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Systematics and biogeography of Spinoctenus, a new genus of wandering

spider from Colombia (Ctenidae)
Article in Invertebrate Systematics · February 2018

DOI: 10.1071/IS17022
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Invertebrate Systematics, 2018, 32, 111–158
https://doi.org/10.1071/IS17022
Systematics and biogeography of Spinoctenus, a new genus

of wandering spider from Colombia (Ctenidae)
Nicolas A. Hazzi A,B,C,H, Daniele Polotow D,E, Antonio D. BrescovitF,

Ranulfo González-Obando A and Miguel SimóG
A
Departamento de Biología, Facultad de Ciencias Naturales y Exactas, Universidad del Valle, Santiago de Cali,
Colombia.
B
Department of Biological Sciences, The George Washington University, Washington, D.C. 20052, United States.
C
Fundacion Ecotonos, Cra 72 No. 13A-56, Cali, Colombia.
D
Arachnology Laboratory, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118,
USA.
E
Museu de Zoologia “Adão José Cardoso”, Universidade Estadual de Campinas, Rua Charles Darwin s/n,
Bloco N - Cx Postal 6109, 13.083-863, Campinas, SP, Brazil.
F
Laboratório Especial de Coleções Zoológicas, Instituto Butantan, Av. Vital Brazil, 1500, 05503-900,
São Paulo, SP, Brazil.
G
Sección Entomología, Facultad de Ciencias, Universidad de la República, Iguá 4225, CP11400, Montevideo,
Uruguay.
H
Corresponding author. Email: nicolashazzi@hotmail.com
Abstract. Among ctenid spiders, ctenines comprise the most diverse subfamily. In this study, a new genus of Cteninae,
Spinoctenus, is proposed to include the type species S. yotoco, sp. nov. Ten new species are also described: S. escalerete,
S. pericos, S. eberhardi, S. spinosus, S. stephaniae, S. nambi, S. florezi, S. tequendama, S. chocoensis and S. flammigerus.
Results of the parsimony and Bayesian phylogenetic analyses using morphological and behavioural characters indicate the
monophyly of this genus, closely related to Phoneutria Perty, 1883 and Ctenus Walckenaer, 1805. This genus can be
distinguished from the remaining Ctenidae by three unambiguous synapomorphies: embolus with folded process, tegulum
with median process, and RTA curved internally close to the cymbium. A dispersal-vicariance biogeographical analysis of
the genus in the Andean and Chocó regions indicates the origin of Spinoctenus in the Western and Central Andean
Cordilleras. From this region, three events of dispersal occurred to the other regions (one to the Chocó and two to the Eastern
Cordillera), which were subsequently followed by three events of vicariance, suggesting that dispersal and vicariance were
equally important in shaping the current distribution patterns of Spinoctenus species. The discovery of this new genus
containing a large number of new species in the Andean and Chocó regions highlights the current poor knowledge of the
Colombian biodiversity.
http://zoobank.org/urn:lsid:zoobank.org:pub:A7DA044C-8A59-4FAE-8F3B-00D3D2498820
Additional keywords: Andes, Araneae, cloud forest, Ctenus, Phoneutria, tropical rainforest, vicariance and dispersion.
Received 24 July 2016, accepted 25 June 2017, published online 1 February 2018
Introduction tropical forests all over the world (Silva 2003). Some of these
Ctenidae is well represented in the Neotropics by small to large species are unique to forest and their densities decrease
wandering spiders that usually inhabit the ground of forests and significantly when forested habitats suffer fragmentation and
understory vegetation; only a few species are known to be disturbance (Rego et al. 2005, 2007). Species of the genus
arboreal (Jocqué and Dippenar-Schoeman 2006). This family Phoneutria are some of the best-known representatives of this
is one of the most diverse in Araneae, comprising 514 species in family, commonly referred to as banana spiders, because they
47 genera (World Spider Catalog 2017), and it is abundant in often inhabit this crop. They are considered aggressive, and
Journal compilation CSIRO 2018 www.publish.csiro.au/journals/is

112 Invertebrate Systematics N. A. Hazzi et al.
among the most medically important spiders in the world Ciencias Naturales-Universidad Nacional de Colombia, Bogota
(Foelix 2010). Their venom has a neurotoxic action (Foelix (E. Flórez); MPUJ, Museo Pontificia Universidad Javeriana,
2010) and many researchers have analysed its components Bogota (D. Forero).
and the epidemiology of bites (Bücherl 1953a, 1953b, 1956;
Simó 1983; Cruz-Höfling et al. 1985; Marangoni et al. 1993; Morphological examination and description of species
Pineda and Florez 2002; Flórez et al. 2003; García et al. 2008;
The following abbreviations are used: AC, aciniform gland
Maguiña et al. 2008).
spigot; ALS, anterior lateral spinneret; C, conductor; CY,
For many years, the study of Ctenidae was hindered by
cylindrical gland spigot; E, embolus; FP, folded process of the
poor original descriptions and a lack of knowledge of the
embolus; IB, internal bulge of the embolus; LF, lateral field; LP,
evolutionary relationships, which prevent the clarification of
lateral process; LL, locking lobe; MA, median apophysis; MAP,
the taxonomic status of most genera. Nevertheless, during the
major ampullate gland spigot; mAP, minor ampullate gland
past three decades significant advances were published in
spigot; MS, median sector; MTP, median tegular process; PI,
multiple generic taxonomic revisions and systematic works,
piriform gland spigot; PLS, posterior lateral spinneret; PMS,
especially in the Neotropics, which have helped to clarify
posterior median spinneret; RSB, retrolateral sclerotised bulge
the phylogenetic relationships and the diagnosis of most of
of male tibia palp; ta, tartipore; t, tegulum.
the neotropical genera (Lachmuth et al. 1985; Simó and von
Specimens were preserved in 75% ethanol. Descriptions and
Eickstedt 1994; Brescovit 1996; Brescovit and Simó 1998, 2007;
terminology follow Höfer et al. (1994) and Simó and Brescovit
Höfer and Brescovit 2000; Simó and Brescovit 2001; Silva
(2001). All measurements were taken with a micrometric
2003, 2004; Martins and Bertani 2007; Polotow and Brescovit
ocular and are in millimetres. Epigyna were digested with
2008, 2009a, 2009b, 2009c, 2013, 2014; Hazzi and Silva
trypsin or pancreatin solution (Álvarez-Padilla and Hormiga
2012; Polotow et al. 2015). Taxonomic development have
2007) to enable study of internal structures. Digital images
also facilitated advances in natural history and ecological
were made with a Nikon MSZ1500 stereomicroscope adapted
studies (Merrett 1988; Simó and Bardier 1989; Höfer et al.
to a Nikon DS-Fi1 digital camera. Extended focal range
1994; Schuster et al. 1994; Almeida et al. 2000; Simó et al.
images were composed with NIS-Elements BR software,
2000; Gasnier and Höfer 2001; Folly-Ramos et al. 2002; Gasnier
ver. 3.1. Digital SEM photographs were taken on a scanning
et al. 2002; Dias and Brescovit 2004; Polotow et al. 2007;
electron microscope from the scanning laboratory, Ingenieria de
Torres-Sánchez and Gasnier 2010; Dias et al. 2011; Hazzi 2014).
Materiales, Universidad del Valle, and with a JEOL JSM 5900
The literature mentioned above mostly focusses on
from Facultad de Ciencias, Universidad de la República,
Brazilian ctenids and those spiders of neighbouring countries,
Montevideo, Uruguay. For scanning electron microscope
but the general knowledge of ctenids in Colombia is still
preparation, structures were cleaned ultrasonically, transferred
incipient. Recently, Hazzi et al. (2013) presented new records,
to 95% and then to 100% ethanol for 10 min in each immersion
geographical distribution extensions and a checklist of
before being critical-point-dried or air-dried. Finally, structures
the current ctenid species in Colombia based on the review of
were mounted on rivets using copper wire then sputter-coated
four arachnological collections and published literature; the
with gold.
species diversity of ctenids in Colombia was increased from
Species descriptions are in alphabetical order in the text,
16 to 25. However, several specimens of this family could not
except for Spinoctenus yotoco, which appears first because is the
be assigned to any of the currently described Ctenidae genera.
type species of the genus.
In the present study, Spinoctenus is newly described to
accommodate some of those species and they are included in
the most recent phylogenetic analysis based on morphological Phylogenetic analysis
characters of Ctenidae (Polotow and Brescovit 2014) to A morphological matrix based on the most recent phylogenetic
test the monophyly of the genus and relationships with the analysis of the family Ctenidae (Polotow and Brescovit
remaining genera of the family. In addition, we present a 2014) was used, which comprises 72 terminal taxa scored for
historical biogeographical analysis of this genus based on the 89 characters. Recently, Polotow et al. (2015), in a phylogenetic
phylogenetic relationships of the species obtained in this study. total evidence analysis, demonstrated that the subfamily
These species are found in forests of the Andean and Chocó Viridasiinae is a family apart from Ctenidae. Thus, we
biogeographical regions in Colombia, which have undergone deactivated in TNT 1.1 (Goloboff et al. 2008) the five species
recent dynamic geological processes (Hoorn et al. 1995, 2010; of viridasiids (Viridasius fasciatus (Lenz, 1886), Vulsor
Kattan et al. 2004; Sedano and Burns, 2010) that may have isoalensis (Ono, 1993), Asthenoctenus borellii Simon, 1897,
influenced modes of speciation and current distributions. Ctenus tarsalis F.O. Pickard-Cambridge, 1902 and C. bulimus
Strand, 1909) and four related characters (char: 28, 38, 54
Methods and 80). Then, 23 characters (Table 1) and 12 species were
added to the dataset, to a total of 108 characters and 79
Museum abbreviations terminals. The additional species were: Phoneutria boliviensis
The material examined belongs to the following Colombian (F.O. P.-Cambridge, 1897), Spinoctenus yotoco, sp. nov.,
museums (curators in parentheses): IMCN, Museo de S. spinosus, sp. nov., S. nambi, sp. nov., S. flammigerus, sp.
Ciencias Naturales Federico Carlos Lehmann-INCIVA, nov., S. stephaniae, sp. nov., S. eberhardi, sp. nov., S. pericos, sp.
Cali (R. Rios); MUSENUV, Museo Entomológico de la nov., S. escalerete, sp. nov., S. florezi, sp. nov., S. chocoensis,
Universidad del Valle, Cali (C. E. Posso); ICN, Instituto de sp. nov. and S. tequendama, sp. nov.
Taxonomy and systematics of Spinoctenus Invertebrate Systematics 113
Table 1. Character descriptions of Spinoctenus species
(90) Epigynum – copulatory ducts orientation: (0) parallel (convergent); (1) divergent (Figs 20D, 23E). Non-parallel copulatory ducts occur in a clade formed
by S. spinosus and S. florezi.
(91) Epigynum – shape of the spermathecae: (0) not comma-shaped; (1) comma-shaped (Fig. 14D, E). The presence of this feature occurs in a clade formed
by S. escalerete and S. chocoensis.
(92) Epigynum – spermathecae with one or two small anterior granules: (0) absent; (1) present (Fig. 14D, E). The presence of this feature occurs in a clade
formed by S. escalerete and S. chocoensis.
(93) Male palp – folded process of the embolus: (0) absent; (1) present (Figs 9B, 26A–K). The presence of this feature is a synapomorphy of Spinoctenus.
(94) Male palp – shape of the folded process of the embolus: (0) large and non-uniform (Fig. 26A–C); (1) thin and uniformly curved (Fig. 26E, G). The thin
and uniformly curved folded process of the embolus occurs in a clade formed by S. nambi and S. flammigerus.
(95) Male palp – external margin of the embolus: (0) small (Fig. 26C); (1) large (Fig. 26A, D). The large external margin of the embolus occurs in a clade
formed by S. yotoco, S. tequendama, S. spinosus, S. florezi, S. escalerete, S. chocoensis and S. pericos.
(96) Male palp – embolus form: (0) curved (Figs 9A, 27A); (1) straight (Figs 11A, 12E, 27H, C). The presence of a straight embolus derived feature occurs
in S. escalerete, S. chocoensis and S. pericos.
(97) Male palp – median tegular process: (0) absent; (1) present (Fig. 9C). This feature is a synapomorphy of Spinoctenus, gen. nov.
(98) Male palp – probasal rounded tegulum: (0) absent; (1) present (Fig. 10A, D). The presence of this feature arises independently in a clade formed by
Phoneutria and Spinoctenus, gen. nov. and in another clade formed by Ctenus dubius and C. nigritus.
(99) Male palp – retrolateral tibial apophysis (RTA) orientation: (0) far from the cymbium; (1) curved internally close to the cymbium (Figs 17B, 27A–K).
The presence of an RTA curved internally close to the cymbium is a synapomorphy of Spinoctenus, gen. nov.
(100) Male palp – triangular RTA: (0) absent; (1) present (Figs 12D, 13G). The presence of this feature occurs in a clade formed by S. stephaniae,
S. flammigerus and S. nambi.
(101) Male palp – small retrolateral sclerotised bulge (RSB) on male tibia: (0) absent; (1) present (Figs 11F, 13G, 28F, J, K). This feature is a synapomorphy of
the clade formed by S. bogotensis, S. nambi, S. flammigerus, S. stephaniae, S. yotoco, S. spinosus, S. florezi, S. escalerete and S. pericos, secondarily lost
in S. pericos.
(102) Male palp – small rounded lobe on male tibia: (0) absent; (1) present (Fig. 11D, F). The presence of this feature occurs in a clade formed by S. tequendama,
S. yotoco, S. spinosus, S. florezi, S. escalerete, S. chocoensis and S. pericos.
(103) Legs – females with scopula on metatarsi I and II: (0) absent; (1) present. The presence of this feature occurs in a clade formed by S. spinosus,
S. florezi, S. escalerete, S. chocoensis and S. pericos, but this character presents missing entries for several terminals.
(104) Legs – males with partial scopulae on metatarsus III: (0) absent; (1) present. The presence of this feature occurs in a clade formed by S. yotoco, S. spinosus,
S. florezi, S. escalerete and S. pericos, and is lost in S. chocoensis. This character presents missing entries for several terminals.
(105) Legs – males with long ventral setae in tibiae I and II: (0) absent; (1) present. The absence of this feature occurs independently in S. eberhardi and
S. tequendama, but this character presents missing entries for several terminals.
(106) Legs – males with long ventral setae on metatarsus III and tibia III: (0) absent; (1) present. The presence of this feature occurs in a clade formed by S. yotoco,
S. spinosus, S. florezi, S. escalerete and S. pericos, and is lost in S. chocoensis, but this character presents missing entries for several terminals.
(107) Legs – males and females with scopulae on tarsi I and II: (0) abundant and thin (Fig. 30B); (1) scarce and thick (Fig. 30C). The presence of this feature
occurs in a clade formed by S. stephaniae, S.nambi and S. flammigerus, but this character presents missing entries for several terminals.
(108) Legs – spination pattern in metatarsus IV of males r1–0-1–1-3–2-1: (0) absent; (1) present (Fig. 29F, I). The presence of this spination pattern occurs in a
clade formed by S. stephaniae, S. nambi and S. flammigerus.
(109) Coloration – males with distal tarsus IV yellowish at the apex: (0) absent; (1) present (Fig. 30A). The presence of this feature occurs in a clade formed
by S. escalerete and S. chocoensis.
(110) Coloration – live specimens with a longitudinal black line on the carapace: (0) absent; (1) present. The presence of this feature is a synapomorphy of
the Phoneutria clade.
(111) Coloration – white setae on the sternum: (0) absent; (1) present. The absence of this feature occurs independently in S. stephaniae and a clade formed
by S. spinosus, S. florezi, S. escalerete and S. pericos, but this character presents missing entries for several terminals.
(112) Coloration – dorsal coloration in live specimens: (0) brown and black designs (Figs 31A–F, 32A, B); (1) orange and reddish designs (Fig. 33C–E).
The presence of orange and reddish coloration occurs in a clade formed by S. nambi and S. flammigerus.
Most of the new characters were easy to score based characters were treated as non-additive (Fitch 1971) to avoid
on species descriptions from previous analyses of the group. previous assumptions on the evolution of the character states.
A combination of binary coding with inapplicable entries and Cladistic analyses were conducted using TNT 1.1. Heuristic
unordered multistate characters was used. In this technique, the search strategies were performed using the new technology
presence or absence of a feature is scored in a first character and options starting from 30 RAS plus TBR followed by tree
a second character that describes the condition of that particular drifting (command: hold 1000; mult 30 = tbr drift). All trees
feature. Thus, taxa that did not present the feature were scored found during searches were collapsed under ‘rule 1’ (minimum
as ‘inapplicable’ for the second character. This method of possible length is zero; Coddington and Scharff 1994).
scoring is preferred over others as it captures (1) the presence The data matrix was analysed under equal weights (EW) and
or absence of the feature and (2) values (i.e. states) of the feature implied weights (IW) analysis (Goloboff 1993). The sensitivity
(Brazeau 2011). of results (sensu Wheeler 1995) was assessed by varying values
The character matrix was edited in the software Mesquite 2.74 of the concavity constant in the IW analyses (k = 1–11 command:
(Maddison and Maddison 2010). Inapplicable and unknown Piwe = N’, where ‘N’ is the k value). In total, 12 analyses were
states are presented as ‘–’ and ‘?’, respectively. Multistate conducted (one EW and 11 IW analyses). As support measures,
we used symmetric resampling frequencies expressed as (k = 1–5). We chose this strict consensus tree (six nodes
frequency differences (Goloboff et al. 2003). WinClada collapsed; L = 249; CI = 0.53; RI = 0.85; Figs 1, 2) as the
10.00.08 (Nixon 2002) was used to study the character preferred hypothesis because it was more stable (less parameter-
optimisations on the preferred cladogram using accelerated dependent) and presented higher average support values of
transformation (ACCTRAN) (Farris 1970; Swofford and symmetric resampling. The sensitivity of the results of IW to
Maddison 1987). Although Agnarsson and Miller (2008) different values of the concavity constant is presented in
demonstrated that there are no theoretical grounds for Table 2. The Bayesian inference topology (Fig. 3) obtained
favouring ACCTRAN or DELTRAN in ambiguous character was congruent with the maximum parsimony analysis. These
optimisations, we used ACCTRAN to try to keep the results are also congruent with the phylogeny of Polotow and
hypothesis of character transformations of Polotow and Brescovit (2014).
Brescovit (2014). Nevertheless, we differentiated ambiguous In all analyses, using different weighting regimes (EW and
character optimisations by ACCTRAN from unambiguous IW) and different optimality criteria (parsimony and Bayesian),
optimisations in the phylogenetic tree using different colours. Spinoctenus appears as a monophyletic group supported by
In addition to the cladistics analysis, we analysed the matrix high symmetric resampling and posterior probability values.
using Bayesian inference with a likelihood model appropriate Spinoctenus is placed within the subfamily Cteninae, closely
for morphological data (Lewis 2001). The analysis was related to Phoneutria with low support, although their grouping
performed in MrBayes (Ronquist and Huelsenbeck 2003) is obtained in all analyses. This clade is morphologically
under the MkvG model (Lewis 2001; Nylander et al. 2004). supported by two homoplastic conditions: embolus with
The G parameter allows rate variation between characters to internal bulge (char. 26) and probasal rounded tegulum (char.
follow a gamma distribution, which may be analogous to those of 98). These two conditions are independently derived in some
the IW parsimony (Nylander et al. 2004). We sampled trees and species of Ctenus, which appears as the sister clade of these
parameters every 1000 generations from four chains (one cold two genera.
and three heated), 25% of the trees were discarded as burn-in The monophyly of Spinoctenus is supported by three non-
and the remainder were used to calculate posterior probabilities. ambiguous synapomorphies: the presence of a tegular median
To check that the run was long enough for the chains to converge, process (char. 97), an RTA curved internally close to the
the probabilities of the marginal parameters were observed in cymbium (char. 99) and embolus with a folded process (char.
Tracer v. 1.5 (Rambaut et al. 2014b). The consensus trees with 93). The genus is also supported by the following homoplastic
branch lengths and node posterior probabilities were edited in condition: male metatarsus IV curved and with several small
FigTree 1.3.1 (Rambaut 2014a). spines (char. 76). In addition, the cladistic and Bayesian analyses
show several well-supported groups among Spinoctenus species.
Biogeographical analysis Spinoctenus eberhardi is sister to the remaining Spinoctenus
Based on the geographical distribution patterns of the species (although with low support) and two well-supported
Spinoctenus species and recent studies on biogeographical clades are identified. The first clade includes the species
regionalisation (Hernández-Camacho et al. 1992; Morrone Spinoctenus stephaniae, S. nambi and S. flammigerus and is
2014), we conducted the analysis using the following supported by the following synapomorphies: RTA triangular
biogeographical units (Fig. 4): Chocó, Central and Western (char. 100); few and thick scopulae on tarsi I and II of both
Cordilleras (Cauca province), and Eastern Cordillera. These males and females (char. 107), and the spination pattern
regions are divided by topographical barriers and endemism of r1–0-1–1-3–2-1 on metatarsus IV of males (char. 108). The
Spinoctenus species. A dispersal-vicariance analysis (DIVA; loss of the internal bulge of the embolus (char. 26) is a
Ronquist 1997) implemented in program RASP 2.0 Beta homoplastic condition also supporting this group. The second
(Yu et al. 2010, 2015) was performed. This program clade includes the remaining species (Spinoctenus tequendama,
complements DIVA with the methods of Nylander et al. (2008) S. yotoco, S. spinosus, S. florezi, S. pericos, S. escalerete and
and Harris and Xiang (2009), which can determine statistical S. chocoensis) and is supported by two non-ambiguous
support for the reconstruction of the ancestral area of each clade synapomorphies: large external margin of the embolus (char.
(‘S-DIVA’ value). As outgroups for the analysis, we used the 95) and small rounded lobe on the RTA (char. 102). This clade
genera Phoneutria and Ctenus (see ‘Results), whose species are is also supported by an ambiguous synapomorphy: males with
mostly distributed on the eastern side of the Andean Cordilleras long ventral setae on metatarsus and tibia III (char. 106).
(cis-Andean region).
Biogeography
Results
The DIVA analysis (Fig. 5) indicated three events of dispersal,
Phylogeny which were subsequently followed by three events of vicariance,
Searches under EW in TNT found 53 most parsimonious trees suggesting that both types of events were equally important in
(L = 256; CI = 0.55; RI = 0.85). In the strict consensus, 13 nodes shaping the current distribution patterns of Spinoctenus species.
collapsed and resulted in a tree with 260 steps (CI = 0.50; This analysis supports an origin of Spinoctenus in the Western
RI = 0.80). In the IW analysis with concavity values and Central Cordilleras of the Andes. One event of dispersal from
k = 1–11, fewer trees of the same length (3) were obtained this area occurred to the Chocó region followed by vicariance,
than with the EW analysis. The topology of the strict consensus originating the three species of Spinoctenus that are found in this
trees was stable in five of the 11 weighting regimes explored region (S. chocoensis, S. escalerete and S. pericos). In addition,
Tengella radiata
Zoropsis spinimana
Acanthocteninae
99 Calocteninae
0.90
Acantheinae
79
0.30
25
0.04
Cteninae
Fig. 1. Strict consensus of the three most parsimonious trees obtained by cladistic analysis under implied weights (k = 4).
Support values for groups are symmetric resampling expressed as GC ‘‘Group present/Contradicted’’ frequency differences
(top) and Bremer support in units of fit 100 (bottom).
Fig. 2. Consensus tree under the implied weights analysis with k = 4. Non-homoplasious, homoplasious and ambiguous transformations (recovered by
ACCTRAN) indicated by black, white and grey circles, respectively along branches with character numbers above and character states below each.
Table 2. Summary of the phylogenetic hypotheses among the most Diagnosis

parsimonious trees (MPTs) found with equal weights (EW) and implied
Males of Spinoctenus can be distinguished from the remaining
weights (IW) with 11 values for the concavity constant (k)
CI, consistency index; RI, retention index; Av. Sym, average support Ctenidae by three unambiguous synapomorphies: embolus with
symmetric resampling folded process (Figs 9B, 26A–K), tegulum with median
sclerotised process (Fig. 9C) and RTA curved internally close
Length MPTs CI RI Av. Sym to cymbium (Figs 17B, 27A–K). Females can be distinguished
from Ctenus and Phoneutria by the combination of the following
EW 256 59 0.55 0.85 49.7 characters: epigynum with reduced posterior area of the median
1 257 3 0.55 0.85 53.2
sector and anterior area less sclerotised and slightly excavated
2 257 3 0.55 0.85 53.9
3 257 3 0.55 0.85 54.0
(Fig. 15D); vulva with copulatory ducts large and strongly
4 257 3 0.55 0.85 54.4 sclerotised (Fig. 15E).
5 257 3 0.55 0.85 54.2
6 256 3 0.55 0.85 54.0 Description
7 256 3 0.55 0.85 53.8
Medium- to large-sized ecribellate spiders, total length
8 256 3 0.55 0.85 53.8
10.00–26.60. Carapace piriform, brown with a pale yellow
9 256 3 0.55 0.85 53.7
10 256 3 0.55 0.85 53.7 longitudinal median line, black pigment around eyes; thoracic
11 256 3 0.55 0.85 53.8 groove longitudinal, in the posterior third. Ctenid eye pattern
2–4-2, with the anterior and posterior row recurved in dorsal view
(Fig. 6A, B). Eyes round, except oval anterior lateral eyes, with
two other events of dispersal took place from the ancestral area to white setae around PLE and PME and lateral of AME. Chilum
the Eastern Cordillera, followed by vicariance, originating divided. Clypeus with long erect black bristles. Chelicerae
Spinoctenus tequendama and S. florezi independently. brown, promargin with three teeth; retromargin with four
similar-sized teeth and three small teeth (Fig. 7A), prominent
Taxonomy basal condyle, intermarginal denticles present. Endites brown
with anterolateral serrula and anteromedian scopulae. Labium
Family CTENIDAE Keyserling dark brown, slightly longer than wide (Fig. 6C, D). Sternum
truncated anteriorly and posteriorly pointing, not extending
between coxae IV (Fig. 6C). Ventral faces of coxae light
Subfamily CTENINAE Keyserling brown with dark brown spots. Male legs more elongated and
slender than female legs. Tarsal organ with central narrow
Spinoctenus, gen. nov. opening (Fig. 7B). Trochanter notched. Tarsus with claw tufts
http://zoobank.org/urn:lsid:zoobank.org:act:19641BD4-4C5B-4A50- (Fig. 7D) composed of tenent setae, specialised in adhesion to
A430-5729ECD0CB31 smooth surfaces (Ramírez 2014). Trichobothria bases with
Type species: Spinoctenus yotoco, sp. nov. many transverse grooves (Fig. 7C). Abdomen oval, brown,
Tengella_radiata
Zoropsis_spinimana
Nothroctenus_marshii
Acanthoctenus_spiniger
Viracucha_andicola
Celaetycheus_flavostriatus
Caloctenus_gracilitarsis
Toca_bossanova
Gephyroctenus_philodromoides
Acantheis_laetus
Chococtenus_miserabilis
Ctenus_inazensis
Enoploctenus_cyclothorax
Ctenus_curvipes
Ctenus_sinuatipes
Afroneutria_velox
Afroneutria_immortalis
Afroneutria_erythrochelis
Leptoctenus_paradoxus
Leptoctenus_byrrhus
Ctenus_inaja
Thoriosa_sp_
Thoriosa_taurina
Trogloctenus_fagei
Amauropelma_trueloves
Amauropelma_torbjorni
Celaetycheus_modestus
Ohvida_vernalis
Ohvida_fulvorufa
Centroctenus_ocelliventer
Centroctenus_acara
Ctenus_lejeunei
Ctenus_amanensis
Africactenus_decorosus
Africactenus_monitor
Petaloctenus_bossema
Petaloctenus_songan
Ctenus_similis
Isoctenus_coxalis
Isoctenus_folliifer
Parabatinga_brevipes
Parabatinga_sp_nov_
Ctenus_bngipes
Ctenus_villasboasi
Ctenus_satanas
Ctenus_fallax
Ctenus_eminens
Ctenus_pergulanus
Ctenus_rectipes
Ctenus_pauloterrai
Ctenus_manauara
Ctenus_nigritus
Ctenus_dubius
Ctenus_crulsi
Ctenus_amphora
Ctenus_minor
Ctenus_paubrasil
Ctenus_medius
Ctenus_ornatus
Ctenus_vehemens
Ctenus_fernandae
Phoneutria_boliviensis
Phoneutria_fera
Phoneutria_nigriventer
Spinoctenus_stephaniae
Spinoctenus_nambi
Spinoctenus_flammigerus
Spinoctenus_eberhardi
Spinoctenus_tequendama
Spinoctenus_yotoco
Spinoctenus_spinosus
Spinoctenus_florezi
Spinoctenus_pericos
Spinoctenus_escalerete
Spinoctenus_chocoensis
Leptoctenus_aff_agalenoides
Anahita_centralis
Anahita_mamma
Anahita_blandini
Fig. 3. Bayesian optimal tree obtained under Mkv+ model. Nodes with posterior probability values.
dorsum with a black anterior border, lighter centrally and with a S. nambi, S. flammigerus and S. stephaniae. Locking lobes
folium-like pale brown longitudinal band, venter brown with present (Figs 9B, 11B); conductor membranous, hyaline and
four divergent series of white dots. Spinnerets (Fig. 8A–D): C-shaped (Fig. 9A, C). Epigynal plate (Figs 9D, 14A, C,
anterior laterals (ALS) long and wide in the apex; posterior 15D–25D) consisted in a median sector and two lateral lobes;
medians (PMS) roughly conical, short and narrow in the apex; median sector with anterior area less sclerotised and slightly
posterior laterals (PLS) long and conical. Female ALS (Fig. 8B) excavated with heavily sclerotised margins; lateral field
with two major ampullate gland spigots and more than 20 with small lateral spurs that originate medially and are
aciniform gland spigots. PMS (Fig. 8C) with two minor directed medially; copulatory opening pocket-like. Internally
ampullate gland spigots plus a tartipore and ~15 cylindrical (Figs 14B, D, 15E–25E), copulatory ducts large and strongly
gland spigots. PLS (Fig. 8D) with about 10 cylindrical and sclerotised; spermathecae large with undifferentiated head
more than 20 aciniform gland spigots. Male palp: tibia and base, with or without anterior granules; fertilisation ducts
without suture line, as long as cymbium, straight with an originate basally.
apical RTA curved internally close to the cymbium (Figs 15B,
16B, 17B, 18B, 19B, 20B, 21B, 22B, 23B, 24B, 25B, 27A–K) Composition
with a small sclerotised rounded lobe (Figs 11D, F, 12H, 13D)
Eleven species: S. yotoco, sp. nov., S. escalerete, sp. nov.,
except in S. stephaniae, S. eberhardi, S. flammigerus and
S. pericos, sp. nov., S. stephaniae, sp. nov., S. eberhardi, sp.
S. nambi (Figs 12D, 13G). Cymbium with elongated tip,
nov., S. florezi, sp. nov., S. tequendama, sp. nov., S. spinosus,
without lateral or dorsal projections nor scopulae path on
sp. nov., S. nambi sp. nov., S. flammigerus and S. chocoensis,
dorsum; subtegulum prolateral, with a basal excavation to fit
sp. nov.
the locking lobe (Fig. 9B), tegulum large and rounded pro-basally
with a median sclerotised process (MTP; Fig. 9A, C); median
apophysis cup-shaped (Fig. 9C), embolus short (Figs 9B, 26A–K) Distribution
with internal bulge (Figs 9B, 10B, F, 11A, 13B) except in Andean and Pacific areas of Colombia (Fig. 36A, B).
78°0′0″W 76°0′0″W 74°0′0″W 72°0′0″W
10°0′0″N
8°0′0″N
6°0′0″N
4°0′0″N
2°0′0″N
0°0′0″
0 37.5 75 150 225 300
Kilometers
Fig. 4. Biogeographical regions of Colombia selected for the DIVA analysis. Blue, Chocó (C); red, Western and Central Cordilleras
(WCC); yellow, Eastern Cordillera (EC).
(CI) Ctenus
78°0′0″W 77°0′0″W 76°0′0″W 75°0′0″W 74°0′0″W 73°0′0″W
(CI) Phoneutria
9°0′0″N (WCC) Spinoctenus eberhardi
(WCC) Spinoctenus stephaniae
8°0′0″N (WCC) Spinoctenus nambi
(WCC) Spinoctenus flammigerus
7°0′0″N (EC) Spinoctenus tequendama

N 1
W E
(WCC) Spinoctenus yotoco
6°0′0″N (EC) Spinoctenus florezi

S
2 and 3 (WCC) Spinoctenus spinosus
5°0′0″N (C) Spinoctenus pericos
(C) Spinoctenus chocoensis
4°0′0″N (C) Spinoctenus escalerete
Cis Andean (CI)

3°0′0″N
Western and Central Cordillera (WCC)
Eastern Cordillera (EC)

2°0′0″N
Chocó (C)
CI + WCC
1°0′0″N
0 25 50 100 150 200
WCC + EC
Kilometers
WCC + EC + C
WCC + C
Fig. 5. Ancestral distributions reconstructed by RASP. Pie charts at the nodes give the relative frequencies of the ancestral-area reconstruction. The numbers
in the map correspond with the numbers at the nodes of the phylogenetic tree and indicate interpretations of three major historical events of dispersal followed
by vicariance.
Etymology as Cupiennius granadensis Keyserling, 1877, Ancylometes

The generic name is a combination of ‘spino’, referring to the bogotensis Keyserling, 1877, Ctenus sp. (sinuatipes-group)
macrosetae on metatarsus IV of males, and ‘ctenus’ in reference and Enoploctenus sp.
of the genus type of the family Ctenidae. The gender is masculine. Spinoctenus spinosus occurs in the valley of Cauca River,
in remnants of dry forest (bs-T) and subtropical wet forest (bh-
ST), between elevations of 900 and 1200 m, in sympatry with
Natural history Phoneutria boliviensis, Cupiennius bimaculatus (Taczanowski,
Spinoctenus species live in a variety of ecosystems and across 1874) and Ctenus sp. (sinuatipes-group). Spinoctenus yotoco,
a wide altitudinal range, from sea level to elevations of 2500 m S. eberhardi and S. stephaniae occur in cloud forests at elevations
(Fig. 34A–D). All Spinoctenus species are nocturnal forest between 1600 and 2200 m and in subtropical wet forest (bh-ST)
dwellers, living on ground covered with leaf litter and in and lower montane wet forest (bh-MB). In one location, S. yotoco
some occasions on vegetation in the understory. During was found living in sympatry with S. stephaniae in the leaf litter,
the day, these spiders can be found hiding under leaf litter or along with Chococtenus miserabilis (Strand, 1916), a smaller
decaying fallen logs. Spinoctenus pericos, S. escalerete and ctenid spider. In the central mountain range, S. eberhardi or
S. chocoensis are found in very humid tropical forest (bmh-T) S. stephaniae are sympatric with Caloctenus albertoi Hazzi &
and tropical rainforest (bp-t) according to Holdridge’s life Silva, 2012. Spinoctenus florezi and S. bogotensis inhabit cloud
zones, between elevations of 40 and 600 m. In the ‘Reserva forests of the eastern mountain range. Spinoctenus bogotensis is
Pericos’, these three species were collected and/or observed the species of the genus with the highest elevation record, and can
in the same parts of the forest and the same microhabitats be found at elevations up to 2500 m.
(i.e. leaf litter on the ground), which indicates that they are Spinoctenus yotoco has been observed on three occasions
syntopic and may have strongly overlapping niches. The three eating cockroaches and on one occasion eating an annelid
Spinoctenus species were also collected with other ctenids, such worm (Fig. 31A). Over the course of two night observations,
(A) (B)
(C) (D)
Fig. 6. Spinoctenus yotoco. (A, B) Front and dorsal view of the eye pattern; (C) sternum, labium and endites; (D) close-up of labium and endites.
we observed four events of cannibalism in S. stephaniae, in two behavioural units were only observed in the first event,
which adult females were detected eating juveniles (in a locality because the female rejected the male in the second event.
only inhabited by this species). In addition, this species was Male approximation: during this stage, the male begins to
recorded as eating a cricket (Gryllidae) and a harvestmen. approach the female slowly with his forelegs raised. When the
In Reserva Pericos, we observed a female S. escalerete eating male contacts the female with his forelegs, the female responds
a juvenile Spinoctenus; however, it was not possible to establish by vigorously shaking her forelegs to remove the male’s legs
whether it was cannibalism or intra-guild predation, because and stay in the same position. The male then moves away and
this reserve is inhabited by two more species of Spinoctenus circumvents the female to approach again from another direction.
(S. pericos and S. chocoensis). Also, S. escalerete was recorded This behaviour occurred several times (up to seven times) and
as eating cockroaches (Fig. 32A). lasted for up to 12 min.
In Reserva Pericos, we observed and recorded with a video Groping: after several approaches of the male, if the female
camera two instances of mating in Spinoctenus escalerete, which finally accepts him, the male touches her very briefly (less than
occurred between 20:00 and 21:00 on different nights. Both 3 s) on the sides of her carapace with his first legs repeatedly
events occurred on the leaves of low vegetation (30–40 cm above (Fig. 35A) and then joins his second and third legs with the
the ground). We identified three behavioural units of the mating female’s first and second legs, respectively, moving them slightly
process: male approximation, groping and copulation. The last in a simultaneous fashion (5 s; Fig. 35B).
Copulation: the male climbs over the female so they face in pronounced (Figs 15C, 28A), median tegular process short
opposite directions (Fig. 35C). The female draws her legs close (Figs 9A, C, 27A); RTA as in Fig. 15C............... Spinoctenus yotoco
to her body, resulting in a configuration where the patellae of all 9(7). Tarsus IV white at apex (Fig. 30A) and metatarsus IV with curvatures
her legs almost touch each other above her carapace. The male (Fig. 29B, C) ....................................................................................10
Tarsus IV without lighter apex, metatarsus IV straight (Fig. 29G) and
moves laterally to the sides of the female’s body and contacts her
embolus with a small external distal bulge (Figs 12E, 27H) ..............
epigynum with his palps. The mating position was type III,
Spinoctenus pericos
according to Foelix (2010). Copulation lasted less than 5 s. 10(9). Embolus with small and non-spiniform internal bulge (Figs 11A, 27D),
Due to difficulties of the field-based observation (i.e. limited folded process pointed in the middle (Figs 11B, 26D); presence of
light) it was not possible to count the number of palpal insertions. a ventral sclerotised tegular process near the median apophysis
Females of Spinoctenus yotoco and S. pericos were observed (Figs 11A, C, 28D); metatarsus IV with few spines and a large
with egg sacs both in captivity and in the field. The egg sacs were distal hook-shaped spine (Fig. 29C)...............Spinoctenus escalerete
white with a flat face attached to the substrate (i.e. litter leaf or Embolus with large and spiniform internal bulge (Figs 10A, B, 27B), folded
the wall of the terrarium) and a convex face. Females protected process curve (Figs 10A, C, 26B); without tegular process near the
the egg sacs by standing over them (Fig. 32B) and it was not median apophysis; metatarsus IV curved and with many small spines
possible to record the emergence of spiderlings. Two females of (Fig. 29B) ...................................................... Spinoctenus chocoensis
S. eberhardi were observed in the field carrying egg sacs in their
chelicerae (Fig. 31E).
Key to identification of species of Spinoctenus – Females
Key to identification of species of Spinoctenus – Males 1. Dorsum of the body orange and reddish (Fig. 32C–E)...........................2
1. Embolus with folded process reduced (Figs 10G, 17A, 26C) and with Dorsum of the body brown-grey (Figs 31A–F, 32A, B, F, 33A–D) ...........3
internal bulge as in Figs 10E, F, 27C............. Spinoctenus eberhardi 2(1). Middle field of the epigynum quadrangular (Fig. 21D); spermathecae
Embolus with folded process large and conspicuous (Figs 9B, 26A, B), with anterior granules (Fig. 21E) ......................... Spinoctenus nambi
internal bulge otherwise or absent.....................................................2 Middle field of the epigynum triangular (Fig. 19D); spermathecae without
2(1). Embolus without internal bulge (Figs 12A, 13E, H, 27G, J); RTA anterior granules (Fig. 19E)........................ Spinoctenus flammigerus
triangular (Figs 12D, 13G, 19C, 21C, 24C) without a small rounded 3(1). Copulatory ducts with non-parallel orientation (Figs 15E, 16E, 17E,
lobe; spination pattern r1–0-1–1-3–2-1 on metatarsus IV (Fig. 29F, I) 18E, 19E, 21E, 22E, 24E, 25E) .........................................................4
...........................................................................................................3 Copulatory ducts with parallel orientation (Figs 20E, 23E) .......................5
Embolus with internal bulge (Figs 9B, 10A, B, 11A, 13A, B); RTA otherwise 4(3). Spermathecae with anterior granules (Fig. 23E), middle field of the
and with a small rounded lobe (Figs 11D, F, 12H, 13D); spination epigynum wide towards the base (Fig. 23D) ... Spinoctenus spinosus
pattern of metatarsus IV otherwise (Fig. 29A–E, G, H, J)................5 Spermathecae without anterior granules (Fig. 20E), middle field of the
3(2). Median apophysis not rounded (Figs 13E, H, 24B, 27J); folded process epigynum narrow towards the base and wider anteriorly (Fig. 20D)
of the embolus large, occupying more than half the area of the embolus .............................................................................. Spinoctenus florezi
(Figs 13E, F, 24A, B); coloration of the dorsum of the body typical of 5(3). Comma-shaped spermathecae present, with one or two small anterior
the genus (Fig. 33C, D) .................................Spinoctenus stephaniae granules (Figs 14D, E, 16E, 18E)......................................................6
Median apophysis rounded (Figs 12A, C, 19B, 21B, 27E, G); folded process Spermathecae otherwise ..............................................................................7
of the embolus thin and curved (Figs 12B, 26E, G); coloration of the 6(5). Middle region of the vulva between the two copulatory ducts narrow
body orange and reddish (Fig. 32C–E) .............................................4 (Fig. 18E); large-sized spiders (carapace length 10.10–10.62)...........
4(3). Embolus with whitish membranous internal margin towards the tip ........................................................................Spinoctenus escalerete
(Figs 21B, 27G) and prolateral margin wide (Figs 21A, 26G) ........... Middle region of the vulva between the two copulatory ducts wide (Fig. 16E);
.............................................................................. Spinoctenus nambi medium-sized spiders (carapace length 6.75–7.93) ............................
Embolus without small membranous whitish internal margin towards the tip ...................................................................... Spinoctenus chocoensis
(Figs 19B, 27E) and thin external lateral margin (Figs 19A, 26E) ..... 7(5). Large-sized spiders (carapace length 12.25–12.62), spermathecae
.................................................................... Spinoctenus flammigerus rounded (Figs 14F, 22E) ................................... Spinoctenus pericos
5(2). Tibiae I, II and III and metatarsus III’without long ventral setae, Medium-sized spiders (carapace length <10.50), spermathecae non-rounded
metatarsus IV unmodified (Fig. 29J).......... Spinoctenus tequendama or slightly rounded.............................................................................8
Tibiae I, II and III and metatarsus III with long ventral setae, metatarsus IV 8(7). Thin copulatory ducts and spermathecae as in Fig. 24E, middle field
modified (Fig. 29A–I) ........................................................................6 of the epigynum with curved margins as in Fig. 24D.........................
6(5). Metatarsus IV strongly curved and with many small spines (Fig. 29H), .......................................................................Spinoctenus stephaniae
internal bulge of the embolus large and rounded (Figs 13A, B, 27I) Vulva with wide copulatory ducts (Figs 15E, 19E, 25E) and spermathecae
.......................................................................... Spinoctenus spinosus otherwise, margins of the middle field of the epigynum slightly curved
Metatarsus IV straight or slightly curved, internal bulge of the embolus (Figs 15D, 19E, 25D) ........................................................................9
smaller or spiniform (Figs 9A, B, 10A, B).........................................7 9(8). Middle field of the epigynum slightly triangular (Fig. 19D),
7(6). Tegulum with elongate projection at the base and embolus curved spermathecae without anterior granules (Fig. 19E) ............................
(Figs 15C, 20C, 25C, 28A, F, K).......................................................8 ........................................................................ Spinoctenus eberhardi
Tegulum without elongate projection at the base (Figs 16C, 18C, 22C, 28B, Middle field of the epigynum slightly quadrangular (Figs 15D, 25D),
D, H) and embolus straight (Figs 10A, B, 11A, 27B, D, H)..............9 spermathecae usually with anterior granules (Figs 15E, 25E)........10
8(7). Metatarsus IV with many small spines (Fig. 29E); elongate projection of 10(9). Spermathecae large and oval-shaped with few anterior granules
the tegulum pronounced (Figs 20C, 28F), median process of the (Fig. 25E), endemic to the Eastern Cordillera.....................................
tegulum long (Figs 11E, 27F); RTA straight and pointed (Figs 11F, .................................................................... Spinoctenus tequendama
20C) ...................................................................... Spinoctenus florezi Spermathecae slightly rounded and small (Fig. 15E), with many anterior
Spination pattern of metatarsus IV as in Fig. 29A, with two large final spines granules, endemic to the Western and Central Cordilleras .................
in the middle region; elongate projection of the tegulum slightly ............................................................................. Spinoctenus yotoco
Spinoctenus yotoco, sp. nov. 76110 2300 W; 1700 m), iv.2012, N. Hazzi (MUSENUV); 1 ,, Pance
(03520 5000 N, 76370 4200 W; 1650 m), viii.2012, N. Hazzi (MUSENUV).
(Figs 7A–D, 8A–D, 9A–D, 15A–E, 26A, 27A, 28A, 29A,
31A, B, 32F)
Diagnosis
http://zoobank.org/urn:lsid:zoobank.org:act:A3D0E9FD-9C79-485A-
Males of Spinoctenus yotoco resemble S. florezi and
AACE-1A887546DFCB
S. tequendama by the elongated projection on the base of the
Material examined tegulum (Figs 15C, 28A), but can be distinguished from them by
the spination pattern on metatarsus IV with two large final spines
Holotype. Colombia: Valle del Cauca: <, Reserva Forestal de Yotoco
in the middle region (Fig. 29A). Females of Spinoctenus yotoco
(03515000 N, 76260 1700 W; 1700 m), xii.2011, N. Hazzi manual capture at
resemble S. eberhardi, but can be distinguished from them
night, on litter in a secondary forest (MUSENUV 27026).
Paratypes. Colombia: Valle del Cauca: 5 <, 5 ,, same data as holotype by a wider and less triangular middle field of the epigynum
(MUSENUV 27027; 1 ,, Chicoral, Yumbo (03330 5900 N, 76340 2500 W; 1900 (Fig. 15D) and by the presence of anterior granules in the
m), iv.2012, N. Hazzi (MUSENUV); Risaralda: 1 <, Estacion La Suiza, spermatheca (Fig. 15E).
Ucumari, Pereira (04430 4800 N, 75340 4800 W; 1850 m), viii.1992,
S. Rodriguez (MPUJ 3138.1). Description
Other material examined. Colombia: Risaralda: 1 <, 1 ,, El Brillante
(5 02 0 5300 N, 7 590 5400 W; 1600 m), xi.2012, L. D. Santa (MAUQ); 1 <, El
Male (MUSENUV 27026)
Tambo (5 010 13N, 7 570 4900 W; 1800 m), viii.2012, L. D. Santa (MAUQ); 2 ,, Total length 15.56. Colour in life (Fig. 31B) as in genus
Verdum (5 000 4100 N, 76 010 5700 W; 2000 m), iv.2012, L. D. Santa 2 description. Carapace 8.06 long, 7.68 wide; ocular area 1.12
(MAUQ); Valle del Cauca: 1 <, El Saladito (03280 5900 N, 76370 0000 W; 2000 long, 2.43 wide, eye diameters AME : ALE : PME : PLE,
m), x.2014, N. Hazzi (MUSENUV); Montañuela, Roldanillo (4270 1800 N, 0.38 : 0.21 : 0.42 : 0.48; AME–AME nearly AME diameter;
(B)
(A)
(C) (D)
Fig. 7. Spinoctenus yotoco. (A) Retrolateral view of the chelicerae; (B) tarsal organ; (C) trichobothria; (D) claw of leg I.
PME separation 0.59 PME diameter; clypeal height 0.14; Paratypes. Colombia: Valle del Cauca: 2 <, 2 ,, same data as
sternum 3.70 long, 3.25 wide; labium 1.25 long, 1.00 wide. holotype (MUSENUV 27051); Chocó: 1 ,, Acandi (90 m), Capurgana,
Leg measurements: I: femur 8.12, patella 3.25, tibia 7.87, Jardin Botanico del Darien, iv.2008, N. Morales (MPUJ-ENT 31385); 1 <,
metatarsus 6.31, tarsus 2.55, total 28.10; II: 6.87, 3.25, 7.44, same locality, iv.2007, C. Duran (MPUJ-ENT 31386).
5.81, 2.12, 25.49; III: 6.19, 2.87, 6.25, 5.31, 1.94, 22.56; IV: 7.50,
2.95, 7.50, 9.87, 3.12, 30.94. Leg spination: tibia I and II v2–2- Diagnosis
2–2-2, r2–2-0, p0; III v2–2-2, r2–2, p2–2; metatarsus I and II
v2–2-2, r1–1-0, p1–1-0; III v2–2-2, r1–1-1–0, p1–1-1; Males of Spinoctenus chocoensis resemble those of S. escalerete
metatarsus IV modified (Fig. 29A). Palp: external margin of and S. pericos by the straight embolus (Figs 10A–C, 16B, 27B),
the embolus curved (Figs 9A, C, 15A, 26A). Tibia with small but differ by the large and spiniform ventral bulge of the embolus
retrolateral sclerotised bulge and RTA with small rounded lobe (Figs 10A, B, 16B, 27B) and by the pattern of spination on
(Fig. 15C). metatarsus IV (Fig. 29B). Females of S. chocoensis resemble
those of S. escalerete by the comma-shaped spermathecae
Female (MUSENUV 27027–1) (Fig. 16E), but they can be distinguished by a wider area
between the two copulatory ducts. Individuals of both sexes
Total length 11.49. Colour in life (Fig. 31A) as in genus are smaller than individuals of S. chocoensis.
description. Carapace 8.5 long, 6.56 wide; ocular area 1.31
long, 2.37 wide; eye diameters AME : ALE : PME : PLE,
0.36 : 0.17 : 0.42 : 0.41 AME diameter; PME separation 0.51 Description
PME diameter; clypeal height 0.13; sternum 3.62 long, 3.19 Male (MUSENUV 27050)
wide; labium 1.25 long, 1.10 wide. Leg measurements: I: femur
6.43, patella 3.12, tibia 6.06, metatarsus 4.69, tarsus 1.87, total Total length 14.94. Colour in life (Fig. 31D) as in genus
16.3; II: 6.19, 3.13, 5.55, 4.69, 1.82, 21.38; III: 5.65, 2.65, 4.50, description. Carapace 8.37 long, 6.25 wide. AME : ALE :
3.00, 1.30, 23.77; IV: 7.00, 3.25, 6.50, 7.15, 2.65, 26.55. Leg PME : PLE 0.32 : 0.22 : 0.47 : 0.45; AME–AME nearly AME
spination: Tibiae I and II v2–2-2–2-2; metatarsi I and II v2–2-2; diameter; PME separation 0.60 PME diameter, sternum 3.55
leg III tibia d1–1-1, v2–2-2, r1–0-1, p1–0-1, metatarsus v2–2-2, long, 3.1 wide; labium 1.25 long, 1.00 wide. Leg measurements:
r0–1-1, p1–1-2; leg IV tibia v2–2-2, p0–2-2–0, r1–1-1–1, d1–1- I: femur 8.62, patella 3.56, tibia 9.31, metatarsus 7.81, tarsus
1, metatarsus v2–2-2, p1–1-2, r1–1-2. Epigynum (Figs 9D, 15D): 2.50, total 31.80; II: 8.00, 3.50, 8.25, 7.25, 2.31, 29.31; III: 7.25,
median sector wide with a posterior depression slightly 3.12, 6.43, 6.87, 2.19, 25.86; IV: 9.50, 3.56, 8.43, 13.44, 2.81,
sclerotised and lateral lobes clearly delimited by heavily 37.74. Leg spination: leg I tibia v2–2-2–2-2, r1–1-0, p0–1-1–0,
sclerotised margins of middle sector, lateral lobes with small metatarsus v2–2-2, r1–1-00, p1–1-0–0; leg II tibia v2–2-2–2-2,
spurs that originate medially and are directed medially. Vulva r1–1-1, p1–1-1, metatarsus v2–2-2, r1–1-0, p1–1-0; leg III tibia
(Fig. 15E): copulatory ducts convergent and spermathecae with d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2,
anterior granules. r1–1-2, p1–1-2; leg IV tibia v2–2-2, d1–1-1, r0–1-1–0, p0–1-
0–1, metatarsus modified (Fig. 29B). Palp: embolus with external
Variation margin straight and large spiniform ventral bulge (Figs 10A–C,
16B, 27B). Tibia with small retrolateral sclerotised bulge and
Females (n = 6): total length 15.85–22.31; carapace 8.00–10.00; RTA with small rounded lobe (Figs 16C, 28C).
femur I 6.19–8.10. Males (n = 5): total length 15.87–18.87;
carapace 8.12–9.62; femur I 8.43–9.44.
Female (MUSENUV 27051–1)
Distribution Total length 14.31. Colour in life (Fig. 31C) as in generic
Western and Central Andean Cordilleras of Colombia (Fig. 36A). description. Carapace 6.75 long, 4.81 wide. AME : ALE :
PME : PLE, 0.32 : 0.22 : 0.40 : 0.37; AME–AME nearly AME
Etymology diameter; PME separation 0.60 PME diameter; sternum 3.00
long, 2.75 wide; labium 1.25 long, 0.94 wide. Leg measurements:
The specific name is a noun in apposition to the Reserva Forestal
I: femur 5.87, patella 3.12, tibia 6.25, metatarsus 4.69, tarsus
Yotoco, in acknowledgement to the people of the Universidad
1.31, total 21.24; II: 5.43, 2.93, 5.50, 4.25, 1.56, 19.68; III: 4.44,
Nacional, who care for and protect this forest.
2.25, 4.12, 4.50, 1.62, 16.93; IV: 6.12, 2.62, 6.00, 7.81, 2.19,
Spinoctenus chocoensis, sp. nov. 24.74. Leg spination: tibiae I and II v2–2-2–2-2; metatarsi I and
II v2–2-2; leg III tibia d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-
(Figs 10A–C, 16A–D, 26B, 27B, 28B, 29B, 36A) 0–2, metatarsus d0–1-0, v2–2-2, r1–1-2, p1–1-2, leg IV tibia
http://zoobank.org/urn:lsid:zoobank.org:act:C47CDEC7-B0F6-4C2B- v2–2-2, d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus d0–1-0-, v2–2-
99D6-7E8E06C41EF4 2–2, r1–1-2, p1–1-2. Epigynum (Fig. 16D): median sector wide,
with a posterior depression slightly sclerotised and lateral lobes
Material examined clearly delimited by heavily sclerotised margins of middle
Holotype. Colombia: Valle del Cauca: <, Reserva Natural Pericos, sector, lateral lobes with small spurs that originate medially
Buenaventura (03500 5500 N, 76470 1400 W; 470 m), v.2015, N. Hazzi and and are directed medially. Vulva (Fig. 16E): copulatory ducts
R. Palacio, manual capture at night, on litter in a secondary forest convergent, spermathecae comma-shaped and with one or two
(MUSENUV 27050). small anterior granules.
(A) (B)
(C) (D)
Fig. 8. Female spinnerets of Spinoctenus yotoco. (A) Overview; (B) anterior lateral spinneret (ALS); (C) posterior median spinneret (PMS); (D) posterior
lateral spinneret (PLS). Abbreviations: AC, aciniform gland spigots; CY, cylindrical gland spigots; MAP, major ampullate gland spigots; mAP, minor ampullate
gland spigots; PI, piriform gland spigots; ta, tartipore.
Variation Material examined

Males (n = 4): total length 14.94–15.37; carapace 8.12–8.43, Holotype. Colombia: Valle del Cauca: <, La Estrella, Sevilla
femur I 7.00–8.43. Females (n = 3): total length 14.12–16.37; (4080 2900 N, 75520 0200 W; 2265 m), xi.2013, N. Hazzi, manual capture at
carapace 6.75–7.93, femur I 5.68–6.44. night, on litter in a secondary forest (MUSENUV 27038).
Paratypes. Colombia: Valle del Cauca: 3 <, 4 ,, same data as holotype
(MUSENUV 27039); 1 male, 2 females, Reserva Natural Cerro del Inglés,
Distribution el Cairo, (4440 3200 N, 76170 3800 W, 2100–2300 m) X. 2016, manual capture
at night, on litter in a primary forest, N.Hazzi, (MUSENUV); Quindio:
Chocó biogeographical region (Fig. 36A).
1 <, Bosque de Bremen, Filandia (04410 0300 N, 75370 0400 W; 1900 m),
IV.1998 (MPUJ 31383), R. Varón; 1 <, same locality, viii.2014, S. Moreno
Etymology (MUSENUV); Risaralda: 1 <, Santuario de Flora y Fauna Otun Quimbaya,
vii.2005, A. Sabogal (04520 N, 75720 W; 2100 m) (Ar-ICN).
The specific name refers to the Chocó biogeographical region in
which the species occurs.
Diagnosis
Spinoctenus eberhardi, sp. nov. Males of Spinoctenus eberhardi can be distinguished from the
remaining species of the genus by the reduced folded process of
(Figs 10E–H, 17A–E, 26C, 27C, 28C, 29C, 33C, D, 36A) the embolus (Figs 10F, G, 17A, 26C), the shape of the internal
http://zoobank.org/urn:lsid:zoobank.org:act:FC009164-41C8-4A17- bulge of the embolus (Figs 10E, F, 17B, 27C) and by the
B017-C6B291D791FB morphology of the retrolateral tibial apophysis (Figs 10H,
(A) (B)
(C) (D)
Fig. 9. Spinoctenus yotoco. (A) Ventral view of male palp; (B) prolateral view of male palp; (C) close-up of bulb; (D) female epigynum, ventral view.
Abbreviations: C, conductor; FP, folded process of the embolus; E, embolus; IB, internal bulge of the embolus; LF, lateral field; LP, lateral process; LL, locking
lobes, median sector; MA, median apophysis; MTP, median tegular process; ST, subtegulum; t, tegulum.
17C, 28C). Females of Spinoctenus eberhardi resemble those of metatarsus 5.56, tarsus 2.37, total 24.49; II: 6.50, 3.10, 6.25,
S. yotoco, but can be distinguished by a more triangular and 4.75, 2.06, 22.66; III: 5.31, 2.56, 5.25, 4.62, 1.62, 19.36; IV: 6.56,
narrower middle field of the epigynum (Fig. 17D) and by the 2.19, 6.81, 8.50, 2.87, 26.93. Leg spination: leg I tibia v2–2-2–2-
absence of anterior granules in the spermathecae (Fig. 17E). 2, r1–1-0, p1–1-0–0, metatarsus v2–2-2; leg II tibia v2–2-2–2-2,
Individuals of both sexes are smaller than individuals of S. yotoco p1–1-1–1-0, r1–1-0, metatarsus v2–2-2, r1–1-0, p1–1-0; leg III
(see morphometric measurements in the ‘Variation’ section). tibia, d1–1-1, v2–2-2, r0–1-1–0 p0–1-1–0, metatarsus d0–1-0,
v2–2-2, r1–1-2, p1–1-2; leg IVtibia d1–1-1, v2–2-2, r0–1-0–1,
Description p0–1-0–1, metatarsus modified (Fig. 29D). Palp: embolus with
Male (MUSENUV 27038) folded process reduced (Figs 10F, G, 17A, 26C), external margin
curved and internal bulge triangular (Figs 10E, F, 17B, 27C).
Total length 12.50. Colour in life (Fig. 33D) as in genus RTA without small rounded lobe (Figs 10H, 17C, 28C).
description. Carapace 6.75 long, 5.62 wide; ocular area 1.08
long, 1.92 wide; eye diameters AME : ALE : PME : PLE,
0.31 : 0.20 : 0.35 : 0.38; AME–AME nearly AME diameter; Female (MUSENUV 27039–1)
PME separation 0.66 PME diameter; clypeal height 0.25; Total length 14.68. Colour in life (Fig. 33C) as in genus
sternum 3.00 long, 2.62 wide; labium 1.00 long, 0.81 wide. description. Carapace 6.87 long, 5.81 wide; ocular area 1.20
Leg measurements: I: femur 6.50, patella 3.25, tibia 6.81, long, 2.37 wide. Eye diameters AME : ALE : PME : PLE,
(A) (B)
(C) (D)
(E) (F)
(G) (H)
Fig. 10. (A–C) Spinoctenus chocoensis, (D) Phoneutria boliviensis and (E–H) S. eberhardi. (A) ventral view of bulb;
(B) close-up of bulb; (C) prolateral view of bulb; (D) ventral view of bulb; (E) ventral view of bulb; (F) close-up of
bulb; (G) prolateral view of bulb; (H) retrolateral tibial apophysis. Abbreviations: C, conductor; FP, folded process of the
embolus; E, embolus; IB, internal bulge of the embolus; LL, locking lobes; MA, median apophysis; MTP, median tegular
process; ST, subtegulum; PRT, probasal rounded tegulum; t, tegulum.
(A) (B)
(C) (D)
(E) (F )
Fig. 11. (A–D) Spinoctenus escalerete and (E, F) S. florezi. (A) Ventral view of bulb; (B) prolateral view of bulb, arrow
indicates the pointed folded process of the embolus; (C) close-up of bulb; (D) retrolateral tibial apophysis, arrow indicates small
rounded lobe; (E) ventral view of bulb; (F) retrolateral tibial apophysis, arrow indicates small rounded lobe. Abbreviations:
C, conductor; FP, folded process of the embolus; E, embolus; IB, internal bulge of the embolus; LL, locking lobes; MA, median
apophysis; MTP, median tegular process; RSB, retrolateral sclerotised bulge; ST, subtegulum; t, tegulum.
0.26 : 0.21 : 0.38 : 0.35; AME–AME nearly AME diameter; PME r1–1-2, p1–1-2. Epigynum (Fig. 17D): median sector wide with a
separation 0.60 PME diameter; clypeal height 0.35; sternum posterior depression slightly sclerotised and lateral lobes clearly
3.12 long, 2.81 wide; labium 1.12 long, 0.94 wide. Leg delimited by heavily sclerotised margins of middle sector, lateral
measurements: I: femur 6.00, patella 3.12, tibia 5.87, lobes with small spurs that originate medially and are directed
metatarsus 4.37, tarsus 1.68, total 21.04; II: 5.69, 2.75, 5.37, medially. Vulva (Fig. 17E): copulatory ducts convergent.
4.19, 1.69, 19.69; III: 5.00, 2.62, 4.25, 3.62, 1.62, 17.11; IV: 6.18,
2.75, 5.81, 6.50, 2.31, 23.55. Leg spination: tibiae I and II v2–2- Variation
2–2-2; metatarsi I and II v2–2-2; leg III tibia d1–1-1, v2–2-2–2, Males (n = 3): total length 10.93–12.65; carapace 5.93–7.00;
r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, r1–1-2, p1–1-2; leg IV femur I 5.95–6.75. Females (n = 3): total length 12.68–14.62;
tibia v2–2-2, d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus v2–2-2–2, carapace 6.12–6.75; femur I 4.62–5.50.
(A) (B)
(C) (D)
(E) (F )
(G) (H)
Fig. 12. (A–D) Spinoctenus nambi and (E–H) S. pericos. (A) Ventral view of bulb; (B) prolateral view of close-up of
bulb; (C) close-up of bulb; (D) retrolateral tibial apophysis; (E) ventral view of bulb; (F) prolateral view of bulb; (G) close-
up of bulb; (H) retrolateral tibial apophysis, arrow indicates small rounded lobe. Abbreviations: C, conductor; FP, folded
process of the embolus; E, embolus; IB, internal bulge of the embolus; LL, locking lobes; MA, median apophysis; MTP,
median tegular process; ST, subtegulum; t, tegulum.
(A) (B)
(C) (D)
(E) (F)
(G) (H)
Fig. 13. (A–D) Spinoctenus spinosus; (E–H) S. stephaniae. (A) Ventral view of bulb; (B) close-up of bulb; (C) prolateral
view of bulb; (D) retrolateral tibial apophysis, arrow indicates small rounded lobe; (E) ventral view of bulb; (F) prolateral
view of bulb; (G) retrolateral tibial apophysis; (H) close-up of bulb. Abbreviations: C, conductor; FP, folded process of the
embolus E, embolus; IB, internal bulge of the embolus; LL, locking lobes; MA, median apophysis; MTP, median tegular
process; RSB, retrolateral sclerotised bulge; ST, subtegulum; t, tegulum.
Distribution 5.00, 53.80. Leg spination: leg I tibia v2–2-2–2-2, r2–2-0, p1–1-
Central Andean Cordillera of Colombia (Fig. 36A). 1–0, metatarsus v2–2-2, r1–1-0, p1–0-0; leg II tibia v2–2-2–2-2,
r-0–2-2–0, p2–2-0, metatarsus v2–2-2, r2–2-0, p2–2-0; leg III
Etymology tibia v2–2-2, r2–0-2, p0–2-2, metatarsus v2–2-2, r1–1-1, p1–1-1;
leg IV tibia v2–2-2, r0–1-1–0, p0–1-1–0, metatarsus IV modified
The specific name is in honour of the arachnologist William (Fig. 29C). Palp: tegulum with a ventral sclerotised process
Eberhard, in acknowledgement of his support and teachings at the base near the median apophysis (Fig. 11B, C), external
about arachnid behaviour to the first author. margin of the embolus straight (Figs 10A, 16B, 27B), with folded
process pointed in the middle (Figs 11B, 18A, 26D); tibia with
Spinoctenus escalerete, sp. nov. small retrolateral sclerotised bulge and RTA with small rounded
(Figs 11A–D, 14C–E, 18A–E, 26D, 27D, 28D, 29C, 30A, lobe (Figs 11D, 18C).
32A, B, 36B)
http://zoobank.org/urn:lsid:zoobank.org:act:3E48102B-FD04-4F9A-
B791-60842D19DF6C Total length 23.00. Colour in life (Fig. 32C): similar to male,
carapace dark brown with a pale brown longitudinal line; dorsum
Material examined with a folium-like golden longitudinal band to middle region.
Holotype. Colombia: Valle del Cauca: <, Reserva San Cipriano, Carapace 12.85 long, 9.00 wide; ocular area 1.90 long, 3.60 wide;
Buanaventura (03520 4700 N, 76550 3200 W; 370 m), ii.2012, N. Hazzi and eye diameters AME : ALE : PME : PLE, 0.40 : 0.25 : 0.55 : 0.60;
J. Moreno, on litter in a secondary forest (MUSENUV 27029). AME–AME nearly AME diameter; PME separation 0.63 PME
Paratypes. Colombia: Valle del Cauca: 2 ,, same data as holotype diameter; clypeal height 0.60; sternum 4.60 long, 4.35 wide;
(MUSENUV 27031); 1 <, 7 ,, Reserva Natural Pericos, i.2014, N. Hazzi, labium 2.50 long, 2.10 wide. Leg measurements: I: femur 10.01,
R. D. Palacio and S. Moreno, Buenaventura (03500 5500 N, 76470 1400 W; 470 patella 4.25, tibia 9.40, metatarsus 7.90, tarsus 2.75, total 34.31;
m), manual capture at night, on litter in a secondary forest (MUSENUV
II: 10.15, 4.75, 9.25, 6.45, 3.00, 33.60; III: 8.50, 4.25, 7.50, 7.25,
27031); 3 ,, same locality, xi.2014, N. Hazzi, R. D. Palacio and S. Moreno
(MUSENUV 27032); Buanaventura: 1 <, Bajo Calima (03570 1200 N,
2.25, 29.75; IV: 9.75, 4.25, 9.50, 11.50, 3.40, 38.40. Leg
76590 2600 W; 40 m), iii.1995, C. Morales (MPUJ 31384). spination: tibiae I and II v2–2-2–2-2; metatarsi I and II v2–2-
2; leg III tibia d0–1-1–0, v2–2-2, p0–2-2–0, r0–2-2–0, metatarsus
Diagnosis v2–2-2, p1–1-2, r1–1-2; leg IV tibia d1–1-1, v2–2-2, r0–1-1–0,
p0–1-1–0, metatarsus d0–1-0 v2–2-2, p1–1-2, r1–1-2. Epigynum
Males of Spinoctenus escalerete resemble those of S. pericos and (Figs 14C, 18D): median sector wide, with a posterior depression
S. chocoensis by the straight external margin of the embolus slightly sclerotised and lateral lobes clearly delimited by heavily
(Figs 11A, B, 18B, 27D), but differ by the folded process pointed sclerotised margins of the middle sector, lateral lobes with small
in the middle (Figs 11B, 18A, B, 26B), by a ventral sclerotised spurs that originate medially and are directed medially. Vulva
tegular process near the median apophysis (Figs 18C, 28B) and (Figs 14C–E, 18E): copulatory ducts convergent, spermathecae
by the curved IV metatarsus (Fig. 29C) with a distal large hook- comma-shaped and with one or two small anterior granules.
shaped spine. Females of Spinoctenus escalerete resemble those
of S. chocoensis by the comma-shaped spermathecae (Figs 14D,
Variation
E, 18D), but can be distinguished by a narrow middle region of
the vulva between the two copulatory ducts (Figs 14D, 18E). Females (n = 7): total length 17.60–23.30; carapace 10.10–10.62;
Individuals of both sexes are larger than individuals of femur I 8.74–9.19.
S. chocoensis (see morphometrics measures in the ‘Variation’
section). Distribution
Pacific region of Valle del Cauca, Colombia (Fig. 36B).
Description
Male (MUSENUV 27029) Etymology
Total length 19.00. Colour in life (Fig. 32D): carapace dark The specific name is a noun in apposition to the Escalerete River,
purple with a longitudinal pale yellow median line in the thoracic which passes through the wet forest near San Cipriano, and is
region, black pigment around eyes, chelicerae black, legs pale protected by the native community in that locality.
brown except femur dark brown; ventral faces of coxae, endites
and labium brown; abdomen black, dorsum with a folium-like Spinoctenus flammigerus, sp. nov.
golden longitudinal band, venter brown with four divergent
series of white dots. Carapace 12.00 long, 10.00 wide; eye (Figs 19A–E, 26E, 27E, 28E, 32C, D, 36A)
diameters AME : ALE : PME : PLE, 0.40 : 0.25 : 0.55 : 0.60; http://zoobank.org/urn:lsid:zoobank.org:act:8349EFEC-0928-4B92-
AME–AME nearly AME diameter; PME separation 0.63 A29A-9F437B67ACFB
PME diameter; clypeal height 0.30; sternum 5.00 long, 4.40
wide; labium 1.65 long, 1.30 wide. Leg measurements: I: femur Material examined
12.55, patella 5.40, tibia 13.30, metatarsus 11.30, tarsus 4.00, Holotype. Colombia: Nariño: <, Reserva Natural Rio Ñambi, Altaquer,
total 46.55; II: 14.10, 5.40, 13.75, 11.25, 3.80, 48.30; III: 12.20, Barbacoas (01180 N, 78050 W; 1400 m), x.2012, M. Medrano, A. Garcia,
4.50, 11.25, 15.80, 3.50, 46.25; IV: 13.50, 4.75, 13.05, 17.50, Y. Cifuentes and D. Martinez (MUSENUV 27129).
(A) (B)
(C) (D)
(E) (F )
Fig. 14. Genitalia of Spinoctenus spp. Spinoctenus pericos. (A) Epigynum, ventral view; (B) vulva, ventral view. Spinoctenus
escalerete. (C) Epigynum, ventral view; (D) vulva, ventral view; (E) close-up of spermatheca, arrow indicated small granule.
Spinoctenus pericos. (F) Close-up of spermatheca. Abbreviations: CD, copulatory duct; FD, fertilisation ducts; LF, lateral field;
LP, lateral process; MS, median sector; S, spermatheca.
Paratypes. Colombia: Nariño: 3 ,, same data as holotype margin (Figs 19C, 26E). Females can be distinguished by the
(MUSENUV 27130). triangular shape of the middle field of the epigynum (Fig. 19D)
and by the lack of anterior granules in the spermathecae
(Fig. 19E).
Diagnosis
Both sexes of S. flammigerus resemble S. nambi by the orange Description
and reddish lines on the dorsum of the body (Fig. 32C, D). Males
of S. nambi resemble those of S. flammigerus by the shape of the Male (MUSENUV 27129)
embolus, but can be distinguished by the smaller membranous Total length 18.74. Colour in life (Fig. 32C): similar to
internal margin of the embolus, which is absent towards the tip of S. nambi. Carapace 10.06 long, 7.62 wide. AME : ALE :
the embolus (Figs 19B, 27E), and by the thinner external lateral PME : PLE, 0.44 : 0.25 : 0.50 : 0.56; AME–AME nearly AME
(A) (B) (C)
(D) (E)
Fig. 15. Spinoctenus yotoco, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Spinoctenus yotoco, female
genitalia. (D) Epigynum, ventral view; (E) vulva, dorsal view.
diameter; PME separation 0.606 PME diameter; sternum 4.19 Palp: embolus with folded process thin and curved, without
long, 3.57 wide; labium 1.31 long, 1.00 wide. Leg measurements: small membranous whitish internal margin towards the tip
I: femur 10.31, patella 4.18, tibia 10.94; metatarsus 9.37, tarsus (Figs 19B, 27E) and thin external lateral margin (Figs 19A, 26E).
3.56, total 38.36; II: 10.00, 4.25, 10.06, 8.75, 3.25, 36.31; III:
8.44, 3.12, 7.50, 7.62, 2.87, 29.55; IV: 11.00, 3.47, 10.31, 13.44,
4.12, 42.34. Leg spination: tibia I d0–0-1, v2–2-2–2-2, r1–1-0, Female (MUSENUV 27130–1)
p1–1-0–0, metatarsus v2–2-2, r1–1-0, p1–1-0–0; leg II tibia Total length 19.80. Colour in life (Fig. 32D): as in S. nambi.
d1–1-1–0, v2–2-2–2-2, r1–1-0, p1–1-0–0, metatarsus v2–2-2, Carapace 10.12 long, 7.75 wide. AME : ALE : PME : PLE,
r1–1-0, p1–1-0, leg III tibia d1–1-1, v2–2-2–2, r0–2-0–2, p0–2- 0.32 : 0.25 : 0.37 : 0.50; PME separation 0.606 PME
0–2, metatarsus v2–2-2, r1–1-2, p1–1-2; leg IV tibia v2–2-2, diameter; sternum 4.25 long, 3.65 wide; labium 1.25 long,
d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus IV modified (Fig. 29F). 1.10 wide. Leg measurements: I: femur 8.43, patella 4.37,
(B) (C)
(A)
(D) (E)
Fig. 16. Spinoctenus chocoensis, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Spinoctenus chocoensis,
female genitalia. (D) Epigynum, ventral view; (E) vulva, dorsal view.
tibia 8.43, metatarsus 6.31, tarsus 2.25, total; II: 8.12, 4.25, 5.12, Vulva (Fig. 19E): copulatory ducts convergent and spermathecae
5.94, 2.06, total; III: 6.70, 3.31, 6.37, 5.94, 2.37, total; IV: without granules.
9.25, 3.81, 1.30, 8.12, 2.87, total. Leg spination: as tibiae I
Distribution
and II v2–2-2–2-2; metatarsi I and II v2–2-2; leg III tibia
d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, Nariño Department, Colombia (Fig. 36A).
v2–2-2, r1–1-2, p1–1-2; leg IV tibia v2–2-2, d1–1-1, r0–1-
1–0, p0–1-0–1, metatarsus d1–1-0–0, v2–2-2–2, r1–1-2, Etymology
p1–1-2. Epigynum (Fig. 19D): triangular median sector with a The specific name is a combination of ‘flamma’ and ‘igo’,
posterior depression, slightly sclerotised and lateral lobes clearly derived from the Latin, which mean ‘carries a flame’,
delimited by heavily sclerotised margins of middle sector, lateral referring to the orange and red colours on the dorsum of
lobes with small spurs originated medially and directed medially. males and females.
(A) (B) (C)
(D) (E)
Fig. 17. Spinoctenus eberhardi, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia.
(D) Epigynum, ventral view; (E) vulva, dorsal view.
Spinoctenus florezi, sp. nov. Paratypes. Colombia: Cundinamarca: ,, same data as holotype
(ICN-AR-7925); 2 ,, Toquiza, Medina (04360 N, 73240 W; 1000 m),
(Figs 11E–H, 20A–E, 26F, 27F, 28F, 29E, 36A) P. Perez (MPUJ 0031388).
http://zoobank.org/urn:lsid:zoobank.org:act:6A0D4258-66D7-482B-
B88C-AAF6D4BED89B
Diagnosis
Material examined Males of Spinoctenus florezi resemble those of S. yotoco and
Holotype. Colombia: Cundinamarca: <, Alban (04520 N, 74290 W; S. bogotensis by the elongated projection at the base of
1500 m), E. Florez (ICN-AR-7924). the tegulum, although in S. florezi it is considerably more
(A) (B) (C)
(D) (E)
Fig. 18. Spinoctenus escalerete, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia.
(D) Epigynum, ventral view; (E) vulva.
pronounced (Figs 20C, 28F). Males of S. florezi also have a larger retrolateral face of metatarsus IV (Fig. 29E). Females of
sclerotised tubercle under the retrolateral tibial apophysis Spinoctenus florezi can be distinguished from the remaining
(Figs 20C, 28F) than the rest of the species and a small peak species by the unique morphology of the middle field of
in the apex of the folded process of the embolus (Figs 11E, 20A, the epigynum, with a narrowed base, wider anteriorly and
26F). In addition, they have large and numerous spines on the with a large non-sclerotised depression in the centre (Fig. 20D).
(A) (B) (C)
(D) (E)
Fig. 19. Spinoctenus flammigerus, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia. (D) Epigynum,
ventral view; (E) vulva, dorsal.
(A) (B) (C)
(D) (E)
Fig. 20. Spinoctenus florezi, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia. (D) Epigynum,
ventral view; (E) vulva, dorsal view. Abbreviations: CD, copulatory duct.
Description 31.86; IV: 11.06, 3.75, 10.81, 14.56, 4.19, 44.37. Leg
spination: leg I tibia v2–2-2–2-2, r1–1-0, p1–1-1–1,
Male (ICN-AR-7924) metatarsus v2–2-2, p1–1-0–0; leg II tibia v2–2-2–2-2, r1–1-0,
Coloration as in the genus description. Total length 18.44. p1–1-1–1, metatarsus v2–2-2, r1–1-0, p1–1-0; leg III tibia
Carapace 9.69 long, 7.75 wide. Eye diameters AME : ALE : d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2,
PME : PLE, 0.41 : 0.30 : 0.50 : 0.52; AME–AME nearly AME r1–1-2, p1–1-2; leg IV tibia v2–2-2, d1–1-1, r0–1-1–0, p0–1-
diameter; PME separation 0.536 PME diameter; sternum 0–1, metatarsus IV modified (Fig. 29H). Palp: tegulum with
4.12 long, 3.44 wide; labium 1.31 long, 1.18 wide. Leg elongated projection at base (Figs 20C, 28F), external margin of
measurements: I: femur 9.87, patella 4.06, tibia 11.37, embolus curved (Figs 11E, 20B, 28F). Tibia with pronounced
metatarsus 10.25, tarsus 3.81, total 39.36; II: 10.00, 4.06, retrolateral sclerotised bulge and RTA with small rounded lobe
10.31, 8.87, 3.12, 36.36; III: 8.87, 3.31, 8.31, 8.06, 3.31, (Figs 11F, 20C, 28F).
Female (ICN-AR-7925) (Fig. 21D) and by the presence of anterior granules in the
Coloration: as in genus description. Total length 19.87. spermathecae (Fig. 21E).
Carapace 9.75 long, wide 7.81. AME : ALE : PME : PLE
0.40 : 0.32 : 0.53 : 0.53; AME–AME nearly AME diameter; Description
PME separation 0.76 PME diameter; sternum 4.06 long, Male (Ar-ICN 7922)
3.43 wide; labium 1.25 long, 1.20 wide. Leg measurements: I:
femur 8.12, patella 3.81, tibia 8.06, metatarsus 6.43, tarsus 2.31, Total length 14.93. Coloration in live specimens as in
total 2.87; II: 7.75, 4.06, 7.50, 6.12, 2.06, 27.49; III: 6.50, 3.19, S. flammigerus. Carapace 7.81 long, 7.00 wide; ocular area
5.31, 5.62, 2.06, 22.68; IV: 8.43, 3.44, 7.62, 9.75, 2.62, 31.86. 1.50 long, 2.25 wide. Eye diameters AME : ALE : PME : PLE,
Leg spination: tibiae I and II v2–2-2–2-2; metatarsi I and II v2–2- 0.36 : 0.25 : 0.38 : 0.43; AME–AME nearly AME diameter;
2; leg III tibia d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus PME separation 0.52 PME diameter; clypeal height 0.30;
d0–1-0, v2–2-2, r1–1-2, p1–1-2; leg IV tibia v2–2-2, d1–1-1, sternum 3.94 long, 3.37 wide; labium 1.37 long, 1.06 wide.
r0–1-1–0, p0–1-0–1, metatarsus d0–1-0-, v2–2-2–2, r1–1-2, Leg measurements: I: femur 9.31, patella 3.56, tibia 10.12,
p1–1-2. Epigynum (Fig. 20D): median sector with lateral metatarsus 7.81, tarsus 3.00, total 33.81; II: 9.31, 4.00, 8.56,
margins curved and with anterior area less sclerotised and 7.50, 3.00, 32.37; III: 8.00, 3.19, 7.50, 6.87, 2.75, 28.31; IV:
deeply excavated, lateral lobes clearly delimited by heavily 10.12, 3.44, 9.50, 11.12, 3.56, 37.74. Leg spination:As
sclerotised margins of middle sector, lateral lobes with small S. flammigerus. Palp: embolus curved and thin, without
spurs that originate medially and are directed medially. Vulva internal bulge (Figs 12A–C, 21B, 27G), folded process with
(Fig. 20E): copulatory ducts divergent and spermathecae uniform shape and external margin curved (Figs 12B, 21A, 26G).
rounded, without granules. Tibia with small retrolateral sclerotised bulge and RTA
triangular (Figs 12D, 21C, 28G).
Variation
Males (n = 3): total length 17.31–18.44; carapace 8.69–9.69; Female (Ar-ICN 7923–1)
femur I 9.87–11.06. Colour in life (Fig. 32E): carapace dark brown with two
longitudinal oranges lines in the cephalic region and usually three
Distribution radial oranges lines in the thoracic region, black pigment around
Eastern Andean Cordillera of Colombia (Fig. 36A). eyes; chelicerae, legs and sternum brown; coxae brown with
white spots; abdomen orange, dorsum with a folium-like golden
Etymology longitudinal band surrounded by black dots, venter brown with
two lateral divergent series of yellow dots. Total length 20.69.
The specific name is in honour of the arachnologist Eduardo Carapace 8.50 long, 7.44 wide. AME : ALE : PME : PLE,
Flórez, in acknowledgement of his unconditional support and 0.33 : 0.25 : 0.38 : 0.40; AME–AME nearly AME diameter;
encouragement of the first author in the study of arachnids. PME separation 0.50 PME diameter; sternum 3.31 long,
2.87 wide; labium 1.00 long, 1.00 wide. Leg measurements: I:
Spinoctenus nambi, sp. nov. femur 6.31, patella 3.31, tibia 6.25, metatarsus 4.62, tarsus 1.75,
(Figs 12A–D, 21A–E, 26G, 27G, 28G, 29F, 32E, 36A) 22.24; II: 6.06, 3.25, 6.00, 4.68, 1.81, 21.80; III: 5.37, 2.62, 5.00,
http://zoobank.org/urn:lsid:zoobank.org:act:8FCD53B5-DD3F-4367-
4.44, 1.81, 19.24; IV: 6.87, 2.87, 6.87, 7.81, 2.37, 26.79. Leg
AB74-414B4D916DB5 spination: As S. flammigerus. Epigynum (Fig. 21D): median
sector wide, with a posterior depression slightly sclerotised and
Material examined lateral lobes clearly delimited by heavily sclerotised margins
Holotype. Colombia: Nariño: <, Reserva Natural Rio Ñambi, Altaquer,
of middle sector, lateral lobes with small spurs originated
Barbacoas (01180 N, 78050 W; 1400 m), vii.2012, M. Medrano, A. Garcia, medially and directed medially. Vulva (Fig. 21E): copulatory
Y. Cifuentes and D. Martinez (Ar-ICN 7922). ducts convergent and spermathecae with anterior granules.
Paratypes. Colombia: Nariño: 3 ,, same data as holotype (Ar-ICN
7923); 2 ,, same locality as holotype, x.2012, D. Martinez, C. Perafan, Variation
C. Castellanos, manual capture at night, on litter in a secondary forest
(MUSENUV 27048). Females (n = 4): total length 15.99–19.12; carapace 7.50–9.62;
femur I 5.94–7.20.
Diagnosis
Both sexes of S. nambi resemble S. flammigerus by the orange Distribution
and reddish lines on the dorsum of the body (Fig. 32E). Males Nariño Department, Colombia (Fig. 36A).
of S. nambi resemble those of S. flammigerus by the shape of
the embolus (Figs 21B, 27G), but can be distinguished by the
whitish membranous internal margin of the embolus that is Etymology
larger towards the tip and by the wider external lateral margin The specific name is a noun in apposition to the Reserva Ñambi,
(Figs 21A, 26E). Females can be distinguished by the in acknowledgement to the people of Fundacion Felca, who care
quadrangular shape of the middle field of the epigynum for and protect this forest.
(A) (B) (C)
(E)
(D)
Fig. 21. Spinoctenus nambi, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia. (D) Epigynum,
ventral view; (E) vulva, dorsal.
Spinoctenus pericos, sp. nov. Material examined

(Figs 12E–H, 14A, B, F, 22A–E, 26H, 27H, 28H, 29G, 33E, F) Holotype. Colombia: Valle del Cauca: <, Reserva Natural Pericos,
Buenaventura (03500 5500 N, 76470 1400 W; 470 m), i.2014, N. Hazzi,
http://zoobank.org/urn:lsid:zoobank.org:act:0B3422BC-F093-42FB- R. D. Palacio and S. Moreno, manual capture at night, on litter in
8959-A2AAFB03B7E5 a secondary forest (MUSENUV 27034).
(A) (B) (C)
(E)
(D)
Fig. 22. Spinoctenus pericos, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia.
Paratypes. Colombia: Valle del Cauca: 2 ,, same data as holotype of the retrolateral tibial apophysis (Figs 12H, 22C, 28H) and
(MUSENUV 27035); 2 ,, same data except xi.2015 (MUSENUV 27036); by the straight, spiny metatarsus IV (Fig. 29C). Females of
1 <, same data except v.2015 (MUSENUV 27037). Spinoctenus pericos differ by the rounder spermathecae
(Figs 14B, F, 22E) and by the triangular middle field of the
Diagnosis epigynum with straight margins (Figs 14A, 22E).
Males of Spinoctenus pericos resemble those of S. escalerete and Description
S. chocoensis by the straight external margin of the embolus, but
differ by the morphology of the embolus, with a small external Male (MUSENUV 27034)
distal bulge (Figs 12E, 22B, 27H) and by a thinner, small and Total length 23.00. Colour in life (Fig. 33F): carapace dark
curved folded process (Figs 12F, 22A, 26H), by the morphology brown carapace, black pigment around eyes, chelicerae black;
legs pale brown, except femur dark brown; ventral face of Etymology
coxae pale orange, endites and labium dark brown, abdomen The specific name is a noun in apposition to Pericos Stream,
black, dorsum with a folium-like golden longitudinal band, which passes through the wet forest of the Reserva Pericos.
venter brown with two lateral divergent series of yellow
dots and two medial large dots. Carapace 12.50 long, 11.00 Spinoctenus spinosus, sp. nov.
wide; ocular area 2.00 long, 62.95 wide; eye diameters
AME : ALE : PME : PLE, 0.50 : 0.30 : 0.55 : 0.55; AME–AME (Figs 13A–D, 23A–E, 26I, 27I, 28I, 29H, 33A, B)
nearly AME diameter; PME separation 0.60 PME diameter; http://zoobank.org/urn:lsid:zoobank.org:act:0E8395A3-1861-4AF8-
clypeal height 0.60; sternum 6.25 long, 4.80 wide; labium 1.90 AE4F-9827EC42EA01
long, 1.60 wide. Leg measurements: I: femur 14.25, patella 6.00,
tibia 14.00, metatarsus 13.10, tarsus 4.50, total 51.85; II: 14.25, Material examined
6.00, 13.90, 13.25, 4.30, 51.70; III: 12.25, 5.15, 12.25, 12.00, Holotype. Colombia: Valle del Cauca: < from Parque Natural Regional
3.75, 45.4; IV: 15.00. 5.50, 14.70, 18.75, 4.70, 58.65. Leg ‘El Vinculo’, Buga (03510 5000 N, 76260 1700 W; 950 m), viii.2013, N. Hazzi,
spination: leg I tibia d1–1-1–1-1, v2–2-2–2-2, r1–1-1–1-1, manual capture at night, on litter in a secondary forest (MUSENUV 27044).
p1–1-1–1-1–1, metatarsus v2–2-2, p1–1-0–0, r1–1-0–0; leg II Paratypes. Colombia: Valle del Cauca: 3 ,, same data as holotype
tibia d1–1-1–1-0, v2–2-2–2-2, p1–1-1–1-1–1-1, r1–1-1–1, (MUSENUV 27045); 1 ,, Mateguada (04010 2900 N, 76090 4500 W; 1127 m),
metatarsus v2–2-2, r1–1-0–0, p1–1-0–0; leg III tibia d1–1-1, ii.2014, S. Moreno (MUSENUV 27046); 2 <, 1 ,, El Dovio (04330 4000 N,
v2–2-2, p0–1-1–1, r1–1-1–1, p1–1-1; leg IV d1–1-1, v2–2-2, 76160 3800 W; 1190 m), ix.2013, N. Hazzi (MUSENUV 27047).
Other material examined. Colombia: Cauca: 1 <, 2 ,, Caloto
r2–2-2–2, p 1–1-1, metatarsus IV modified (Fig. 29G). Palp:
(2590 4900 N, 76240 3500 W; 1200 m), ix.2009, N. Muriel (MUSENUV).
external margin of the embolus straight with distal small bulge
in the embolus (Figs 12E, 22B, 27H), folded process thinner,
Diagnosis
small and curved (Figs 11F, 22C, 26H); tibia without small
retrolateral sclerotised bulge and RTA with small rounded lobe Males of Spinoctenus spinosus can be easily recognised
(Figs 11H, 22C, 28H). from other species by the large internal bulge of the embolus
(Figs 13A, B, 23B, 27I) and by the strongly curved metatarsus IV
with numerous small spines (Fig. 29H). Females of Spinoctenus
Female (MUSENUV 27035–1) spinosus resembles those of S. yotoco by the presence of
Total length 27.75. Colour in life (Fig. 33E): carapace greyish anterior granules in the spermatheca, but can be distinguished
brown with a longitudinal pale brown longitudinal median line by the curved shaped of the lateral margins of the middle field
in the thoracic region, legs reddish brown, sternum and labium of the epigynum (Fig. 23D) and by the non-parallel disposition
bluish black, coxae with pale yellow spots; abdomen greyish of the copulatory ducts (Fig. 23E).
dorsally, venter as in male. Carapace 13.5 long, 10.5 wide;
ocular area 1.95 long, 3.65 wide; eye diameters AME : ALE : Description
PME : PLE, 0.50 : 0.40 : 0.50 : 0.60; AME–AME nearly AME Male (MUSENUV 27044)
diameter; PME separation 0.60 PME diameter; clypeal
height 0.65; sternum 5.65 long, 4.70 wide; labium 1.90 long, Total length 16.87. Colour in life (Fig. 33B) as in the genus
1.85 wide. Leg measurements: I: femur 11.75, patella 5.0, tibia description. Carapace 9.37 long, 7.56 wide; ocular area 1.19
12.00, metatarsus 9.50, tarsus 3.60, total 41.85; II: 12.50, 5.25, long, 2.50 wide; eye diameters AME : ALE : PME : PLE,
11.00, 9.35, 3.50, 41.60; III: 10.50, 5.00, 9.00, 9.50, 3.05, 34.00; 0.35 : 0.28 : 0.55 : 0.57; AME–AME nearly AME diameter;
IV:13.25, 5, 12.50, 14.50, 3.90, 49.40. Leg spination: tibiae I PME separation 0.50 PME diameter; clypeal height 0.28;
and II v2–2-2–2-2; metatarsi I and II v2–2-2; leg III tibia d1–1-1, sternum 4.43 long, 3.50 wide; labium 1.00 long, 1.12 wide.
v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, r1–1-2, Leg measurements: I: femur 9.12, patella 4.37, tibia 8.75,
p1–1-2; leg IV tibia v2–2-2, d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus 7.81, tarsus 2.68, 32.73; II: 9.18, 4.00, 9.25, 7.69,
metatarsus v2–2-2–2, r1–1-1, p1–1-2. Epigynum (Figs 14A, 2.50, 32.62; III: 8.12, 3.62, 7.87, 6.75, 2.25, 28.61; IV: 9.62, 4.37,
22D): median sector more or less triangular with a posterior 12.06, 10.93, 3.68, 40.66. Leg spination: leg I tibia v2–2-2–2-2,
depression slightly sclerotised and lateral lobes clearly delimited r1–1-0, p1–1-0–0, metatarsus v2–2-2, r1–1-0, p1–1-0–0; leg II
by heavily sclerotised margins of the middle sector, lateral tibia d1–0-1–0, v2–2-2–2-2, r1–1-0, p1–1-0–0, metatarsus v2–2-
lobes with small spurs that originate medially and are directed 2, r1–1-0, p1–1-0; leg III tibia d1–1-1, v2–2-2–2, r0–2-0–2,
medially. Vulva (Figs 14B, F, 22E): copulatory ducts convergent p0–2-0–2, metatarsus v2–2-2, r1–1-2, p1–1-2; leg IV tibia v2–2-
and spermathecae rounded without anterior granules. 2, d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus IV modified
(Fig. 29H). Palp: tibia with retrolateral sclerotised bulge and
RTA with small rounded lobe (Figs 13D, 23C, 28I). Embolus
Variation with external margin curved and pronounced ventral bulge
Females (n = 3): total length 22.75–24.50; carapace 12.25–12.62; (Figs 13A–C, 23B, 26I).
femur I 10.62–10.75.
Total length 18.49. Coloration in life (Fig. 33A) as in genus
Distribution description. Carapace 8.81 long, 7.50 wide; ocular area1.37
Pacific region of Valle del Cauca, Colombia (Fig. 36B). long, 3.00 wide; eye diameters AME : ALE : PME : PLE,
(A) (B) (C)
(D) (E)
Fig. 23. Spinoctenus spinosus, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia.
(D) Epigynum, ventral view; (E) vulva, dorsal view. Abbreviations: CD, copulatory duct.
0.33 : 0.33 : 0.46 : 0.50; AME–AME nearly AME diameter; metatarsus 5.43, tarsus 1.87, total 25.54; II: 7.25, 3.56, 6.93,
PME separation 0.70 PME diameter; clypeal height 0.43; 5.00, 1.87, 24.61; III: 5.62, 3.40, 5.00, 4.25, 1.50, 19.77; IV: 8.12,
sternum 4.00 long, 3.18 wide; labium 1.35 long, 1.37 wide. 3.50, 7.81, 7.50, 8.12, 2.37, 37.42; Leg spination: tibiae I
Leg measurements: I: femur 7.31, patella 3.75, tibia 7.18, and II v2–2-2–2-2; metatarsi I and II v2–2-2; leg III tibia
(A) (B) (C)
(D) (E)
Fig. 24. Spinoctenus stephaniae, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia.
d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, Distribution

v2–2-2, r1–1-2, p1–1-2; leg IV tibia v2–2-2, d1–1-1, r0–1- Western and Central Andean Cordilleras of Colombia
1–0, p0–1-0–1, metatarsus v2–2-2–2, r1–1-2, p1–1-2. (Fig. 36A).
Epigynum (Fig. 23D): median sector with lateral margins
curved and lateral lobes clearly delimited by heavily
sclerotised margins of middle sector, lateral lobes with small Etymology
spurs originated medially and directed medially. Vulva The specific name refers to the numerous and small spines on
(Fig. 23E): copulatory ducts divergent and spermathecae metatarsus IV in males.
rounded, with numerous anterior granules.
Variation Spinoctenus stephaniae, sp. nov.
Males (n = 2): total length 17.12–18.43; carapace 8.87–9.68; (Figs 13E–H, 24A–E, 26J, 27J, 28J, 29J, 33C, D, 36B)
femur I 8.75–9.12. Females (n = 3): total length 16.68–18.11; http://zoobank.org/urn:lsid:zoobank.org:act:CBF81465-C70B-4D23-
carapace 7.50–9.30; femur I 6.00–7.50. B333-203C8881F87D
(A) (B) (C)
(D) (E)
Fig. 25. Spinoctenus tequendama, left male palp. (A) Prolateral view; (B) ventral view; (C) retrolateral view. Female genitalia.
Material examined Other material examined. Colombia: Valle del Cauca: 1 <, El Otoño, la
Buitrera, Cali (3210 400 N, 76370 2200 W; 1800 m), S. Moreno, (MUSENUV);
Holotype. Colombia: Valle del Cauca: <, Chicoral, Bitaco, La Cumbre
1 <, Finca Alejandría, Cali (3300 N, 76340 W; 1850 m), x.2014, S. Moreno
(3340 2900 N, 76340 5300 W; 1900 m), i.2015, S. Sandoval and N. Hazzi,
(MUSENUV).
manual capture at night, on litter in a secondary forest (MUSENUV
27041).
Paratypes. Colombia: Valle del Cauca: 5 <, 13 ,, collected with Diagnosis
holotype (MUSENUV 27042); 2 <, Hacienda Bello Horizonte
(03380 4700 N, 76100 1100 W; 2080 m), El Cerrito, ii.2015, S. Moreno Males of Spinoctenus stephaniae can be distinguished from the
(MUSENUV 27043). remaining species of the genus by the non-rounded shape of the
(A) (B) (C)
(D) (E) (F)
(G) (H) (I )
(J) (K )
Fig. 26. Prolateral view of left male palp of Spinoctenus spp. (A) Spinoctenus yotoco; (B) S. chocoensis;
(C) S. eberhardi; (D) S. escalerete, arrow indicates pointed process of the embolus; (E) S. flammigerus; (F) S. florezi;
(G) S. nambi; (H) S. pericos; (I) S. spinosus; (J) S. stephaniae; (K) S. tequendama. Abbreviations: EM, external
margin of the embolus.
(A) (B) (C)
(D) (E) (F )
(G) (H) (I )
(J) (K)
Fig. 27. Ventral view of left male palp of Spinoctenus spp. (A) Spinoctenus yotoco; (B) S. chocoensis;
(C) S. eberhardi; (D) S. escalerete; (E) S. flammigerus; (F) S. florezi; (G) S. nambi; (H) S. pericos;
(I) S. spinosus; (J) S. stephaniae; (K) S. tequendama. Abbreviations: EM, external margin of the embolus.
(A) (B) (C)
(D) (E) (F)
(G) (H) (I )
(J) (K )
Fig. 28. Retrolateral view of left male palp of Spinoctenus species. (A) Spinoctenus yotoco; (B) S. chocoensis;
(C) S. eberhardi; (D) S. escalerete; (E) S. flammigerus; (F) S. florezi; (G) S. nambi; (H) S. pericos; (I) S. spinosus;
(J) S. stephaniae; (K) S. tequendama. Abbreviations: RSB, retrolateral sclerotised bulge.
(A) (B)
(C) (D)
(E) (F )
(G) (H)
(I )
(J)
Fig. 29. Retrolateral view of the metatarsus IV of males of Spinoctenus. (A) Spinoctenus yotoco; (B) S. chocoensis; (C) S. escalerete; (D) S. eberhardi;
(E) S. florezi; (F) S. nambi; (G) S. pericos; (H) S. spinosus; (I) S. stephaniae; (J) S. tequendama.
median apophysis (Figs 13E, H, 24B, 27J), by the robust base of without internal bulge (Figs 13E, H, 24B, 27J), external margin
the tegulum (Figs 13E, 24B, 28J) and by the large folded process curve and folded process large occupying more than half the
of the embolus occupying more than half the area of the embolus area of the embolus (Figs 13E, F, H, 24A). Median apophysis
without an internal bulge (Figs 13E, F, H, 26J, 27J). Females can subquadrangular in ventral view (Figs 13E, H, 24B, 27E). Tibia
be distinguished by the curved margins of the middle field of with small retrolateral sclerotised bulge and RTA triangular
the epigynum (Fig. 24D) and by the vulva (Fig. 24E) with large (Figs 13G, 24C, 28J).
spermathecae and thin copulatory ducts.
Female (MUSENUV 27042)
Description
Total length 11.62. Colour in life (Fig. 33C) as in genus
Male (MUSENUV 27041) description. Carapace 5.50 long, 4.25 wide; ocular area 1.07
Total length 10.50. Colour in life (Fig. 33B) as in genus long, 1.80 wide; eye diameters AME : ALE : PME : PLE,
description. Carapace 5.50 long, 4.75 wide; ocular area 0.625 0.27 : 0.17 : 1.62 : 0.32; AME–AME nearly AME diameter;
long, 1.62 wide; eye diameters AME : ALE : PME : PLE, PME separation 0.70 PME diameter; clypeal height 0.10;
0.30 : 0.20 : 0.27 : 0.32; AME–AME nearly AME diameter; sternum 1.55 long, 2.17 wide; labium 0.70 long, 0.75 wide.
PME separation 0.70 PME diameter; clypeal height 0.125; Leg measurements: I: femur 4.50, patella 2.37, tibia 4.50,
sternum 2.57 long, 2.25 wide; 0.60 long, 0.67 wide. Leg metatarsus 3.20, tarsus 1.30, total 15.87; II: 4.12, 2.15, 4.12,
measurements: I: femur 6.25, patella 2.62, tibia 6.00, 3.00, 1.37, 13.40; III: 3.87, 2.37, 3.00, 2.75, 1.50, 13.49; IV: 5.12,
metatarsus 4.87, tarsus 2.00, total 21.74; II: 5.62, 2.25, 5.37, 2.00, 5.00, 4.75, 1.50, 18.37. Leg spination: tibiae I and II v2–2-
4.75, 1.87, 19.86; III: 4.75, 1.87, 4.25, 4.00, 2.37, 17.24; IV: 7.00, 2–2-2; metatarsi I and II v2–2-2; leg III tibia d1–1-1, v2–2-2–2,
2.50, 7.37, 3.12, 7.50, 27.49. Leg spination: tibiae I and II v2–2- r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, r1–1-2, p1–1-2; leg IV
2–2-2, r1–1-0–0, p1–1-0–0, metatarsus v2–2-2, r1–1-0–0, p1–1- tibia v2–2-2, d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus v2–2-2–2,
0–0; leg III tibia d1–1-1, v2–2-2, r0–1-1–0, p0–1-1–0, metatarsus r1–1-2, p1–1-2. Epigynum (Fig. 24D): median sector wide with
d0–1-0, v2–2-2, r1–1-2, p1–1-2; leg IV d1–1-1, v2–2-2, r0–1- lateral margins curved and lateral lobes clearly delimited by
0–1, p0–1-0–1, metatarsus IV modified (Fig. 29I). Palp: embolus heavily sclerotised margins of the middle sector, lateral lobes
(A)
(B)
(C)
Fig. 30. Tarsi of male Spinoctenus spp. (A) Tarsus IV of S. chocoensis; (B) tarsus I of S. yotoco; (C) tarsus I
of S. stephaniae.
with small spurs that originate medially and are directed Spinoctenus tequendama, sp. nov.
medially. Vulva (Fig. 24E): spermathecae large and rounded, (Figs 25A–E, 26K, 27K, 28K, 29K, 36B)
without anterior granules. http://zoobank.org/urn:lsid:zoobank.org:act:EE1461DB-C0DE-413F-
961E-B6580CA9BD4B
Variation
Males (n = 5): total length 9.93–10.93; carapace 5.50–6.18; Material examined
femur I 5.62–6.25. Females (n = 11): total length 10.68– Holotype. Colombia: Cundinamarca: <, Bosque el Ermitaño, Vereda la
14.06; carapace 5.18–6.25; femur I 4.37–5.18. Rapida, San Antonio del Tequendama (04380 N, 74190 W; 2000 m),
vii.1998, S. Forero (ICN-AR-7925).
Distribution Paratypes. Colombia: Cundinamarca: 2 <, 2 ,, same data as holotype
(Ar-ICN-7926); 1 ,, same locality, vii.2014, S. Galvis and D. Mohna (ICN-
Western and Central Andean Cordilleras of Colombia (Fig. 36B).
AR 7927); 1 ,, Granja Ecologica el Porvenir, Vereda San Francisco, Soacha
Etymology (04350 N, 74160 W; 2550 m), vii.2009, C. Cantor and D. Martinez (Ar-ICN-
7928); 1 ,, same locality, iv.2013, C. Cantor (Ar-ICN-7929); 1 ,, Vereda
Patronym dedicated to the first author’s girlfriend, Stephania Anatoli, La Mesa (04410 N 74260 W; 1200–1600 m), J. Sanchez (Ar-ICN
Sandoval, because she is small and charming. 7930); 1 ,, La Vega, Vereda el Cural (05010 N, 74170 W; 1700 m), iv.2012
(A) (B)
(C) (D)
(E) (F )
Fig. 31. Habitus of Spinoctenus spp. Spinoctenus yotoco. (A) Female; (B) male. Spinocenus chocoensis. (C) Female; (D) male.
Spinoctenus eberhardi. (E) Female; (F) male.
(Ar-ICN 7932); Boyaca: 1 ,, Reserva Natural el Secreto, Garagoa Description

(05070 3900 N, 73160 6400 W, 2000 m), xii 2011, A. Chirivi (MPUJ-ENT
0031387); Santander: 1 ,, Cueva del Naranjo, Vereda Santa Rosa, La Male (ICN-AR-7925)
Aguada (06100 600 N, 73320 2700 W; 2019 m), viii.2008, D. Casallas-Pabon Total length 13.87. Coloration as in genus description.
and Y. Muñoz (Ar-ICN 7931). Carapace 7.06 long, 5.50 wide. Eye diameters AME : ALE :
Diagnosis diameter; PME separation 0.58 PME diameter; sternum 3.43
Males of Spinoctenus tequendama resemble those of S. yotoco long, 2.75 wide, labium 0.81 long, 0.69 wide. Leg measurements:
and S. florezi by the elongated projection at the base of tegulum I: femur 6.87, patella 3.06, tibia 7.94, metatarsus 6.37, tarsus
(Figs 25C, 28K), but differ from them by the unmodified 2.44; total 26.68; II: 7.18, 3.05, 7.00, 5.62, 1.94, 24.79; III: 6.25,
metatarsus IV (Fig. 29J) and by the absence of long setae in 2.56, 5.06, 4.62, 2.75; 21.22; IV: 7.50, 3.00, 6.87, 8.94, 2.87,
metatarsi and tibiae I and II. Females of Spinoctenus tequendama 29.18. Leg spination: leg I tibia v2–2-2–2-2, r1–1-0, p0–1-1–0,
differ from S. yotoco by the large and oval-shaped spermathecae metatarsus v2–2-2, r1–1-0–0, p1–1-0–0; leg II tibia v2–2-2–2-2,
(Fig. 25E). r1–1-1, p1–1-1, metatarsus v2–2-2, r1–1-0, p1–1-0; leg III tibia
(A) (B)
(C) (D)
(E) (F )
Fig. 32. Habitus of Spinoctenus spp. Spinoctenus escalerete. (A) Female; (B) male. Spinoctenus flammigerus. (C) Male;
(D) female. Spinoctenus nambi. (E) Female. S. yotoco. (F) Female protecting the egg sac.
d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, 2.50, 3.43, 3.87, 1.68, 16.48; IV: 6.25, 2.50, 5.68, 6.56, 2.56,
r1–1-2, p1–1-2; leg IV tibia v2–2-2, d1–1-1, r0–1-1–0, p0–1- 23.55. Leg spination: tibiae I and II v2–2-2–2-2; metatarsi I and II
0–1, metatarsus IV modified (Fig. 29J). Palp: tegulum with v2–2-2; leg III tibia d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2,
elongated projection at the base (Figs 22C, 28H), external metatarsus d0–1-0, v2–2-2, r1–1-2, p1–1-2; leg IV tibia v2–2-
margin of embolus curved (Figs 25B, 27K). Tibia with small 2, d1–1-1, r0–1-1–0, p0–1-0–1, metatarsus d0–1-0-, v2–2-2–2,
retrolateral sclerotised bulge and RTA with small rounded lobe r1–1-2, p1–1-2. Epigynum (Fig. 25D): median sector wide, with
(Figs 25C, 28K). a posterior depression slightly sclerotised and lateral lobes
clearly delimited by heavily sclerotised margins of middle
sector, lateral lobes with small spurs that originate medially
Female (Ar-ICN-7926–1) and are directed medially. Vulva (Fig. 25E): copulatory ducts
Total length 15.43. Coloration as in genus description. divergent and spermathecae large, occupying most of middle
Carapace 6.62 long, 5.62 wide. Eye diameters AME : ALE : vulva, with or without anterior granules.
diameter; PME separation 0.70 PME diameter; sternum 3.00 Variation
long, 2.62 wide; labium 1.00 long, 0.94 wide. Leg measurements: Males (n = 5): total length 12.31–14.00; carapace 5.50–7.06;
I: femur 5.62, patella 3.00, tibia 5.87, metatarsus 4.19, tarsus femur I 5.94–6.87. Females (n = 11): total length 14.75–17.55;
1.50, total 20.18; II: 5.62, 3.06, 5.18, 4.00, 1.62, 19.48; III: 5.00, carapace 6.37–8.12; femur I 5.69–6.12.
(A) (B)
(C) (D)
(E) (F )
Fig. 33. Habitus of the female of Spinoctenus spp. Spinoctenus spinosus. (A) Female; (B) male. Spinoctenus stephaniae.
(C) Female; (D) male., Spinoctenus pericos, (E) Female; (F) male.
Distribution with the remaining genera of Ctenidae. The results indicate

Eastern Andean Cordillera of Colombia (Fig. 51B). that the monophyly of Spinoctenus is supported by three
unambiguous synapomorphies with high metrics of support,
both in the parsimony and Bayesian analyses. The results
Etymology also show some well-supported groups among Spinoctenus
The species epithet is a toponym in apposition. species. This genus is the sister group to Phoneutria (although
with weak support) within the subfamily Cteninae. In addition,
Discussion Phoneutria is supported by a new unambiguous synapomorphy
not observed in previous studies: the presence of a longitudinal
Phylogeny median black line on the carapace, which can be easily observed
The present phylogenetic analysis was designed to test the in live specimens of all species (Martins and Bertani 2007; Hazzi
hypothesis of monophyly of Spinoctenus and its relationships 2014; Höfer et al. 2016). Other ctenid genera do not present this
(A) (B)
(C) (D)
Fig. 34. Habitat of Spinoctenus spp. (A, B) Reserva Natural Rio Ñambi, subtropical rainforest habitat of S. nambi and S. flammigerus, note the high density of
epiphytic and ground-based plants in B; (C) PNN Farrallones de Cali, Topacio, cloud forest habitat of S. yotoco and S. stephaniae; (D) Reserva Mateguadua,
Tulua, tropical dry forest habitat of S. spinosus.
black narrow line, but usually a brown or pale yellow proposed by Hoorn et al. (1995, 2010). The main uplift of the
longitudinal wide median band. Moreover, the addition of northern Andes may have influenced the current distributional
these new taxa did not affect the tree and we recovered patterns of Spinoctenus species from the early Miocene
the same results as the previous morphological phylogeny of (12 million years ago) to the Pleistocene (2 million years ago).
Ctenidae proposed by Polotow and Brescovit (2014). At this time, the Western and Central Cordilleras together
were an isolated unit and the elevation was less than present-
day, which could have facilitated the dispersal from the north-
Biogeography western Andes to the Chocó region. The subsequent uplift of
The genus Spinoctenus is endemic to the Andean and Chocó this area to its current elevation may have caused the breakdown
biogeographical regions. According to the DIVA results, of gene flow between these populations and caused speciation
the ancestral area of this genus is the Cauca province by vicariance. Contemporaneously, in the late middle Miocene
(Morrone 2014), which corresponds with the union of the (between 10 and 7 million years ago) the Eastern Cordillera
Western and Central Cordilleras, and reaches into southern started developing, and connected with the Central Cordillera in
Ecuador. Probably, there are still new Spinoctenus species the late Miocene (Hoorn et al. 1995, 2010). The connection of
yet to be discovered in this region, which has only been the Eastern Cordillera with the Central Cordillera may have
sampled in Colombia. The biogeographical analysis also caused the two events of dispersal towards the Eastern Cordillera.
shows three dispersal events from this region (one to Chocó These biogeographical patterns are also congruent with those
and two to Eastern Cordillera), which were subsequently identified by Mendoza et al. (2015) for Pristimantis (Anura),
followed by vicariance events. These events could be related which indicate this region of the Andes as its origin centre, from
to the palaeographical scenario of the northern Andes uplift which several dispersion events occurred to different regions of
(B)
(A)
(C)
Fig. 35. Mating sequence of Spinoctenus escalerete. (A, B) Groping; (C) copulation.
South America including the Chocó and the Eastern Cordillera. Therefore, it is possible that species of Spinoctenus do not
However, molecular studies should be integrated to this analysis possess venom that is dangerous to humans; however,
and might be used to test the robustness of the morphological the sample size is too small for confirmation (i.e. different
phylogeny among Spinoctenus species and whether the temporal people could react to the venom in different ways), and other
sequence of diversification is congruent with the biogeographical species of Spinoctenus may be harmful to humans.
hypothesis presented here. Most of the Spinoctenus species were found in sympatry
with least with one other species. The most extraordinary
case occurred in the Reserva Pericos where three species of
Field observations and ecology Spinoctenus (S. pericos, S. escalerete and S. chocoensis) were
Although Spinoctenus is closely related to Phoneutria, a genus found inhabiting the same parts of the forest and the same
of venomous species with potential medical significance to microhabitats. Moreover, the phylogenetic analysis indicates
humans, the first author was bitten by some species of that these three species are more closely related than the
Spinoctenus (a female S. yotoco, a female S. escalerete and remaining species. As expected in evolution and ecology
a male S. flammigerus) during this study, and only presented theory, the three species should not co-occur due to
symptoms of pain and swelling in the area of the bite (fingers). competition (Krebs 2008). In the Amazonia region, a similar
(a) (b)
78°0′0″W 77°0′0″W 76°0′0″W 75°0′0″W 74°0′0″W 73°0′0″W 78°0′0″W 77°0′0″W 76°0′0″W 75°0′0″W 74°0′0″W 73°0′0″W
9°0′0″N 9°0′0″N
8°0′0″N 8°0′0″N
7°0′0″N 7°0′0″N
N N
6°0′0″N 6°0′0″N
W E W E
S S
5°0′0″N 5°0′0″N
4°0′0″N 4°0′0″N
3°0′0″N 3°0′0″N
2°0′0″N 2°0′0″N Spincotenus pericos

Spinoctenus chocoensis
Spinoctenus eberhardi Spinoctenus escalerete
Spinoctenus florezi Spinoctenus spinosus

1°0′0″N 1°0′0″N
Spinoctenus nambi Spinoctenus stephaniae
Spinoctenus yotoco Spinoctenus tequendama
0°0′0″ 0 25 50 100 150 200 0°0′0″ 0 25 50 100 150 200

Kilometers Kilometers
Fig. 36. (A, B) Geographical distribution of Spinoctenus spp.
case occurs with three sympatric species of Phoneutria Gasnier et al. 2002; Rego et al. 2005, 2007; Torres-Sanchez
(P. boliviensis, P. fera and P. reidyi); however, ecological and Gasnier 2010). The discovery of this new genus containing
studies have demonstrated that their niches differ in the a large number of species in the Andean and Chocó regions
same habitat. According to Torres-Sanchez and Gasnier highlights the immensity of the deficit in our knowledge of
(2010) and Höfer et al. (2016), P. boliviensis seems to be Colombian biodiversity.
restricted to inundation forests while the two larger species
(P. fera and P. reidyi) appear to be abundant in forests Conflicts of interest
(‘terra firme’). In the case of these two species, P. fera prefers
the ground surface of the forest (leaf litter) as microhabitat and The authors have declared that no competing interests exist.
P. reidyi shows preference for lower vegetation, especially
acaulescent palms. In addition, these species present different Acknowledgements
reproductive cycles (Torres-Sanchez and Gasnier 2010). Our We are grateful to Raul Rios (IMCN), Carmen E. Posso (MUSENUV),
preliminary observations indicate no differences between the Dimitri Forero (MPUJ) and Eduardo Flórez (ICN) for making available
microhabitats of S. pericos, S. escalerete and S. chocoensis, the spiders for this study. We are deeply indebted to Juan F. Ortega and the
as all the specimens were collected in the same parts of the Laboratorio de Imágenes, Departamento de Biologiía, Universidad del Valle
forest, in the same microhabitat (i.e. leaf litter on the ground) for their assistance with the stereoscopic images. The authors thank Abel
and both males and females at the same time of year. Further, we Pérez, Diana Silva and Jimmy Cabra for their valuable comments throughout
report several cases of cannibalism and possible intra-guild the development of the project. N. Hazzi thanks Stephania Sandoval, Ruben
predation between these spiders, suggesting that competition D. Palacio, Sebastian Moreno and Jairo Moreno for their valuable company
in the field. We also thank Ruben D. Palacio, Jorge Cañas and Sebastian
may be strong. Future studies are necessary to furnish the basis
Moreno for taking the photos of Figs 32C, 33F and 2, respectively and Juan
for understanding this coexistence. David Escobar for the drawings of Fig. 35. We also are grateful to S. Villar,
Spinoctenus may be keystone species in the forests where they M. Rodao and A. Laborda for their assistance with the use of the scanning
occur due to their habitat preferences. Therefore, these spiders microscope. We thank V. Townsend for help with the English version of
may be good indicators of disturbance or forest fragmentation, as the manuscript. M. Simó acknowledges financial support from Programa de
occurs with other species of ctenids (Gasnier and Höfer 2001; Desarrollo de las Ciencias Básicas (PEDECIBA) and Sistema Nacional de
Investigadores (SNI) de la Agencia Nacional de Investigación e Innovación Flórez, E., Ortiz, A., and Montoya, M. (2003). Accidentes por mordedura de
(ANII). The manuscript was greatly improved by critical comments from la araña de las bananeras Phoneutria boliviensis (Araneae: Ctenidae) en
Robert Raven and an anonymous reviewer. la región de Urabá, Colombia. Entomólogo 96, 1–4.
Foelix, R. (2010). ‘Biology of the Spiders’. 3rd edn. (Oxford University
Press: New York, NY.)
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Systematics and biogeography of Spinoctenus, a new genus of wandering

Article in Invertebrate Systematics · February 2018

Nicolas Hazzi Daniele Polotow

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Antonio Domingos Brescovit Ranulfo González-Obando

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Systematics and biogeography of Spinoctenus, a new genus

Nicolas A. Hazzi A,B,C,H, Daniele Polotow D,E, Antonio D. BrescovitF,

Journal compilation CSIRO 2018 www.publish.csiro.au/journals/is

Table 1. Character descriptions of Spinoctenus species

Table 2. Summary of the phylogenetic hypotheses among the most Diagnosis

78°0′0″W 76°0′0″W 74°0′0″W 72°0′0″W

9°0′0″N (WCC) Spinoctenus eberhardi

(WCC) Spinoctenus stephaniae

8°0′0″N (WCC) Spinoctenus nambi

(WCC) Spinoctenus flammigerus

7°0′0″N (EC) Spinoctenus tequendama

6°0′0″N (EC) Spinoctenus florezi

2 and 3 (WCC) Spinoctenus spinosus

5°0′0″N (C) Spinoctenus pericos

(C) Spinoctenus chocoensis

4°0′0″N (C) Spinoctenus escalerete

Cis Andean (CI)

Eastern Cordillera (EC)

Etymology as Cupiennius granadensis Keyserling, 1877, Ancylometes

Variation Material examined

(A) (B) (C)

(A) (B) (C)

(A) (B) (C)

(A) (B) (C)

(A) (B) (C)

(A) (B) (C)

Spinoctenus pericos, sp. nov. Material examined

(A) (B) (C)

(A) (B) (C)

(A) (B) (C)

d1–1-1, v2–2-2–2, r0–2-0–2, p0–2-0–2, metatarsus v2–2-2, Distribution

Variation Spinoctenus stephaniae, sp. nov.

(A) (B) (C)

(A) (B) (C)

(D) (E) (F)

(A) (B) (C)

(A) (B) (C)

(D) (E) (F)

(Ar-ICN 7932); Boyaca: 1 ,, Reserva Natural el Secreto, Garagoa Description

Distribution with the remaining genera of Ctenidae. The results indicate

2°0′0″N 2°0′0″N Spincotenus pericos

Spinoctenus florezi Spinoctenus spinosus

Spinoctenus yotoco Spinoctenus tequendama

0°0′0″ 0 25 50 100 150 200 0°0′0″ 0 25 50 100 150 200

Fig. 36. (A, B) Geographical distribution of Spinoctenus spp.

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