu Nacional, Rio de Janeiro, Brazil;
MPEG—Museu Paraense Emilio Goeldi, Belém, Brazil;
MZSP—Museu de Zoologia, São Paulo, Brazil.
PUCRS—Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Brazil.
Systematic accounts
Heterophrynus Pocock
Heterophrynus Pocock 1894: 527.
Type species: Phrynus cheiracanthus Gervais, 1842, by original designation.
Included species. 1) Heterophrynus alces Pocock, 1902 (Brazil, Guyana, Suriname); 2) Heterophrynus armiger
Pocock, 1902 (Colombia, Ecuador); 3) Heterophrynus batesii (Butler, 1873) (Brazil, Colombia, Ecuador); 4)
Heterophrynus cervinus Pocock, 1894 (Colombia, Ecuador); 5) Heterophrynus cheiracanthus (Gervais, 1842)
(Colombia, Trinidad and Tobago, Venezuela); 6) Heterophrynus elaphus Pocock, 1903 (Peru); 7) Heterophrynus
gorgo Wood, 1869 (Peru); 8) Heterophrynus longicornis (Butler, 1873) (Brazil); 9) Heterophrynus seriatus
Mello-Leitão, 1939 (Brazil); 10) Heterophrynus vesanicus Mello-Leitão, 1931 (Brazil); 11) Heterophrynus
boterorum sp. nov. (Colombia); 12) Heterophrynus silviae sp. nov. (Colombia).
Heterophrynus batesii (Butler, 1873)
Phrynus batesii Butler 1873: 120–122, plate 6, figs 8–9.
Heterophrynus batesii: Pocock 1894: 287.
330 · Zootaxa 3647 (2) © 2013 Magnolia Press GIUPPONI & KURY
Heterophrynus nicefori Amado & Morales 1986: 33–40, figs 1–4, 7, 9. NEW SYNONYMY
♂
Type data. Phrynus batesii: 1 , 1 juvenile sy♂ntypes (BMNH, not examined), from [BRAZIL: Amazonas State],
“Upper Amazons”. Heterophrynus nicefori: 1 hotype (MLS, not examined) from “Colombia, Villavicencio”.
♂ ♀
Sample of material examined. BRAZIL; Acre; Rio Iquiri, Mun. Xapuri; (MZSP 09449); 1 ; 1 ; 1♂.ix.1951;
Exped. Permanente da Amazônia col. / Rio Branco, Reserva Extrativista de Catuaba; (IBSP 0057); 1 ; 1 juv.;
iv.1996; Equipe I♀nstituto Butantan col. / Amazonas; Usina Hidrelétrica de Balbina, Rio Uatumã (1°55'S 5♂9°28'W);
(IBSP 0117); 1 ; 1987; Equipe Instituto Butantan col. / Parque Nacional do Jaú; (IBSP 0069); 1 ; 1995;
♀ ♀
Oliveira, M. E. E. S. col. / (SMNS A 2579); 1 ; 08.vii.1977; R. M. Shelley col. / Aruanã; (PUCRS); 1 ;
2.xi.1991; Peralta, F. col. / Paraná do Urariá, Fortalezas; (MZSP 09458); 1 juv.; 24.ii.1972; Exped. Permanente da
♀
Amazônia col. / Ia♀uaretê; (IBSP 0042); 1 ; i.1971; Hoge, A. R. col. / Presidente Figueiredo, UHE Balbina;
(MNRJ 09003); 1 ; iv.1988; ELETRONORTE col. / Pará; Jurutí, drenagem do Igarapé Mutum (0236’10, 6’’ S,
♂
56º12’25,8’’ W) (Pitfall de répteis); (MPEG 0062); 5 ; 2 juv.; 02-20.ix.2002; T. C. A. Pire♀s et al. col. / Óbidos;
(MZSP 09457); 1 juv.; iii.1958; Oliveira, F. col. / Rondônia; Porto Velho; (MZSP 09452); 1 ; xi.1962; Herrmann
♂ ♀
col. / COLOMBIA; Amazonas; ♂Rio Apaporis Aná Pirá, 0º25' S, 70º15' W.; (CAS); 2 ; 1 ; 07-16.ii.1989; V. B.
Roth col. / (SMNS A 2577); 1 ; 08.vii.1977; J. W. Gentry col. / Meta; Villavicencio, Finca La Loma; (MNRJ
♂ ♀ ♂ ♀
9121); 6 ; 2 ; 1 juv.; viii.2006; A. Giupponi col./ Villavicencio, Restrepo; (MNRJ 9141); 2 ; 2 ; 2 juv.;
viii.2006; A. Giupponi col./ ECUADOR; Francisco de Orellana; Via Auca 55 km of Coca + 14km; (CAS); 1 juv.;
30.vi.03.vii.1996; A. I. Cognato col. / Napo; Mondaña; (CAS); 2 juv.; 25-29.vii.1996; A. I. Cognato col. / 20 Km
E. Puerto Napo, Aliñahui, 450 m, 1º0 S, 77º25 W.; (CAS); 1 ?; ii.1984; E. S. Ross col. / 20 Km E. Puerto Napo,
♀
Aliñahui, 450 m, 1. 0. S, 77. 25. W.; (CAS); 1 ; xi-xii.1995; E. S. Ross col. / 20 Km E. Puerto Napo, Aliñahui,
450 m, 1. 0. S, 77. 25. W.; (CAS); 1 juv.; xixii. 1995; E. S. Ross col.
Name formation. The specific name “batesii” with –ii is the correct Latin genitive of the form “Batesius”, as
used by Butler. The correction to “batesi” (as sometimes seen in the literature) with –i is an unjustified
emendation.
FIGURE 1. Male genitalia, colorized to show structures as defined here. See text for more details. Heterophrynus vesanicus
Mello-Leitão, 1931 male (MNRJ 9056) gonopod in apical view; Fi = fistula (gonopod tube); GO = genital operculum; LaM =
lamina medialis; LoD = lobus dorsalis; LoL1 = lobus lateralis primus ; LoL2 = lobus lateralis secundus; PI = processus internus
of gonopod.
TWO NEW HETEROPHRYNUS FROM COLOMBIA Zootaxa 3647 (2) © 2013 Magnolia Press · 331
FIGURE 2. Male genitalia, colorized to show structures as defined here. See text for more details. 2. Heterophrynus silviae sp.
nov., male (MNRJ 09115) paratype, gonopod in ventral view; 3. Heterophrynus boterorum sp. nov., male (MNRJ 09124)
paratype, gonopod in dorsal view. Fi = fistula (gonopod tube); GO = genital operculum; LaM = lamina medialis; LoD = lobus
dorsalis; LoL1 = lobus lateralis primus ; LoL2 = lobus lateralis secundus; PI = processus internus of gonopod.
Rationale of the synonymy. The original description of Amado & Morales does not contain any meaningful
evidence to keep H. nicefori as distinct from H. batesii. The whereabouts of the type specimens of Heterophrynus
nicefori are unknown, following a survey conducted in many Colombian potential repositories. However, material
from the type locality has been examined (see list above) and deemed conspecific with H. batesii.
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FIGURE 3. Heterophrynus batesii (Butler, 1873): comparison between specimens from Brazil and Colombia: a-c (MNRJ 9126
from Brazil, Pará) and d–g (topotype of H. nicefori; MNRJ 9121) from Colombia, Meta; Villavicencio; a. Male gonopod, left
side, ventral view; b. Same, detail of LoD apex; c. Same, detail of LoL 1 microsculpture; d. Male gonopod, left side, ventral
view; e. Same, detail of LoD apex; f. Same, detail of LoL 1 microsculpture; g. Female gonopod, general dorsal view.
Heterophrynus boterorum sp. nov.
Etymology. Species name is a tribute to our Colombian friends and colleagues Ricardo Botero Trujillo and Juan
Pablo Botero, who aid♂ed in many ways i♂n the discovery of this species.
Type material. 1 holotype and 1 paratype: COLOMBIA: Tolima, Ibague, viii-2006, Col., A. Giupponi, R.
♂ ♀
Botero e J. P. Botero (ICN). Other paratypes same data 1 and 1 MNRJ 09124.
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Diagnosis. Tibia I with 35 pseudoarticles and tarsus I with 79-80 pseudoarticles. Male gonopod: tegument of
lobus lateralis secundus with densely clustered small acuminate denticles and length almost equal to lobus lateralis
primus (Figs. 5b–c). Female gonopod: sclerotized claw bifid. Distalmost (third) inner tooth of basal article of
chelicera bifid. Small body size for the genus (Total body length: 23.0 mm).
Description of female (holotype): Carapace (Fig. 4a): flattened, wider than long; prominent eyes, median
eye tubercle dark-brown, without apical setae, lateral eye tubercle smaller, with around half its size. Margin of
carapace entirely with sinuous lines and coarse granules, anterior margin in a very smooth, practically straight
curve. Overall densely covered by small granules mostly anteriorly, bundles of bigger and more sparse granules
departing from the fovea towards the border. Frontal process well-developed, with greatly widened base and
abruptly getting thin in distal half, ending in a very acute hook, not exceeding front line of carapace, this way not
visible in dorsal view.
Sternum (Fig. 4b): Tri-segmented, typical of Phrynidae. Tritosternum extremely long projecting between
pedipalp coxae, reaching almost half of their length. Covered with bristles on its entire length, from apex to base,
with an increase in the density at base. Mesosternum convex, reduced to a small tubercle, ranging from slightly
oblong to sharply oblong, with six to eight bristles. Metasternum is a small convex mound with two bristles
Abdomen (Figs.4a–b): oblong, with almost indistinguishable punctuations, thinner than in the carapace.
Chelicera: Cheliceral furrow (Figs. 6a–b) with 3 internal teeth, the distal one bifid, the first cusp bigger than
the second one II<I=IIIa>IIIb. The third tooth (middle) much smaller than the others. With 3 external teeth, the
distal one bifid, the first cusp bigger than the second one II>I=IIIa>IIIb. Claw with five denticles, with the two
distalmost much reduced, the second smaller than first and third.
Pedipalp: Trochanter (Figs. 4c–d): with well-developed retro-ventral apophysis, armed with four spines, the
one adjoining the retro-ventral apophysis is the largest, but not more than twice the size of others, which are
subequal. The biggest spine is of comparable size to the fourth ventral spine of pedipalpal femur. Femur (Figs. 4c–
d): three dorsal spines in the main series (1>2>3), the first one three times bigger than the last one; prominent
geminate spines before the first spine, the dorsalmost subequal to spine 2 and the other subequal to spine 3; four
ventral spines (1>2>3>4) of approximately the same length of the dorsal ones; one spine placed proximally to the
first one, which is located more interiorly than the main series of spines, with a similar length of the third one, but
less robust; last spine ½ the length of the first one. Tibia (Figs. 4c–d): six spines, main series with four spines
(I=II=III>IV); two accessory spine anterior to spine I, with its length similar to spine IV; five ventral spines
(III>IV>II>I>V), Basitarsus (Figs. 4c–d): two dorsal spines and two ventral spines dorsal distal>dorsal
basal=ventral distal>ventral basal. The distal dorsal spine subequal to the claw. Claw (Figs. 4c–d): long, with an
acute, curved tip. The males do not exhibit considerable secondary sexual dimorphic features, and the pedipalps of
males and females are of comparable size.
Legs: All densely setose. Ventral corner of the prolateral face of femora II–IV projecting in a distinct spiniform
process. Femur length I>III>IV>II. Tibia I with 35 pseudoarticles and tarsus (basitarsus+distitarsus) I with 79-80
pseudoarticles in both legs. Leg IV: (Fig. 6c): Basitibia: 4 pseudoarticles, one trichobothrium at the first and one at
the last pseudoarticle. Distitibia: 16 trichobothria. Basitibia-distitibia length BT1>DT>BT3=BT4>BT2.
Distitarsus: with 3 pseudoarticles.
Measurements. Females (n=1): Total body length: 23.0, Cephalothorax: Length: 8.0 mm, Width: 10,5 mm.
Abdomen: 13.0 mm. Pedipalp: Femur 9.0 mm, Tibia 10.0 mm, Tarsus 5.0 mm, Tarsal claw 3.5 mm. Males (n=1):
Total body length: 24.2, Cephalothorax: Length: 10.0 mm, Width: 11,0 mm. Abdomen: 17.0 mm. Pedipalp: Femur
11.0 mm, Tibia 14.5 mm, Tarsus 5.1 mm, Tarsal claw 4.0 mm.
Color pattern (in alcohol): Chelicerae, pedipalps, legs and carapace reddish-brown. Coxae and ventral surface
of abdomen lighter colored.
Genitalia: Male (Figs. 5a–f). PI longer than LaM. LoL1 and LoL2 subequal in length, both densely covered
with minute denticles (but larger than in H. silviae). In dorsal view the LoD is subequal in size to LoL1. Female.
(Fig. 6d) Gonopod claws bifid. Inner branch probably arising from basal third of claw complex.
Natural history. The specimens described were collected on rocks and ravines in a fragment of Andean
montane secondary forest in a very disturbed area surrounded by coffee plantations and rural residential areas.
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FIGURE 4. Heterophrynus boterorum sp. nov., male (MNRJ 09124) paratype: a. Habitus, dorsal view; b. Same, ventral view;
c. Right pedipalp, dorsal view; d. Right pedipalp, ventral view. Scale bars = 5 mm.
Heterophrynus silviae sp. nov.
Etymology. Species name honors the Colombian arachnologist Silvia Vanegas, who paved the way for the 2006
Colombian expeditio♂n. ♀
Type material: holotype, and juv. paratypes: COLOMBIA: Valle del Cauca, Dagua, El Salto, 13-viii-
♂ ♀
2006, Col., A. Giupponi, S. Vanegas e C. Prieto (ICN). Other paratypes 6 , 1 and 1 juv., same data (MNRJ
09115).
TWO NEW HETEROPHRYNUS FROM COLOMBIA Zootaxa 3647 (2) © 2013 Magnolia Press · 335
Diagnosis. Tibia I with 29 pseudoarticles and tarsus I with 67 pseudoarticles. Male gonopod: tegument of
lobus lateralis secundus with texture of concentric ridges, resembling a Pycnoporus mushroom or a fingerprint
(Figs. 8b–c). Female gonopod (Figs. 8e-f) with sclerotized claws with basal portion wide, getting abruptly thin in
the distal third, strikingly similar to the female gonopods of Phryninae. Distalmost (third) inner tooth of basal
segment of chelicera bifid (Figs. 9a-b). Median body size for the genus (total body length: 28.0).
FIGURE 5. Heterophrynus boterorum sp. nov., male (MNRJ 09124) paratype, genitalia and surroundings: a. Male gonopod,
general ventral view; b. Same, detail of left and right lobes; c. Same, detail of left distal part of lateral lobe (LoL2), spiny-
tongue-like with only one layer covered with denticles; d. Same, detail of basal part of Lol2; e. Male gonopod, general dorsal
view; f. Same, detail of right dorsal and lateral lobes.
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Description of female (holotype): Carapace (Figs. 7a–b): flattened, wider than long; prominent eyes, median
eye tubercle dark-brown, without apical setae, lateral eye tubercle smaller, with around half its size. The entire
margin of carapace with sinuous lines and coarse granules, anterior margin with higher sides and center, having
between these shallow pits forming an ill-defined “W”. Overall great amount of small granules mostly anteriorly,
with bundles of larger and sparser granules arranged from the fovea towards the border. Frontal process well-
developed, with greatly widened base and getting abruptly thin in distal half, ending in a very acute hook, not
exceeding frontal line of carapace, this way not visible in dorsal view.
FIGURE 6. Heterophrynus boterorum sp. nov., male (MNRJ 09124) paratype: a. Right chelicera, inner lateral view; b. Same,
outer lateral view; c. Leg IV, basitibia and distitibia showing trichobothria, dorsal view; d. Female gonopod, general dorsal
view. Scale bars = 1 mm (Figs. a–b); 5 mm (Fig. c); 0.5 mm (Fig. d).
Sternum (Fig. 7b): Tri-segmented, typical of Phrynidae. Tritosternum extremely long projected between
pedipalp coxae, reaching almost half their length. With bristles on its entire length, from apex to base, with an
increase in the density at base. Mesosternum convex, reduced to a small tubercle, with six to eight bristles.
Metasternum practically nonexistent, reduced to a flat spot, smaller than mesosternum, with two bristles.
Abdomen (Figs.7a–b): oblong, with almost indistinguishable punctuations, thinner than in the carapace.
Chelicera: Cheliceral furrow (Figs. 9a–b) with 3 internal teeth, the distal one bifid, the first cusp bigger than
the second one II<I=IIIa>IIIb. The third tooth (middle) much smaller than the others. With 3 external teeth, the
distal one bifid, the first cusp bigger than the second one II>I=IIIa>IIIb. Claw with five denticles, with the two
distalmost much reduced, the second smaller than first and third.
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FIGURE 7. Heterophrynus silviae sp. nov., male (MNRJ 09115) paratype: a. Habitus, dorsal view; b. Same, ventral view; c.
Right pedipalp, dorsal view; d. Right pedipalp, ventral view. Scale bars = 5 mm.
338 · Zootaxa 3647 (2) © 2013 Magnolia Press GIUPPONI & KURY