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Finding hot singles: matching males to females in dimorphic
spiders (Araneidae : Micrathena) using phylogenetic
placement and DNA barcoding
Ivan L. F. Magalhaes
A,B,D
, Pedro H. Martins
B
, André A. Nogueira
C
and Adalberto J. Santos
B
A
División Aracnología, Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’–CONICET,
Av. Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina.
B
Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais,
Av. Antônio Carlos 6627, CEP 31270-901, Belo Horizonte, Minas Gerais, Brazil.
C
Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão,
Travessa 14, n. 321, CEP 05508-900, São Paulo, São Paulo, Brazil.
D
Corresponding author. Email: magalhaes@macn.gov.ar
Abstract. Many orb-weaving spiders exhibit remarkable sexual dimorphism, hampering the matching of males and
females in taxonomic studies. This is the case for the spiny Micrathena spiders, a species-rich Neotropical genus with 27% of
its species known from a single sex. In this paper we document several undescribed Micrathena specimens, and test whether
they belong to some of those incompletely known species. In order to do so, we: (1) tested the phylogenetic position of males
and their putative females using a previous morphological dataset; (2) calculated genetic distances among individuals based
on a fragment of the mitochondrial gene cytochrome coxidase subunit I; and (3) examined their geographical distributions.
These approaches, isolated or in combination, allowed us to identify and describe the previously unknown males of M. embira
Levi, M. reimoseri Mello-Leitão, M. exlinae Levi, M. miles Simon, M. spinulata F.O. Pickard-Cambridge, M. yanomami
Magalhães & Santos and M. cornuta (Taczanowski), and the female of M. beta di Caporiacco. We found that the males
previously associated with M. bicolor (Keyserling), M. cornuta and M. lata Chickering had been incorrectly matched with
females. The latter actually belongs to a hitherto unnamed species, herein described as Micrathena perfida, sp. nov. New
geographical data are given for these and other Micrathena species. Our study highlights the importance of using different
sources of data for matching the sexes in diverse groups with strong sexual dimorphism.
http://zoobank.org/urn:lsid:zoobank.org:urn:lsid:zoobank.org:pub:AFEDA522-D0B4-4902-A747-9721DEED8B89
Additional keywords: Araneae, cladistics, taxonomy.
Received 18 December 2015, accepted 17 June 2016, published online 16 March 2017
Introduction
Orb-weaving spiders are known for their remarkable sexual
dimorphism. The Araneidae family contains classical examples
of extreme sexual size dimorphism, such as Nephila Leach,
1815 and Argiope Audouin, 1826, in which females can be
more than 10 times larger than the males (Vollrath 1998).
Sexual dimorphism may also include significant differences in
body morphology and colouration patterns (Cheng and Kuntner
2015). Matching males to females, especially in diverse groups,
is a challenging task for taxonomists (Edwards 2013), and the
situation is particularly difficult in groups with pronounced
sexual dimorphism (e.g. Przybyłowicz and Tarcz 2015).
A good example of this situation is the araneid genus Micrathena
Sundevall, 1833, which contains 116 species (World Spider
Catalog 2016) occurring in forested habitats from the southern
USA to northern Argentina. While females are a conspicuous
element of Neotropical spider communities, with their spiny
and colourful abdomens, males are usually much smaller and
lightly built, with darker and unarmed abdomens (Figs 1–5). This
makes males much harder to collect and, consequently, rarer in
collections than females. For instance, in two recent inventories
in the Brazilian Amazon, 134 and 1040 adult Micrathena
individuals were collected, but of those only 18% and 12.8%
were males (AAN, unpubl. data). Also, as of 2016, the spider
collection at Centro de Cole¸cões Taxonômicas, Universidade
Federal de Minas Gerais, Belo Horizonte, Brazil, housed
113 Micrathena males, as opposed to 579 females. This male–
female disparity in specimen availability leads to two main
Journal compilation CSIRO 2017 www.publish.csiro.au/journals/is
CSIRO PUBLISHING
Invertebrate Systematics, 2017, 31,8–36
http://dx.doi.org/10.1071/IS15062
(A)
(C)
(E) (F)
(D)
(B)
Fig. 1. Living Micrathena specimens. (A)Micrathena miles, female with egg-sac from Maynas, Loreto, Peru. (B)Micrathena miles, male from
Maynas, Loreto, Peru (MUSM-ENT 510057). Approximately to scale regarding figure part A.(C)Micrathena yanomami, female from Tambopata,
Madre de Dios, Peru (UFMG 19567). (D)Micrathena yanomami, female from Paucartambo, Cusco, Peru (MUSM-ENT 510056). (E)Micrathena
embira, female from Porto Velho, Rondônia, Brazil. (F)Micrathena embira, male from the same locality. Photos A–Dby A. Anker and P. H. Martins,
E,Fby F. U. Yamamoto.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 9
consequences: (1) several species are known only from females
(31 out of 116; only Micrathena beta di Caporiacco, 1947 is
known only from the male); and (2) males are usually described
subsequently, and their matching to existing females is often
based on poor evidence.
The particularly difficult task of defining male–female
associations in Micrathena has already been highlighted by
Levi (1985) in his extensive revision of the genus. Prior to
Levi’s study, more than 77% of Micrathena species were
known only from females, a proportion that decreased to
32% after his work. He also commented that matching males
to females was the ‘biggest problem of the genus’due to marked
sexual dimorphism and high diversity. He overcame this matter
by dividing the species (both males and females) into species-
groups and associating them based on overlapping distributions
(Levi 1985: 433). However, this geographic criterion can be
difficult to apply as many species may occur in sympatry. For
instance, an inventory in Porto Velho, in the Brazilian Amazon,
revealed that as many as 26 species of Micrathena can be found
in the same area (AAN, unpubl. data). Levi (1985) was aware of
this problem, and admitted that the association between males
and females was tentative in several cases.
Newly collected specimens are very important to complement
the descriptions of species known from a single sex, and also to
confirm the association of species known from both sexes.
On recent field expeditions, we collected specimens of some
poorly known Micrathena species: M. beta,M. embira Levi,
1985, M. reimoseri Mello-Leitão, 1935, M. exlinae Levi, 1985,
M. miles Simon, 1895, M. spinulata F.O. Pickard-Cambridge,
1904 and M. yanomami Magalhães & Santos, 2011. All of these
species have so far been known from a single sex, and most from
only a few records (see Levi 1985; Magalhães and Santos 2011).
In addition, several previously undescribed specimens, including
males and females, were collected in these same areas (Table 1).
It seemed clear that at least some of the undescribed specimens
probably belonged to these species known from a single sex;
however, matching them was not straightforward. For instance,
our undescribed male #1 was collected in the same area as
M. miles and M. embira, lacks a terminal apophysis, as would
be expected for the male of M. embira, but has a hook on the
coxa I, as expected for the male of M. miles (see Levi 1985).
Additionally, in the recent inventory in Porto Velho several
specimens of M. bicolor (Keyserling, 1864) and M. cornuta
(Taczanowski, 1873) were collected. Although the two species
are known from both sexes, the field sampling consistently
yielded only males of the first species, and only females of
the latter. Could the matching proposed by Levi (1985) for
these species be wrong?
We considered that analysing males and females in a
phylogenetic context would offer more robust evidence of
association between the sexes than considering solely the
geographic distribution and some morphological characteristics.
Micrathena species can be sorted into species-groups, whose
delimitation is reasonably clear for both sexes (Magalhães and
Santos 2012). Although the use of morphological data to match
dimorphic sexes may seem contradictory, we consider that the
placement of both sexes in the same species-group along with
their co-occurrence at collecting sites may represent strong
evidence of conspecificity (see Edwards 2013; Barone et al.
2016). Another possibility is the use of molecular data, in which
DNA sequences from both sexes can be compared (as opposed to
morphological features, which more often than not are present
in only one of the sexes). Molecular data have proven to be
very useful in resolving taxonomic questions for groups in
which the analysis of morphology alone is inconclusive.
Examples include separating cryptic species in groups with low
morphological variation (Clouse and Wheeler 2014), clarifying
ecological interactions (Hulcr et al.2007), associating different
developmental stages of taxa with life cycles that exhibit
remarkably different forms (e.g. cnidarians; Miranda et al.
2010) and, of course, associating males to females in sexually
dimorphic species (Tanikawa 2011; Przybyłowicz and Tarcz
2015; Barone et al. 2016).
In this paper, we test male–female conspecificity hypotheses
for M. beta,M. embira,M. reimoseri,M. exlinae,M. miles,
M. spinulata and M. yanomami (all known from a single sex), and
M. cornuta,M. lata and M. bicolor (known from both sexes, but
with a dubious association between males and females).
Specifically, we wanted to test whether some of the recently
collected and yet undescribed specimens belong to the species
known from a single sex, and whether the current male–female
matching in M. cornuta,M. lata and M. bicolor could be wrong.
We followed three approaches to verify our hypotheses:
(1) determining the phylogenetic placement of each of the
seven species and each of the undescribed specimens by
including them in the morphological matrix of Magalhães
and Santos (2012); (2) calculating genetic distances between
males and females based on a fragment of the mitochondrial
gene coding for the cytochrome coxidase subunit I (COI); and
(3) checking geographic distributions, with data from recent
field surveys. After matching males to their females, we
describe and illustrate them for the first time, and provide new
records for these species.
We dedicate this paper to the recently deceased Herbert
W. Levi, who devoted most of his life to studying the
systematics of orb-weaving spiders, such as Micrathena.
Although we herein suggest some modifications to
combinations he proposed, it is needless to say that he got
most of the species right, and that without his immense
contribution, studies like our own would not be possible. We
hope and believe that his fine work will continue to inspire
generations of spider taxonomists.
Materials and methods
Phylogenetic inference based on morphology
To infer the phylogenetic position of the undescribed specimens
and their candidate males or females, we included them in
the discrete character matrix of Magalhães and Santos (2012).
At first, we scored the undescribed specimens and their
putative pairs (M. beta,M. embira,M. reimoseri,M. exlinae,
M. miles,M. spinulata,M. yanomami,M. cornuta,M. lata,
M. bicolor) as separate terminal taxa. Thus, at this phase each
‘species’was represented by two terminal taxa, e.g. ‘M. embira
female’(with all male characters coded as missing data) and
‘Undescribed male #1’(with all female characters coded as
missing data). This approach is similar to that employed by
Platnick and Shadab (1978) and Barone et al. (2016), and the
10 Invertebrate Systematics I. L. F. Magalhaes et al.
(A) (B)
(C) (D)
(E) (F)
Fig. 2. Living Micrathena specimens. (A)Micrathena beta, female from Requena, Loreto, Peru (MUSM-ENT 510055). (B)Micrathena
cornuta, female from Requena, Loreto, Peru (UFMG 19564). (C)Micrathena reimoseri, female from Miguel Calmon, Bahia, Brazil.
(D)Micrathena shealsi, female from Paucartambo, Cusco, Peru (MUSM-ENT 510061). (E,F)Micrathena perfida, sp. nov., holotype
male and paratype female from Duque de Caxias, Rio de Janeiro, Brazil (UFMG 19049, 19048). All photos by A. Anker and P. H. Martins,
except for Cby G. F. B. Pereira.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 11
latter authors call this a ‘split matrix’.Micrathena cucharas
(Levi, 1985) and M. woytkowskii (Levi, 1985) have also been
included in this analysis as both seem closely related to
M. cornuta and the males and females of the three species
could potentially be mixed up; we have found some specimens
of them misidentified in collections, including a male of
M. cornuta collected with a female M. woytkowskii in a MCZ
vial. We took this approach of including sexes separately so we
could check whether each would be found to belong in the same
Micrathena species-groups. After this initial analysis and
considering genetic and geographic evidence, males were
matched to females. We then re-ran analyses joining males
and females of each species into a single terminal taxon (this
time coded for all characters) to infer their phylogenetic
placement with fewer missing data (a ‘merged matrix’; Barone
et al. 2016). We also took this opportunity to test the monophyly
of the kirbyi species-group as defined by Magalhães and Santos
(2012) by including Micrathena macfarlanei Chickering, 1961
in the phylogeny. Laboratory procedures, coding schemes
and characters are described in detail by Magalhães and Santos
(2012). We searched for most-parsimonious trees under implied
weight (Goloboff 1993) using TNT 1.1 (Goloboff et al.2008),
with heuristic searches starting with 10 000 random addition
sequences, retaining 100 trees per replication, followed by
tree-bisection-and-reconnection branch-swapping (TBR).
Analyses were run with a kvalue of 5 (the preferred value
over a wide range tested in Magalhães and Santos 2012).
Results of analyses under equal weights and other values of k
are not included here because preliminary runs yielded very
similar results to the ones we present, and because the
topologies and conclusions we take from them are robust to
different search parameters (see also Magalhães and Santos
2012). We evaluated branch support using Bremer’s decay
index (estimated using the bremer.run script, which is part of
the TNT package) and symmetric resampling (Goloboff et al.
2003). Symmetric resampling calculations were based on 100
pseudoreplicates with 500 random addition sequences, saving
100 trees per replication, followed by TBR.
Due to the inclusion of new taxa, we made some additions
to the character list developed by Magalhães and Santos (2012).
We added a fifth state to character 126 (palpal bulb, median
apophysis, basal projection, shape): (4) pointed towards embolus
tip. We also propose four new characters:
Ch. 137. Male carapace, pars cephalica, colouration: (0) same
colour as the rest of the carapace; (1) with darker colouration
around median eyes (scored 1 only for M. kirbyi (Perty, 1833),
M. macfarlanei and M. miles).
Ch. 138. Palpal bulb, conductor, base shape: (0) small, not
extending beyond tegulum; (1) enlarged, extending beyond
tegulum.
Ch. 139. Carapace, tubercles, shape: (0) a swelling with
projected setal bases; (1) spiniform. Only applicable to species
with carapace tubercles.
Ch. 140. Abdomen, second pair of posterior spines, position in
relation to first pair: (0) posterior/ventral; (1) anteriorly displaced.
Genetic distances
We used specimens collected by us or our colleagues in the field
and stored in 96% ethanol at 20C, or museum samples
collected from no earlier than 2010 and stored in 75% ethanol
at room temperature. We extracted DNA from muscle tissue of
1–4 legs of each individual using a Wizard Genomic DNA
purification kit (Promega, Madison, WI, USA) following the
manufacturer’s instructions. We amplified a ~1200-bp fragment
of the mitochondrial gene coding for COI using the primers
LCOI1490 (50-GGT-CAA-CAA-ATC-ATA-AAG-ATA-TTG-
G-30) (Folmer et al.1994) and C1-N-2776 ‘SPID’(50-GGA
TAA TCA GAA TAT CGT CGA GG-30) (Hedin and
Maddison 2001). For specimens that failed to amplify for this
fragment, we used the alternative pair C1-N-2568 (50-GCT ACA
ACA TAA TAA GTA TCA TG-30) and C1-J-1751 ‘SPID’(50-
GAG CTC CTG ATA TAG CTT TTC C-30) (Hedin 1997), which
amplifies a ~720-bp fragment. We amplified the fragments using
the following PCR conditions: 5 mLof5buffer (Promega Gotaq
Flexi buffer), 2 mL of 25 mM MgCl
2
, 2.5 mL of dNTP mix at
2mM (Promega), 2.5 mL of each primer at 5 mM, 0.25 unit of
Taq DNA polymerase (Promega Gotaq) and 1 mL of extracted
DNA for a final volume of 25 mL (or 10 mL with proportional
quantities). PCR cycles consisted of 10 min at 94C, 35–40
(30 s at 95C, 45 s at 48C (primer pair 2568–1751) or 52C
(1490–2776), and 45 s to 1 min at 72C) and 7–10 min at 72C. An
alternative PCR program (including five initial cycles at 46C)
was used when the first PCRs failed to succeed for museum
samples. We checked PCR success on a 1% agarose gel stained
with GelRed (Biotium, Fremont, CA, USA) and purified the
products using a 20% 8000 polyethyleneglycol + NaCl solution
followed by washes in 80% ethanol. Sequencing reactions
consisted of 4 mL of PCR product, 1 mL of either forward or
reverse primers, 1.6 mLof5sequencing buffer and 0.8 mLof
BigDye Terminator Cycle Sequencing Kit (Applied Biosystems,
Foster, CA, USA) for a 10 mLfinal volume. Automated
sequencing was carried out on an ABI 3130xl Genetic
Analyzer (Applied Biosystems) following the manufacturer’s
instructions. We checked chromatograms by eye in SeqScape
2.6.0 (Applied Biosystems). We aligned sequences using Muscle
(Edgar 2004) as implemented in MEGA6 (Tamura et al.2013).
Additional Micrathena sequences from McHugh et al.(2014)
were obtained from GenBank; newly generated sequences
are deposited in the same database under accession numbers
KX687305–KX687332. We used MEGA6 for inferring a
neighbour-joining tree for the sequence set using a Kimura 2-
parameter model of evolution. Branch support was evaluated
using 100 bootstrap pseudoreplicates.
Specimen descriptions
The format of descriptions follows Magalhães and Santos (2011).
Specimens were illustrated with the aid of a Leica M205C
stereoscopic microscope with a camera lucida. Photographs
were taken using a DFC295 digital camera attached to the
same microscope and mounted as a single, multifocal image
using Leica Application Suite 3.8. Measurements were taken on
the left side of specimens, except when otherwise specified, and
are expressed in millimetres.
Most new records were georeferenced in situ by the original
collectors. Previously known and new records without
coordinates were georeferenced post hoc using Google Earth
and are approximate (and indicated between square brackets,
rather than parentheses).
12 Invertebrate Systematics I. L. F. Magalhaes et al.
Fig. 3. Phylogeny obtained under implied weights analysis (k= 5) of morphological data, with newly described males and females codedas separate
taxa (‘split matrix’). Groups relevant to the pairings made here highlighted in grey, and codes for undescribed specimens followTable 1; species are
labelled with names that have been applied to them before the conclusions of our study. Values next to branches are group frequencies of symmetric
resampling (only values higher than 50 are shown).
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 13
The material examined for this study is deposited in the
following museum collections: CAS, California Academy of
Sciences, San Francisco, USA; CHNUFPI, Cole¸cão de
História Natural da Universidade Federal do Piauí, Floriano,
Brazil; IBSP, Instituto Butantan, São Paulo, Brazil; INPA,
Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil;
MACN, Museo Argentino de Ciencias Naturales Bernardino
Rivadavia, Buenos Aires, Argentina; MCZ, Museum of
Comparative Zoology, Cambridge, USA; MNRJ, Museu
Nacional do Rio de Janeiro, Rio de Janeiro, Brazil; MPEG,
Museu Paraense Emílio Goeldi, Belém, Brazil; MUSM-ENT,
Museo de Historia Natural de la Universidad Nacional Mayor
de San Marcos, Lima, Peru; MZSP, Museu de Zoologia
da Universidade de São Paulo, São Paulo, Brazil; and UFMG,
Centro de Cole¸cões Taxonômicas, Universidade Federal
de Minas Gerais, Belo Horizonte, Brazil. Specimens used for
DNA extraction are listed together with collection and depository
information and GenBank accession numbers in Supplemenary
Table S1.
Results and discussion
Morphological phylogenetic inference with sexes coded
separately (split matrix)
The analysis with sexes coded separately yielded two fittest trees
differing only in the position of M. exlinae females, which were
either sister to M. plana (C.L. Koch, 1836) or M. excavata (C.
L. Koch, 1836), length = 599, fit = 95.64, CI = 0.275, RI = 0.714
(Fig. 3). Despite the addition of new taxa, the morphological
topologies obtained in this analysis are very similar to those
found by Magalhães and Santos (2012) (Fig. 3). The females of
M. embira,M. reimoseri,M. exlinae,M. miles,M. spinulata and
M. yanomami were recovered as members of the schreibersi,
militaris,plana,guerini,gracilis and triangularispinosa
species-groups, respectively, as anticipated by Levi (1985) and
Magalhães and Santos (2012). The male of M. beta was recovered
as sister to the male currently assigned to M. lata, both in the
militaris-group (Fig. 3).
The undescribed specimens had contrasting behaviours. Most
appeared in the same morphological groups as the females they
had been collected with: this is true for undescribed males #1, 2, 3,
5 and 6 (Table 1; Fig. 3). Other males appeared in unexpected
positions. For example, male #4 was collected with M. miles
females (guerini-group), but was recovered as a member of the
kirbyi-group (Fig. 3). Male #7 and M. bicolor males were
collected in the same locality as M. yanomami, and all were
recovered in the triangularispinosa-group. However, male #7
was recovered in a relatively apical position within this group,
while M. yanomami females and M. bicolor males were recovered
in a politomy at the base of the group (Fig. 3). Finally, the two
undescribed females were recovered in the triangularispinosa-
group; one of them had been collected along with males of M. beta
(militaris-group) (Table 1; Fig. 3).
Genetic distances
The neighbour-joining dendrogram based on a COI fragment
shows that undescribed male #1, collected in the same locality as
M. miles and M. embira females, is genetically more similar to the
latter (Fig. 4). Similarly, male #2 grouped with M. reimoseri, male
#5 grouped with females of M. cornuta, and male #6 grouped with
M. spinulata (Fig. 4)–each of these pairs coming respectively
from the same localities. Undescribed male #7 is most similar
to females identified as M. triangularispinosa (De Geer, 1778)
(Fig. 4). Surprisingly, the females of M. yanomami, which we
initially believed to be conspecific with this latter male, grouped
with males of M. bicolor (Fig. 4). Undescribed female #1 and a
M. beta male have almost identical COI sequences, as do
undescribed female #2 and the male currently assigned to
M. lata; these two pairs form a cluster, although with more
than 4% COI sequence divergence between them (Fig. 4).
Matching sexes using different sources of data
Associating males to females in spider groups with several
sympatric species is always challenging. Levi (1985) stated
that the chances of successfully matching couples are higher in
the context of a revision, but that does not prevent association
mistakes. Fortunately, most of Levi’s pairings have proved to be
correct over the years. Nowadays we are lucky to obtain data
that were not available to him, such as additional specimens
deposited in museums, data from recent field expeditions, a robust
phylogeny of the genus and DNA sequences. With these, we can
review some of the old pairings and propose new ones.
The utility of DNA sequences for matching sexes (or different
developmental stages) in spiders had already been anticipated by
Barrett and Hebert (2005). It is now routinely used in taxonomic
papers (e.g. Faynel et al.2012; Trivinho-Strixino et al.2012;
Przybyłowicz and Tarcz 2015; Barone et al. 2016). Our study
shows that DNA barcoding can be successfully used for matching
sexes in organisms with marked sexual dimorphism. Even in
cases where a single male and a single female of a particular
species could be sequenced (such as M. reimoseri or M. embira;
Fig. 4), the information was useful and concordant with other
sources of data. This is in part because, at least in Micrathena,
divergences in COI correlate well with morphology-based
identification (McHugh et al.2014; this study, Fig. 4). On the
other hand, coding different life stages (or sexes) as separate
terminal taxa (= semaphoront coding) in a phylogenetic analysis
is controversial, although it may be useful in some situations
(e.g. Steyer 2000; Gilbert et al.2009; Meier and Lim 2009; Wolfe
and Hegna 2014). At least two studies have successfully used
this approach to associate the males with females in anyphaenids
(Barone et al.2016) and anapids (Platnick and Shadab 1978),
although in the latter case the authors noted some conflicts
between male and female characters. Sharma et al.(in press)
argue against semaphoront coding. We do agree with them that it
violates several assumptions of cladistic analysis and should not
be used to infer phylogenies in the strict sense. On the other hand,
it might be useful in situations such as the present one, to infer
if different stages present synapomorphies of a particular group,
which might represent evidence of conspecificity. Needless
to say, after matching the sexes it is fundamental to abandon
semaphoront coding and re-analyse the data, and that is why we
present the tree in Fig. 5. Here, we demonstrate that semaphoront
coding, in combination with DNA sequences and geographic
information, allowed us to either confirm existing or propose
new matches for seven species of Micrathena. Details of each
particular case are given in the ‘Taxonomy’section.
14 Invertebrate Systematics I. L. F. Magalhaes et al.
Fig. 4. Neighbour-joining dendrogram based on Kimura 2-parameter distances. Numbers next to nodes represent bootstrap supports (based on 1000
pseudoreplications). Codes for undescribed specimens follow Table 1; species are labelled with names that have been applied to them before the
conclusions of our study.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 15
It is clear that male–female matching requires a good deal of
logical reasoning and ‘detective work’. This is especially true
in cases like Micrathena, in which data are incomplete or
ambiguous for some of the species. For example, male #1 was
collected with M. embira females, and both DNA sequences and
the morphological phylogeny indicate that it belongs to this
species (Table 1). The decision was unequivocal in this case.
However, male #4 was collected with M. miles, but we could not
obtain sequences from the male, and males and females were
placed apart in the morphological phylogeny (Fig. 3). In this case,
we had to rely solely on geography to make a decision. In the
cases of the cornuta and triangularispinosa species-groups,
species are morphologically homogeneous and morphological
phylogenetic placement was not of great help; having sequence
data was crucial to identify the correct males of M. cornuta and
M. yanomami. A similar case occurred with M. miles and M. beta,
in which females and males were not placed in the same groups
in the morphological analysis. This suggests that sometimes
using only female characters for estimating the position of a
species may be misleading, and that sequence data should be
used whenever possible. Finally, it is interesting that having data
on closely related species can help decision-making for a poorly
known taxon. This is the case of M. beta and M. perfida, sp. nov.:
recognising the female of M. beta allowed us to notice that
the male considered to belong with M. lata by Levi (1985)
was actually wrongly matched, and that it belongs to a so
far undescribed species (see details in the ‘Taxonomy’section
below).
Based on the results and arguments presented in the
‘Taxonomy’section, we were able to match six of the seven
undescribed males to the females of M. embira,M. reimoseri,
M. exlinae,M. miles,M. spinulata and M. cornuta, respectively.
The male previously assigned to M. cornuta belongs to
M. woytkowskii instead. Unidentified male #7, which we
initially suspected to be conspecificwithM. yanomami females,
was placed within a complex formed by specimens identified
by us as M. triangularispinosa. On the other hand, females of
M. yanomami were associated with males previously matched by
Levi (1985) to the females of another species, M. bicolor.Oneof
the undescribed females is matched to M. beta. Finally, the male
previously described as M. lata does not belong with this species
and is here described as Micrathena perfida, along with its
true female. For a detailed justification of the matching of each
species, diagnoses, descriptions and new geographical records,
see the ‘Taxonomy’section.
Phylogenetic analysis with both sexes coded (merged matrix)
The phylogenetic analysis with both male and female characters
coded for every species known from both sexes had similar results
(Fig. 5) (length = 605, fit = 83.16, CI = 0.273, RI = 0.710) to
the previous analysis (Fig. 3). In the cases in which males and
females of the same species were placed in different groups
in the previous analysis, the species retained the position
previously found for the male (M. miles in the schreibersi-
group; M. lata and M. perfida in the militaris-group) (Fig. 5).
In this dataset, there are fewer male characters than female
characters (see Magalhães and Santos 2012), but they seem to
be especially important to correctly estimate the phylogenetic
position of Micrathena species. This highlights the importance of
describing unknown males. One of the unexpected results of
our analyses is the position of M. beta and M. perfida, which share
male characters with the militaris-group, but whose females
definitely resemble those of the triangularispinosa-group. This
means that the limits and relationships of this group might change
upon the addition of more data.
Pending taxonomic issues in Micrathena
Micrathena is arguably one of the best-known Neotropical
spider genera with regard to its systematics: it has been
thoroughly reviewed, a phylogeny is available and few
taxonomic novelties have appeared over the years (reviewed in
Magalhães and Santos 2011; Argañaraz and Rubio 2011; this
study). Nevertheless, some problems remain to be solved.
The most obvious one, which we partially tackled with
this contribution, is the lack of knowledge on the males of
many species. Another major issue is the taxonomy of the
triangularispinosa-group, which currently includes 11 small
species. Some species are difficult to distinguish from others,
such as M. annulata Reimoser, 1917 and M. jundiai Levi, 1985.
On the other hand, many of the nominal species, as currently
recognised, have a variable morphology and may represent
species complexes; examples include M. evansi Chickering,
1960 and M. flaveola (Perty, 1839). To address this, we
believe it is necessary to carry out a comprehensive revision of
this species-group based on exhaustive examination of museum
specimens (including types of junior synonyms). As these
issues are gradually solved, an even more solid base will be
erected for the taxonomy of Micrathena –an excellent outcome,
as these spiders are as much conspicuous and charismatic as
they are excellent models for studies in ecology, evolution and
biogeography.
Taxonomy
Family ARANEIDAE Clerck
Genus Micrathena Sundevall
Micrathena perfida, sp. nov.
(Figs 2E,F,6,7A–C,15)
http://zoobank.org/urn:lsid:zoobank.org:act:3A9E9EB7-3534-4A07-
BBBD-1535AF51C5FA
Micrathena lata Chickering: Levi, 1985: 565, figs 608–615 (in part,
misidentified male); Magalhães & Santos, 2012: 51 (in part,
misidentified male).
Material examined
Holotype. Brazil: Rio de Janeiro: <, Duque de Caxias, Parque Natural
Municipal da Taquara (2235035.300S, 4314017.800 W, 36 m), 6–7.i.2016,
A. Anker, P. H. Martins and R. Brito (UFMG 19409).
Paratypes. Same data as the holotype, 1 ,(UFMG 19408). Brazil:
Espírito Santo: Santa Teresa [19550S, 40360W], 29.ix.1942, B. A. Soares,
1<(MZSP7982); Linhares, Reserva NaturalVale [198015.600 S, 40303600W],
5–12.i.1998, A. D. Brescovit et al.,1<(IBSP 16749). São Paulo: Iguapé,
Serra de Itimirim, divisa com Miracatu [242203000S, 47310800 W],
20.xii.1998, R. Pinto-da-Rocha and S. Bérnils, 1 <(MZUSP 17212);
16 Invertebrate Systematics I. L. F. Magalhaes et al.
Fig. 5. Phylogeny obtained under implied weights analysis (k= 5) of morphological data with all characters coded for each species (‘merged matrix’).
Values below branches represent Bremer support values multiplied by 100, values above branches represent group frequency values for symmetric
resampling. Males and females to scale with each other, but not among different species. Codes for undescribed specimens follow Table 1. Species are
labelled with the names that should be applied to them after the conclusions of our study.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 17
Iporanga, Parque Estadual Turístico do Alto Ribeira [242909.330S,
4838047.1100W], 12–18.x.2001, Biota team, 1 <(IBSP 97016); Peruíbe,
Esta¸cão Ecológica de Juréia-Itatins, Núcleo Arpoador (2423013.600S,
41103.300W, 13 m), 21–26.iv.2012, G. H. F. Azevedo and J. P. P. Pena-
Barbosa, 1 ,(UFMG 12666).
Additional material (not types). BRAZIL: Bahia: Ilhéus, 1 ,(IBSP
19025); Una, Reserva Biológica do Una, 1 ,(IBSP 46447), 1 ,(IBSP 47029),
1,(IBSP 46946). Minas Gerais: Marliéria, Parque Estadual do Rio Doce, 2 ,
(IBSP 94984), 1 ,(IBSP 94903), 1 ,(IBSP 94956).
Diagnosis
Micrathena perfida males differ from all other Micrathena
species, except for M. beta, by the elaborate shape of the
palpal tibia, the folded paracymbium, the large hook-shaped
conductor and by the spine-shaped digitiform projection of
the median apophysis (Fig. 7B,C; DP). It can be distinguished
from M. beta by the truncate embolus (Fig. 7B) (pointed and
curved in M. beta; Fig. 7D) and by the apex of the terminal
apophysis surpassing the tegulum edge (Fig. 7B,C; see also Levi
and Santos 2013:fig. 3). Females differ from all species of the
triangularispinosa-group, except M. beta, by the second pair of
posterior spines being anteriorly displaced (Fig. 6A; ps). They
differ from M. beta by the narrower and more pointed epigynal
lobe (Fig. 6F).
Description
Holotype male from Duque de Caxias, Rio de Janeiro (UFMG
19049) (Fig. 2E). Prosoma dark brown, light brown in the ocular
area and with two light brown markings in the pars thoracica;
clypeus pale yellow, chelicerae light brown and sternum dark
brown. Endites, labium and coxae yellow, femora proximally
yellow and distally dark brown; distal articles pale yellow.
Abdomen brown with a white band medially, and posteriorly
with dark patches on each side; venter dark brown with white
patches. Posterior median eyes the largest. Carapace glabrous
with rounded pars thoracica, and an elongated depression as a
fovea, without dimples, slightly flattened dorsoventrally in lateral
view. First coxae without hooks, and legs without macrosetae.
Abdomen rectangular, widest posteriorly, without spines. Total
length 3.7; carapace 1.78 long, 1.27 wide at its widest point;
abdomen 2.23 long; length of femur I 1.17, patella 0.41, tibia
0.81, metatarsus 0.61, tarsus 0.46; femur II 0.91, patella 0.46, tibia
0.76; femur III 0.86, patella 0.30, tibia 0.51; femur IV 1.47, patella
0.36, tibia 0.81, metatarsus 0.81, tarsus 0.51. Palp (Fig. 7B) with
membranous, apically flattened, spoon-like terminal apophysis.
Embolus thick and truncate in apex. Conductor with large
sclerotised hook, and a membranous bifid lobe close to apex.
Paramedian apophysis absent. Median apophysis fused to radix,
basal projection of the median apophysis well developed,
rhomboid and slightly sclerotised, with a proximal spine
posteriorly directed. Paracymbium enlarged and folded onto
itself, tibia modified, with three projections, one rounded, one
conical and another digitiform (Fig. 7C).
Female paratype from Esta¸
cão Ecológica da Juréia, Peruíbe,
São Paulo (UFMG 12666). Carapace light brown, with median
dark brown line and dark brown sides. Chelicerae black, and
light brown proximally. Endites and labium pale yellow. Sternum
dark brown with yellowish blots. Coxae yellowish, distal articles
of legs dark brown, femora yellowish on the bases and with a
longitudinal dark brown band. Dorsum of abdomen white, sides
white with dark line following the apodemes; venter white, with
a black sclerotised ring around the spinnerets. The four pairs of
spines yellowish. Carapace dome-shaped, glabrous and without
dimples; thoracic fovea circular. Abdomen triangular, widest
posteriorly. Four pairs of spines on the abdomen, the first
and the third the longest; the second pair of posterior spines
directed posterolaterally (Fig. 6A,B). Total length 6.14; carapace
2.05 long, 1.91 wide at its widest point; abdomen 4.9 long.
The specimen lacks the first and second legs on the left side,
and all the legs on the right side. Length of femur III 1.53, patella
Table 1. Data on the new Micrathena specimens described here and the new pairings proposed for males and females, which had been incorrectly
matched in the past
‘Collected with’refers to which specimens have been collected in the same localities as the specimens of dubious assignment; ‘final assignment’refers to
our final taxonomic decision; and ‘evidence’to the reasons why we propose the new pairings (P= phylogenetic position, L = locality data, B = DNA barcoding);
all localities in Brazil except where noted
Dubious specimens Collected with Localities Final assignment Evidence
Undescribed female #1 (Fig. 6)M. lata <Bahia; Minas Gerais; São Paulo; Rio de Janeiro M. perfida, sp. nov. LB
Micrathena lata male (Fig. 7A–C) Undescribed female #1 Santa Teresa (Espírito Santo); Rio de Janeiro M. perfida, sp. nov. LB
Undescribed female #2 (Fig. 8)M. beta <Pará; Porto Velho (Rondônia); Requena
(Loreto, Peru)
M. beta LB
Undescribed male #1 (Fig. 9)M. embira ,Porto Velho (Rondônia) M. embira LBP
Undescribed male #2 (Fig. 10)M. reimoseri ,Castro Alves (Bahia); Miguel Calmon (Bahia) M. reimoseri LBP
Undescribed male #3 (Fig. 11)M. exlinae ,Caracaraí (Roraima) M. exlinae LP
Undescribed male #4 (Fig. 12)M. miles ,Senador Guiomard (Acre); Maynas (Loreto, Peru) M. miles L
Undescribed male #5 (Fig. 13)M. cornuta ,Porto Velho (Rondônia); Tambopata
(Madre de Dios, Peru)
M. cornuta LB
M. cornuta male (Levi 1985:
figs 809–811)
M. woytkowskii ,Puerto Asis (Putumayo, Colombia) M. woytkowskii L
Undescribed male #6 (Fig. 14)M. spinulata ,Pico de Orizaba (Veracruz, Mexico) M. spinulata LBP
Undescribed male #7 M. yanomami ,Porto Velho (Rondônia) triangularispinosa
group
LBP
M. bicolor male (Levi 1985:
figs 453–455)
M. yanomami ,Juruti (Pará); Porto Velho (Rondônia) M. yanomami LBP
18 Invertebrate Systematics I. L. F. Magalhaes et al.
(A)
(B)
(D)
(E)
(F)
(C)
Fig. 6. Micrathena perfida, sp. nov. (A–E) Female paratype from Peruíbe, São Paulo, Brazil (UFMG 12666): (A) habitus, dorsal; (B) lateral view.
(C–E) Epigynum: (C) ventral; (D) lateral; (E) posterior. (F) Female from Una, Bahia, Brazil (IBSP 46447), epigynum lateral view. Scale bars = 1 mm
(A,B) and 0.1 mm (C–F). Abbreviation: PS, second posterior spine.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 19
(A)(C)
(B)
(D)
(E)
Fig. 7. (A–C)Micrathena perfida, sp. nov., male paratype from Santa Teresa, Espírito Santo, Brazil (MZSP 7982): (A) habitus, dorsal;
(B) palpal bulb, prolateral; (C) palpal bulb, retrolateral. (D,E)Micrathena beta, male from Requena, Loreto, Peru (MUSM-ENT 510054):
(D) palpal bulb (right, mirrored), prolateral; (E) palpal bulb, retrolateral. Scale bars = 1 mm (A) and 0.1 mm (B–E). Abbreviations: BP, basal
projection of the median apophysis; C, conductor; CL, conductor lobe; Cy, cymbium; DP, digitiform projection; E, embolus; P, paracymbium;
R, radix; ST, subtegulum; TA, terminal apophysis.
20 Invertebrate Systematics I. L. F. Magalhaes et al.
0.51, tibia 1.02, metatarsus 0.78, tarsus 0.51; femur IV 2.81,
patella 0.75, tibia 1.73, other articles missing. Epigynum with a
domed bulge, lacking posterior lobe (Fig. 6E,D). Posterior
median plate a little wider than long, mushroom-shaped and
with two well-separated, inflated black marks corresponding to
the region where the copulatory openings lie (Fig. 6C–E).
Female paratype from Duque de Caxias, Rio de Janeiro
(UFMG 19048). Colouration and abdomen shape as described
above. Length of femur I 2.59, patella 0.82, tibia 1.64, metatarsus
1.57, tarsus 1.02; femur II 2.23, patella 0.61, tibia 0.86; femur III
1.75, patella 0.61, tibia 1.01; femur IV 3.04, patella 0.76,
tibia 1.78, metatarsus 1.62, tarsus 0.86.
Intraspecific variation
Males (n= 2) vary in total body length from 3.6 to 3.7 and
carapace length from 1.62 to 1.78. The male from Santa
Teresa is lighter than the male from Duque de Caxias,
probably because of the better conservation state of the latter.
In the paratype female from Juréia (UFMG 12666), the right
second posterior spine was much smaller than the left one. This
female lacks the epigynum lobe, which is present on the other
analysed specimens (Fig. 6F). Total length varies from 5.59 to
6.14, carapace length from 2.05 to 2.52.
Distribution
This species is known from the Atlantic Forest in eastern Brazil,
from Bahia to São Paulo states (Fig. 15).
Remarks
The male of M. perfida was described by Levi (1985) under the
name M. lata. The only evidence he used for matching this male to
the female of M. lata was that both presented characteristics of
his militaris-group, and he admitted that the association was
uncertain. We here present evidence that the male identified by
Levi (1985)asM. lata actually should be matched to undescribed
female #2: they have similar COI sequences (Fig. 4) and have
been collected in the same locality at Duque de Caxias, Rio de
Janeiro. Also, the male is similar to M. beta and the female is
similar to undescribed female #1 (Figs 3,6–8), which are another
pair for which we have good evidence of a correct match
(see notes under M. beta below). As the holotype of M. lata is
a female, it is this sex that retains Chickering’s(1960) name. We
could not find any name that has been applied to the Atlantic
Forest specimens here described as M. perfida, and thus name it
for the first time.
Etymology
The specific name is a Latin adjective meaning ‘perfidious’or
‘treacherous’and refers to the fact that males of this species were
incorrectly matched to females of Micrathena lata for a long
period of time.
Micrathena beta di Caporiacco
(Figs 2A,3,5,7D,E,8A–E,15)
Micrathena beta di Caporiacco, 1947: 26 (holotype male from Two
Mouths, Essequibo River, Guyana, deposited in Museo di Storia
Naturale dell’Univerisità di Firenze, not examined); di Caporiacco,
1948: 668–669, fig. 81; Levi, 1985: 446 (transferred to Linyphiidae
incertae sedis); Levi & Santos, 2013: 223–224, figs 2–4 (returned to
Araneidae).
Micrathena triangularispinosa (De Geer): Levi, 1985:fig. 472
(misidentified in part, only some females).
Micrathena lata Chickering: Dierkens, 2011:figs 36, 44 (misidentified).
Material examined
BRAZIL: Acre: Cruzeiro do Sul, Parque Nacional da Serra do Divisor, 6 ,
(IBSP 12180, 12345, 12432, 20294, 20295); Senador Guiomard, Esta¸cão
Experimental Catuaba, 1 ,(UFMG 19038). Amazonas: Manaus, 1 <(MZSP
7687), 1 ,(IBSP 80052); Porto Urucu, 1 <(MPEG 20065). Pará: Bagre 1 <
(MPEG 31656), 7 ,(MPEG); Belterra, 1 <(MPEG 30309); Transgarimpeiro,
1<(MPEG 25000); Fazenda Euclides Gallo, 1 <(MPEG 30301); Juruti, 1 <
(MPEG 31666), 1 ,(MPEG 31667). Rondônia: Porto Velho, Abunã, 1 ,
(MZSP 39760), 1 ,(MZSP 35008), 1 ,(MZSP 35441), 1 ,(MZSP 35490),
1,(MZSP 41399); Porto Velho, Cai¸cara, 1 ,(MZSP 40267), 1 ,(MZSP
40407), 1 ,(MZSP 37867), 1 ,(MZSP); Porto Velho, Mutum, 1 ,(MZSP
35153); Porto Velho, 5 <(MZSP 40277, 40434, 40525, 44146, 50567), 3 ,
(MZSP 33907, 33911). ECUADOR: Napo: Napo-Galeras, 1 <(UFMG
3356). PERU: Loreto: Requena, Saquena, 1 <(MUSM-ENT 510054), 1 ,
(MUSM-ENT 510055), 1 ,(UFMG 19560); Requena, Jenaro Herrera, Centro
de Investigaciones Jenaro Herrera, 1 <(IBSP 165538).
Diagnosis
Male diagnosed by Levi and Santos (2013: 223): ‘This is an
unusual species of Micrathena, with the palpus having an
enlarged and modified paracymbium and a strangely modified
tibia (fig. 4). It closely resembles M. lata Chickering, 1960 (Levi
1985: 567, figs 614–615), but differs by having a narrower hook
as a conductor (fig. 3), a more elaborate paracymbium and a
slightly differently shaped tibia (Fig. 7D,E)’. Females resemble
M. perfida in having the second pair of posterior spines displaced
anteriorly (Fig. 8A; ps); they differ from it by the shape of the
epigynal lobe, which is thicker and more rounded in lateral view
in M. beta (Fig. 8D).
Description
For the male description, see Levi and Santos (2013).
Female from Requena, Loreto, Peru (UFMG 19560).
Carapace light grey, with median dark line and dark sides.
Chelicerae black. Endites black and labium dark brown.
Sternum yellowish with white and black blots. Coxae pale
yellow, remaining leg segments dark brown, yellowish on the
basis of femora. Dorsum of abdomen white, sides dark with
white spots, venter white with black blots and two dark red
blots on the sides of the sclerotised ring of the spinnerets, and
with black stripes in posterior view; sclerotised ring around
spinnerets black. Anterior spines pale yellow, lateral spines
black, the first pair of posterior spines orange-brown and
reddish, and the second posterior pair dark red. Carapace
dome-shaped, glabrous, without dimples; thoracic fovea an
indistinct circular marking. Abdomen trapezoidal, widest
posteriorly. Four pairs of spines on the abdomen, the first and
the third the longest; fourth pair directed posterolaterally
(Fig. 8A,B). Total length 5.32; carapace 2.18 long, 1.63 wide
at its widest point; abdomen 3.0 long; length of femur I 2.11,
patella 0.75, tibia 1.36, metatarsus 1.16, tarsus 0.55; femur II 1.91,
patella 0.68, tibia 1.30; femur III 1.36, patella 0.55, tibia 0.82;
femur IV 2.52, patella 0.68, tibia 1.50, metatarsus 1.43, tarsus
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 21
(E)
(D)
(C)
(A)
(B)
Fig. 8. Micrathena beta di Caporiacco, 1947, female from Requena, Loreto, Peru (MUSM-ENT 510054). (A) Habitus, dorsal; (B) lateral.
(C–E) Epigynum: (C) ventral; (D) lateral; (E) posterior. Scale bars = 1 mm (A,B) and 0.1 mm (C–E). Abbreviation: PS, second posterior spine.
22 Invertebrate Systematics I. L. F. Magalhaes et al.
0.68. Epigynum with a domed bulge, from which a rounded
lobe projects posteriorly. Posterior median plate as wide as long,
mushroom-shaped and with two well-separated, inflated, black,
kidney-shaped spots corresponding to the region where the
copulatory openings lie (Fig. 8C–E).
Intraspecific variation
The lobe of the epigynum is broken in a female from Requena.
Females (n= 5) vary in total length from 5.32 to 6.2; carapace
length varies from 2.11 to 2.50. The populations found in Porto
Velho have the half-distal part of the first pair of posterior spines
black, and they lack the red colour in the abdomen observed in the
population from Peru; individuals of these two colour variants
have been collected in the same locality in Juruti, Pará.
Distribution
This species is known from the western Amazon, from French
Guiana to Peru (Fig. 15).
Natural history
At Porto Velho, Rondônia, specimens were captured in the
understorey of dense ombrophilous forest at 50, 1050 and
2050 m from the Madeira River. In Requena, specimens were
collected in floodpain forest next to the Cumaceba River, with
sparse understorey; the webs were connected to shrubs.
Remarks
Females of this species have been misidentified as
M. triangularispinosa in collections, mainly because Levi
(1985) depicted a female M. beta as a variation of the first
species (see his figs 472, 473). We have checked the original
description of M. triangularispinosa and some of its synonyms
and none mention the anteriorly displaced posterior spine; so it
seems that di Caporiacco’s name has indeed been the first to be
applied to this species. Males and females of M. beta are here
matched because they have been collected in the same localities
(Bagres and Juruti, Pará; Porto Velho, Rondônia; Manaus,
Amazonas, Brazil; Requena, Loreto, Peru): at one of them the
male was hanging from a long thread connected to a female’sweb.
Also, males and females share almost identical COI sequences
(Fig. 4).
Micrathena embira Levi
(Figs 1E,F,3–5,9,15)
Micrathena embira Levi, 1985: 552, figs 551–555 ‘holotype female from
‘Mouth of rio Embira, Rio Jurura, N. Amazonia’(sic), Western
Amazon, Brazil, deposited in the American Museum of Natural
History, not examined).
Material examined
BRAZIL: Amazonas: Coari, Base de Opera¸cões Geólogo Pedro de Moura,
PortoUrucu, 1 ,(MPEG20061); São Gabrielda Cachoeira, Picoda Neblina,1
,(INPA 6290). Rondônia: Porto Velho, Abunã, 1 <(MZSP 39156), 1 ,
(MZSP 39942), 1 ,(MZSP 45587), 1 ,(MZSP 39155); Porto Velho, Cai¸cara,
1<(MZSP 40691), 1 ,(MZSP 34455), 1 <(MZSP 46089). Roraima:
Caracaraí, Arquipélago Mariuá, 1 ,(IBSP 165519).
Diagnosis
Males of M. embira share with other species in the schreibersi-
group the lack of a terminal apophysis (Fig. 9B) and a flattened
paracymbium (Fig. 9C,D). It can be distinguished from other
Micrathena species in this group by the rectangular abdomen,
much longer than wide and without a median constriction
(Fig. 9A); by the long and thin embolus held by the large and
membranous conductor (Fig. 9B); and by the large basal
projection of the median apophysis, which is divided into two
parts (Fig. 9B; BP). Female diagnosed by Levi (1985: 552): ‘The
fourteen spines of the abdomen (figs 551, 552) and the epigynum,
with its small projecting lobe on the anterior face of the globular
bulge (figs 553–555), separate M. embira from others of the
schreibersi-group’.
Description
Male from Abunã, Porto Velho, Rondônia, Brazil (MZSP 39156).
Carapace, sternum, labium, endites, chelicerae and abdomen
dark brown. Legs light brown, except for dark brown first
and second femora. Carapace coffin-shaped, wider at the
anterior third. Thoracic groove round and shallow. First coxa
with a hook and second femur with a corresponding prolateral
groove. First and second femora and tibiae ornamented
with strong macrosetae; second femora with single ventral row
of macrosetae. Abdomen rectangular, with two tiny tubercles at
the posterior end (Fig. 9A). Total length 6.58; carapace 2.29
long, 1.25 wide at its widest point; abdomen 4.41 long; length
of femur I 1.73, patella 0.48, tibia 1.19, metatarsus 0.97, tarsus
0.46; femur II 1.5, patella 0.41, tibia 0.94; femur III 1.02, patella
0.31, tibia 0.53; femur IV 2.03, patella 0.36, tibia 1.22, metatarsus
1.02, tarsus 0.53. Palp (Fig. 9B) lacking terminal apophysis.
Embolus thin, long, ending near tip of conductor. Conductor
long, narrow, membranous, extending beyond tegulum
margin. Paramedian apophysis present, digitiform. Conductor
lobe present, digitiform. Median apophysis with lobe, bent
triangular rim, and basal projection divided in two parts.
Paracymbium enlarged and folded, with flat retrolateral lobe
(Fig. 9C,D).
For female description, see Levi (1985).
Intraspecific variation
Males (n= 3) vary in total length from 5.92 to 6.58, carapace
length from 2.26 to 2.29 and femur I length from 1.73 to 1.75.
Females (n= 6) vary in carapace length from 3.0 to 3.5 and femur I
length from 3.07 to 3.5.
Distribution
This species is known from the western Amazon, in the states of
Roraima, Rondônia and Amazonas, Brazil (Fig. 15).
Natural history
A single female was captured at the base (100 m above sea level)
of Pico da Neblina, São Gabriel da Cachoeira, Amazonas (see
Nogueira et al.2014). At Porto Velho, Rondônia, 11 individuals
were collected, of which 10 were found in the sampling plots
located 50 m from the Madeira River, and one was found in a
sampling plot situated 1050 m away from the river. In both
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 23
localities, all individuals were captured in areas of terra firme old-
growth forest.
Remarks
The association of M. embira with its putative male seems
unequivocal. Both sexes were repeatedly collected in the same
area, and one male was found hanging from a female’s web
(S. Outeda-Jorge, pers. obs.). The morphological phylogeny
shows both males and females at a relatively basal position
in the schreibersi-group (Fig. 3). Finally, males and females
have similar COI sequences (Fig. 4). This is corroborated by
some morphological features present in both sexes, such as
the coffin-shaped, shiny and dark carapace and the colouration
of the legs. Interestingly enough, both sexes present features
synapomorphic for the schreibersi-group (such as epigynal keels
and lacking a terminal apophysis), but also plesiomorphic
features not shared by the higher members of this group (such
as a rounded tegulum, a coxal hook and straight spermathecae).
The original type locality is somewhat confusing; we interpret
it as the confluence between the Juruá and Envira rivers,
approximately at 7300S70
060W.
(B)
(A)
(C)
(D)
Fig. 9. Micrathena embira Levi, 1985, male from Porto Velho, Rondônia, Brazil (MZSP 39156). (A) Habitus, dorsal;
(B) palpal bulb, prolateral; (C) palpal bulb, retrolateral; (D) palpal bulb, subapical. Scale bars = 1 mm (A) and 0.1 mm
(B–D). Abbreviations: BP, basal projection of the median apophysis; C, conductor; Cy, cymbium; E, embolus; M, median
apophysis; P, paracymbium; PM, paramedian apophysis; R, radix; ST, subtegulum; Ti, palpal tibia.
24 Invertebrate Systematics I. L. F. Magalhaes et al.
Micrathena reimoseri Mello-Leitão
(Figs 2C,3–5,10,15)
Micrathena reimoseri Mello-Leitão, 1935: 97, fig. 14 (female lectotype
and one female paralectotype designated by Levi (1985) from
Petrópolis, Rio de Janeiro, Brazil, deposited in Museu Nacional do
Rio de Janeiro (MNRJ 41999), not examined); Levi, 1985: 565, 566,
figs 616–619.
Material examined
BRAZIL: Bahia: Castro Alves, Serra da Jibóia, 1 <(UFMG 19676) 1 ,
(CHNUFPI 1601); Miguel Calmon, Parque Estadual das Sete Passagens, 1 <
(IBSP 162301), 3 ,(UFMG 11031), 5 ,(IBSP 162302); Serra Bonita,
1,(MZSP).Espírito Santo: Santa Teresa, Reserva Biológica Augusto Ruschi,
1,(MZSP).
Diagnosis
Males of M. reimoseri are unique among Micrathena due to the
large tegular projection inserted near the base of the embolus
(Fig. 10B; TP) and the folded and complex conductor lobe
(Fig. 10B; CL). Female diagnosed by Levi (1985: 566): ‘This
species differs from M. lata by having the fourth pair of spines
minute (fig. 616, in M. lata they are half the length of the third
spine) and in having small knob on the tip on the epigynum
(fig. 617–619). They differ from others of the species-group by
the wide abdomen (fig. 616)’.
Description
Male from Parque Estadual Sete Passagens, Miguel Calmon,
Bahia, Brazil (IBSP 162301). Carapace yellowish orange, with
dark brown sides. Sternum, labium, endites and chelicerae
yellow. Legs orange, except for light yellow coxae and
trochanters. Abdomen dorsum light yellow with dark brown
lateral margins, venter orange. Carapace oval, with shallow
and round thoracic groove. Abdomen rectangular, slightly
wider in the posterior end. No coxal hook, femoral groove, or
strong macrosetae on any of the legs or tubercles at the posterior
end of the abdomen (Fig. 10A). Total length 3.29; carapace 1.4
long, 1.04 wide at its widest point; abdomen 1.88 long; length of
femur I 1.04, patella 0.38, tibia 0.79, metatarsus 0.64, tarsus 0.43;
femur II 1.04, patella 0.36, tibia 0.74; femur III 0.69, patella 0.2,
tibia 0.38; femur IV 1.12, patella 0.31, tibia 0.66, metatarsus 0.58,
tarsus 0.38. Palp (Fig. 10B) with a flat, sinuous terminal apophysis
(B)
(A)
(C)
Fig. 10. Micrathena reimoseri Mello-Leitão, 1935, male from Miguel Calmon, Bahia, Brazil (IBSP
162301). (A) Habitus, dorsal; (B) palpal bulb, prolateral; (C) palpal bulb, retrolateral. Scale bars = 1 mm
(A) and 0.1 mm (B,C). Abbreviations: C, conductor; CL, conductor lobe; Cy, cymbium; E, embolus;
M, median apophysis; P, paracymbium; R, radix; ST, subtegulum; TA, terminal apophysis; Ti, palpal tibia;
TP, tegular projection.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 25
ending in a rounded tip. Embolus thick, long, S-shaped with
accompanying membrane and distal knob. Conductor short,
hook-shaped, heavily sclerotised. Conductor lobe present,
complex, cup-shaped with posteriorly directed tip. Paramedian
apophysis absent. Median apophysis reduced to basal projection,
which bears a short projection pointing towards the embolus.
Paracymbium enlarged, laterally directed (Fig. 10C). Palpal tibia
with bifid prolateral projection.
For the female description, see Levi (1985).
Intraspecific variation
Females (n= 9) vary in carapace length from 3.1 to 3.5 and femur I
length from 3.5 to 4.05. The genitalic variation mentioned by Levi
(1985) has not been observed by us in specimens from Miguel
Calmon, Bahia.
Distribution
This species is known from the Atlantic rainforests of southern
and eastern Brazil, from Rio de Janeiro to Bahia states.
Natural history
Individuals from Castro Alves and Miguel Calmon, Bahia, were
collected in rainforest enclaves located in the Brazilian Caatinga,
a dry tropical forest not suitable for most Micrathena species.
Most females were found in webs in low (30–50 cm) plants with
a bromeliad-like architecture.
Remarks
Males and females have been collected in two areas in Bahia,
Brazil. The only other Micrathena collected in one of those areas
was M. digitata (C.L. Koch), an unrelated and very common
species whose male–female association is unequivocal. Both
sexes of M. reimoseri possess synapomorphic features of the
militaris-group (such as a reduced median apophysis, palpal tibia
with projections, and abdomen apodemes arranged in a single
line) and are therein placed (Fig. 3). Furthermore, the COI
sequences of males and females are nearly identical (Fig. 4).
Micrathena yanomami Magalhães & Santos
(Figs 1C,D,3,5,15)
Micrathena bicolor Keyserling: Levi, 1985 (partially misidentified, males
only).
Micrathena triangularispinosa (De Geer): Dierkens, 2011:figs 8, 23
(misidentified).
Micrathena yanomami Magalhães & Santos, 2011: 43, figs 9–13
(holotype female from Bebedouro Novo, Pico da Neblina, São
Gabriel da Cachoeira, Amazonas, Brazil, deposited in INPA 6286,
examined).
Material examined
BRAZIL: Acre: Senador Guiomard, Fazenda Experimental Catuaba, 2 ,
(UFMG 10623, 15518). Amazonas: Coari, Porto Urucu, 1 ,(MPEG 20062);
São Gabriel da Cachoeira, Pico da Neblina, 2 ,(INPA 6286, holotype; INPA
6287, paratype). Pará: Fazenda Euclides Gallo, 1 ,(MPEG 24944); Juruti, 1 ,
(MPEG31673), 1 <(MPEG 31674); ParNa da Amazônia, 1 ,(MPEG30300).
Rondônia:Porto Velho, Abunã, 1 ,(MZSP 34041),7 <(MZSP 43160,50839,
51345, 51601, 53727, 55224, 55266), 1 ,(MZSP 34042), 1 ,(MZSP
34515), 1 ,(MZSP 37715), 1 ,(MZSP 46381), 1 ,(MZSP 38718), 1 ,
(MZSP 34933); Porto Velho, Mutum, 1 ,(MZSP 38142). PERU: Cusco:
Paucartambo, Kosñipata, 1 ,, 1 juvenile (MUSM-ENT 510056), 1 ,,
1 juvenile (MUSM-ENT 510060), 1 ,(UFMG 15502), 1 ,(UFMG
19566). Madre de Dios: Tambopata, Inotawa Lodge, La Torre, 1 ,
(MUSM-ENT 510059), 1 ,(UFMG 19567). Loreto: Iquitos, San Juan
Bautista, Zungarococha, 1 <(IBSP 165534).
Diagnosis
Males differ from all other species by having a black band on the
carapace, starting in the ocular region, passing in the lateral edges
of the pars cephalica and nearly touching each other near the
pedicel insertion (Fig. 4). They also differ from the other species
of the triangularispinosa-group by the wide and weakly
sclerotised lobe of the conductor (Levi 1985:fig. 454) and
by the shape of the paracymbium, which is divided into two
lobes, the dorsal one with an indentation (Levi 1985:fig. 455).
Females diagnosed by Magalhães and Santos (2011: 43):
‘Micrathena yanomami females differ from the other species
of the triangularispinosa-group by the colouration pattern of
the carapace, with a black band along its entire edge, by the
granulation in the carapace edges and by the setal bases of
the femora, which are more projected and domed (figs 9, 10).
The epigynum is similar to that of M. triangularispinosa
(De Geer) (Levi 1985:figs 474–476), but differs by the more
robust lobe, which is also more detached from the bulge (figs 11,
12), and by the median plate of the epigynum in posterior view,
which is wider than long and has a mushroom-shaped clear area
embracing two dark spots (fig. 13)’.
Description
Male described by Levi (1985) under the name Micrathena
bicolor. Female described by Magalhães and Santos (2011).
Intraspecific variation
Females vary in total length from 5.92 to 6.58 (n= 3), carapace
length from 2 to 2.5 and femur I length from 1.14 to 1.86 (n= 9).
The females collected in Cusco, Peru, have a rather deviant
somatic morphology: the carapace is uniformly dark, they
have three pairs of lateral spines (a characteristic not found in
the entire swainsoni clade; Magalhães and Santos 2012), and the
abdomen and legs do not have the colouration pattern typical of
this species (Fig. 1C; compare with Fig. 1B). These were initially
thought to represent a different species, but genital morphology
and COI sequences (Fig. 4; Myanomami193) suggest they are a
morphological variant of M. yanomami.
Distribution
This species is known from the Amazon, in French Guiana (see
Dierkens 2011; under M. triangularispinosa; these records have
not been included in the map because we could not verify their
exact placement), Amazonas, Acre and Rondônia states, Brazil,
and Cusco, Loreto and Madre de Dios states, Peru.
Natural history
At Pico da Neblina, 17 individuals were collected. Four were
found at the first altitude sampled, at the base of the mountain
(100 m above sea level), two were located at 400 m, and the
remaining 11 were captured at 860 m. At Porto Velho, specimens
26 Invertebrate Systematics I. L. F. Magalhaes et al.
were captured throughout the entire year. Of the 111 individuals
captured, most of them (86) were found in the sampling plots 50 m
from the Madeira River, while 12 were located at 1050 m and 13
were captured 2050 m from the river. In both localities, all
individuals were captured in areas of terra firme old-growth
forest.
Remarks
We here present evidence that the male previously assigned to
M. bicolor by Levi (1985) is actually the male of M. yanomami.
We had initially thought that undescribed male #7 belonged with
M. yanomami; however, both phylogenetic placement (Fig. 3)
and COI sequences (Fig. 4) show that M. yanomami females
are closer to M. bicolor males than to male #7. Moreover, we also
have evidence for a M. yanomami female–M. bicolor male
association based on a 4-year spider inventory conducted in
Porto Velho, Rondônia (AAN, unpubl. data). During this
period, we collected 13 male specimens of M. bicolor and not
a single female, which is very unusual given that females are
more conspicuous and easier to collect than males. On the other
hand, 121 female individuals of M. yanomami were obtained in
this inventory. Micrathena bicolor males and M. yanomami
females have also been collected in Juruti, Pará. Levi (1985:
536) observed that ‘the placement of the male with M. bicolor is
uncertain’. Although we do not have genetic data for M. bicolor
females (currently known from two localities in Colombia),
current evidence is stronger in favour of the association of this
malewith M. yanomamithan it is with M. bicolor. The true male of
Micrathena bicolor is thus unknown and remains to be found.
On the other hand, we could not confidently associate male #7
with any species, and its placement must await a revision of the
triangularispinosa-group.
Micrathena exlinae Levi
(Figs 3–5,11,15)
Micrathena exlinae Levi, 1985: 514, 516, figs 342–346 (female holotype
and female paratype from Cucharas, Huallaga Valley, Huánuco, Peru,
deposited at the MCZ, not examined).
Material examined
BRAZIL: Amazonas: São Gabriel da Cachoeira, Pico da Neblina,1 ,(INPA),
1,(INPA), 1 ,(INPA), 1 ,(INPA), 2 ,(INPA), 1 ,(INPA), 1 ,(INPA), 1 ,
(INPA), 1 ,(INPA), 1 ,(IBSP), 1 ,(IBSP), 2 ,(IBSP), 1 ,(IBSP). Roraima:
Caracaraí, Arquipélago Mariuá, 1 <(IBSP 165527), 4 ,(IBSP 165521);
Tatocuara, 1 ,(IBSP 165522).
Diagnosis
Males of M. exlinae resemble the unrelated M. patruelis
(C. L. Koch, 1839) due to the shape of the terminal apophysis,
which is anteriorly rounded and acuminate distally (Fig. 11B;
TA). They differ from M. patruelis and all other Micrathena with
similar palpi by the combination of a smoothly rounded edge
of the median apophysis rim (Fig. 11B; M) and by a large and
distally widened paramedian apophysis (Fig. 11B; PM). Female
diagnosed by Levi (1985: 514, 516): ‘This species differs from
M. huanuco by having a bottle-shaped septum on the posterior
face of the epigynum (fig. 345). It differs from M. triangularis by
having only two posterodorsal spines (figs 342, 343)’.
Description
Male from Comunidade Caicubi, Caracaraí, Roraima, Brazil
(IBSP 165527). Carapace yellow, with lateral dark brown
bands. Sternum whitish yellow, labium, endites and chelicerae
yellow. Legs yellow at the base, gradually becoming brown in the
three apical articles. Abdomen whitish yellow, with lateral dark
brown bands ventrally and dorsally. Carapace rounded, with
round thoracic groove. Abdomen rectangular. First coxa with a
hook and second femur with a corresponding prolateral groove.
First femur with a row of dorsal macrosetae and first and second
tibiae ornamented with ventral macrosetae. No tubercles in the
abdomen (Fig. 11A). Total length 3.43; carapace 1.73 long, 1.37
wide at its widest point; abdomen 2.19 long; length of femur I
1.55, patella 0.48, tibia 1.17, metatarsus 0.99, tarsus 0.43; femur II
1.27, patella 0.41, tibia 0.89; femur III 0.79, patella 0.28, tibia
0.48; femur IV 1.53, patella 0.38, tibia 0.94, metatarsus 0.97,
tarsus 0.41. Palp (Fig. 11B) with short terminal apophysis,
with rounded apex that acuminates distally. Embolus thick,
sclerotised, partially fused to terminal apophysis. Radix drop-
shaped. Conductor short, folded near tip of embolus. Conductor
lobe present, digitiform. Paramedian apophysis present, large
and wide, with subsquarish tip. Median apophysis with small
lobe, gently rounded and bent rim, and large but lightly sclerotised
basal projection. Tegular projection small. Paracymbium
enlarged, but simple (Fig. 11C).
For the female description, see Levi (1985).
Intraspecific variation
Females (n= 3) vary in carapace length from 2.0 to 2.1 and femur I
length from 2.3 to 2.4.
Distribution
This species is known from the western Amazon, in Brazil and
Peru.
Natural history
At Pico da Neblina, São Gabriel da Cachoeira, Amazonas, all the
15 females collected were found at the base of the mountain
(100 m above sea level) in an area covered by terra firme old-
growth forest.
Remarks
The single putative male (undescribed male #3) did not amplify
for the COI fragment we used. It was collected in the same area as
five adult females of M. exlinae. Both sexes present features
synapomorphic with the plana-group (e.g. a folded conductor and
four pairs of lateral abdominal spines). This male–female
association must be taken with caution, as there are several
species of the plana-group known only from females (notably
M. marta Levi, 1985;M. bananal Levi, 1985;M. alvarengai
Levi, 1985; and M. huanuco Levi, 1985). Although none has
been recorded from the particular region where this male was
collected, it is possible that some of them have wider distributions
than those reported by Levi (1985). However, the fact that
M. exlinae females are common in the region suggests that our
association is correct.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 27
Micrathena miles Simon
(Figs 1A,B,3–5,12,15)
Micrathena miles Simon, 1895: 852, fig. 907 (female syntypes from Fonte
Boa and Tefé, Amazonas, Brazil, deposited at the Museum National
d’Histoire Naturelle, Paris (catalogue 1172), not examined); Simon,
1897: 468; Levi, 1985: 482–483, figs 164–168.
Micrathena cuminamensis Mello-Leitão, 1930: 62, fig. 16. Synonymised
by Levi, 1985;
Micrathena miles nigra di Caporiacco, 1948: 667. Synonymised by Levi,
1985.
Material examined
BRAZIL: Acre: Senador Guiomard, Esta¸cão Experimental Catuaba 2 ,
(UFMG 15648), 5 ,(UFMG 15611), 1 <(UFMG 15612). Rondônia:
Porto Velho, Cai¸cara, 1 ,(MZSP 35448), 1 ,(MZSP 33394). PERU:
Loreto: Maynas, Indiana, near San Rafael, 1 <(MUSM-ENT 510057), 1 ,
(MUSM-ENT 510062); Iquitos, San Juan Bautista, Zungarococha, 1 ,(IBSP
165530).
Diagnosis
Males of M. miles are most similar to those of M. macfarlanei
and M. kirbyi due to the black pigmentation around the eyes, the
presence of a retrolateral membrane on the conductor, and the
basal projection of the median apophysis, which is distally
detached and points towards the embolus. It differs from these
two species by the shorter, more rounded terminal apophysis
(Fig. 12B; TA) and by the median apophysis rim, whose apex is
more acuminate (Fig. 12B; M). Female diagnosed by Levi (1985:
483): ‘This species differs from Micrathena raimondi by lacking
a frame on each side of the transverse barand lobe of the epigynum
(fig. 166). It differs from M. kirbyi by lacking anterior spines
overhanging the carapace (fig. 165)’.
Description
Male from Senador Guiomard, Acre, Brazil (UFMG 15612).
Carapace orange, suffused with black, ocular area black.
(B)
(A)
(C)
Fig. 11. Micrathena exlinae Mello-Leitão, 1935, male from Caracaraí, Roraima, Brazil (IBSP 165527).
(A) Habitus, dorsal; (B) palpal bulb, prolateral; (C) palpal bulb, retrolateral. Scale bars = 1 mm (A) and
0.1 mm (B,C). Abbreviations: BP, basal projection of the median apophysis; C, conductor; Cy, cymbium;
E, embolus; M, median apophysis; P, paracymbium; PM, paramedian apophysis; R, radix; ST, subtegulum;
TA, terminal apophysis.
28 Invertebrate Systematics I. L. F. Magalhaes et al.
Chelicerae, endites, labium and sternum black. Legs I and II light
brown, III and IV dark brown. Abdomen dark brown ventrally,
dorsally beige with four median dark-brown spots. Carapace
coffin-shaped, wider at the anterior third. Carapace with a
shallow, round thoracic groove. First coxa with a hook and
second trochanter with a corresponding prolateral groove. First
femur with two ventral rows of macrosetae, stouter on the apical
quarter. First and second tibiae with two ventral rows of stout
macrosetae. Abdomen subrectangular, truncated anteriorly,
gently rounded posteriorly, without tubercles (Fig. 12A). Total
length 5.12; carapace 2.23 long, 1.52 wide at its widest point;
abdomen 2.89 long; length of femur I 2.23, patella 0.51, tibia 1.37,
metatarsus 1.57, tarsus 0.61; femur II 1.83, patella 0.41, tibia 1.01;
femur III 1.02, patella 0.34, tibia 0.54; femur IV 2.99, patella 0.46,
tibia 1.78, metatarsus 2.28, tarsus missing. Palp (Fig. 12B) with
short,mango-shaped terminal apophysis. Embolus partially fused
to terminal apophysis. Conductor large, sclerotised with a pointed
apex. Conductor lobe present, digitiform, slightly truncate on the
apex. Paramedian apophysis present, narrow, digitiform. Median
apophysis with large lobe, pointed and bent rim, and large and
heavily sclerotised basal projection, which points towards
embolus. Tegular projection small. Paracymbium enlarged but
simple (Fig. 12C).
For the female description, see Levi (1985).
Intraspecific variation
Males (n= 2) vary in total length from 4.09 to 5.12 and in carapace
length from 1.74 to 2.23.
Distribution
Known from the Amazon, in Brazil, Guyana and Peru.
(A)
(B)
(C)
Fig. 12. Micrathena miles Simon, 1895, male from Maynas, Loreto, Peru (MUSM-ENT 510057).
(A) Habitus, dorsal; (B) palpal bulb, prolateral; (C) palpal bulb, retrolateral. Scale bars = 1 mm (A) and
0.1 mm (B,C). Abbreviations: BP, basal projection of the median apophysis; C, conductor; CL, conductor
lobe; CM, conductor retrolateral membrane; Cy, cymbium; E, embolus; M, median apophysis; P,
paracymbium; PM, paramedian apophysis; R, radix; ST, subtegulum; TA, terminal apophysis.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 29
Natural history
A female from Maynas has been observed with a white, disc-like
egg sac attached to the underside of a leaf (Fig. 1A). At Porto
Velho, Rondônia, 30 individuals were captured throughout
the year. Most of them (26) were found at sampling plots
located 50 and 1050 m from river banks.
Remarks
We were unable to extract DNA from the putative males
(undescribed male #4), which came from two areas (one in
Peru and another in Brazil) where several females of
this species were collected. Regarding the morphological
phylogeny, females were inferred as members of the guerini-
group, while males were members of the kirbyi-group (Fig. 3).
However, females are genetically similar to M. macfarlanei
Chickering, which belongs in this latter group (Fig. 5), and a
preliminary phylogeny based on three molecular markers
strongly suggests the placement of M. miles females in the
kirbyi-group (ILFM, unpubl. data). It seems to us that the
morphological placing of M. miles females in the guerini-
group we observed (Fig. 3) is an artefact caused by missing
data, particularly of the key characters associated with the male
genitalia. With this in mind, it seems likely that this undescribed
male does indeed belong with M. miles. Due to an unfortunate
laboratory incident, the male specimen from Peru has been lost;
the only part remaining is its left palp, which was being kept in a
separate vial.
Micrathena cornuta (Taczanowski)
(Figs 2B,3–5,13,15)
Acrosomacornuta Taczanowski, 1873: 268, pl. 5, fig. 22 (female holotype
from Cayenne, French Guiana (Polska Akademia Nauk, Warszawa),
not examined).
Chaetacis cornuta (Taczanowski): Simon, 1895: 854, figs 920, 921;
Soares & Camargo, 1948: 379, fig. 36; Levi, 1985: 604, figs
(B)
(A)
(C)
Fig. 13. Micrathena cornuta (Taczanowski, 1873), male from Requena, Loreto, Peru (UFMG 19561).
(A) Habitus, dorsal; (B) palpal bulb (right, mirrored), prolateral; (C) same, retrolateral. Scale bars = 1 mm
(A) and 0.1 mm (B,C). Abbreviations: BP, basal projection of the median apophysis; C, conductor; CL,
conductor lobe; CM, conductor retrolateral membrane; Cy, cymbium; DL, dorsal lobe of the paracymbium;
E, embolus; M, median apophysis; P, paracymbium; R, radix; ST, subtegulum; TP, tegular projection.
30 Invertebrate Systematics I. L. F. Magalhaes et al.
803–808 (in part, only females); Dierkens, 2011: 102, figs 17, 38, 47,
50.
Micrathena cornuta (Taczanowski): Magalhães & Santos, 2012: 31.
Material examined
BRAZIL: Acre: Cruzeiro do Sul, Parque Nacional da Serra do Divisor, 1 ,
(IBSP 12404); Sena Madureira, Rio Purus, 1 ,(MCZ 91711); Rio Branco,
Reserva Extrativista Humaitá, 2 ,(IBSP 15716); Senador Guiomard, Esta¸cão
Experimental Catuaba, 3 ,(UFMG 15608, 15644). Pará: Bagre, 1 <(MPEG
31655); Paragominas, Aldeia Araw, 1 <,1,(MZSP 3273); Cachoeira, 1 ,
(IBSP 42). Rondônia: Porto Velho, 2 <,2,(MZSP 33864, 38362, 41312,
42432). Roraima: Alto Alegre, Aldeia Yanomami Palimi-U, 1 ,(IBSP
55482). PERU: Madre de Dios: Tambopata, Tambopata, La Torre,
Inotawa Lodge, 1 <(UFMG 19562), 2 ,(UFMG 19563); Manu, Madre
de Dios, Reserva Particular Bonanza, 1 <1,(MUSM-ENT 510058), 1 <
(UFMG 19561), 1 ,(UFMG 19564). Loreto: Requena, Saquena, Río
Cumaceba, 1 ,(UFMG 19565); Requena, Jenaro Herrera, Centro de
Investigaciones Jenaro Herrera, 1 juvenile (IBSP 165539).
Diagnosis
Males are most similar to those of M. cucharas (Levi, 1985) and
M. woytkowskii (Levi, 1985), due to the very long tegular
projection, slightly bent at the apex, and by the shape of the
paracymbium (Fig. 13B,C). They can be differentiated from them
by the shape of the basal projection of the median apophysis:
undetached in M. cucharas (see Levi 1985:fig. 819), hook-
shaped in M. woytkowskii (see Levi 1985:fig. 810), and round
and tooth-shaped in M. cornuta (Fig. 13B; BP). Also, the
sclerotised tip of the conductor is thick and bifid, with a hook-
shaped apical projection in M. cornuta (visible in mesal view;
Fig. 13B) (thinner and simple in M. cucharas and
M. woytkowskii). Finally, M. cornuta also can be differentiated
from M. cucharas and M. woytkoskii by the more triangular,
tapering shape of the retrolateral lobe of the paracymbium
(Fig. 13C). Female diagnosed by Levi (1985: 604): ‘Females
differ from most other species by having a spine on each side of
the head and by having only twelve spines on the abdomen
(figs 803, 804). It differs from M. woytkowskii by having the
abdomen squarish and the truncate eqigynum [sic](figs 804,
808)’.
Description
Male from Reserva Particular Bonanza, Manu, Madre de Dios,
Peru (UFMG 19561). Carapace, chelicerae, legs, abdomen,
endites, labium and sternum orange-brown. Carapace with a
round thoracic groove, one pair of dimples, two pairs of sulci
and tiny denticles on the edge and dorsum of the pars thoracica
(Fig. 13A). First coxa without hook; fourth coxa with denticles.
First pair of tibiae straight, not modified; first and second pairs
of tibiae with a few strong macrosetae, only on the ventral side
on the second pair; other legs without strong macrosetae in the
tibiae; femora with strong macrosetae in the distal extremity
(Fig. 13A). Abdomen rectangular, widest posteriorly, with six
pairs of spines. First, third and sixth pairs of spines orange-brown,
the remaining black (Fig. 13A). Total length 3.2; carapace 1.29
long, 0.95 wide at its widest point; abdomen 1.66 long; length
of femur I 0.95, patella 0.37, tibia 0.81, metatarsus 0.75, tarsus
0.41; femur II 0.95, patella 0.34, tibia 0.68; femur III 0.61,
patella 0.24, tibia 0.37; femur IV 1.08, patella 0.31, tibia
0.68, metatarsus 0.71, tarsus 0.37. Palp without terminal or
paramedian apophyses. Embolus long, sclerotised, slightly and
proximally curved. Conductor with sclerotised apex bifurcated,
apical projection hook-shaped, membranous and spiniform
lobe parallel to embolus, and membranous basal projection.
Median apophysis lightly sclerotised, except for dark and
tooth-shaped basal projection. Tegular projection long and
strongly bent in apex (Fig. 13B). Paracymbium with rounded,
lightly sclerotised dorsal lobe and tapering retrolateral lobe,
which is highly sclerotised (Fig. 13C).
For female description, see Levi (1985).
Intraspecific variation
Males (n= 6) vary in total length from 2.81 to 3.32; carapace
length varies from 1.22 to 1.39. The orange colouration of the
specimen in Fig. 2Bis very unusual; most females have dark-
brown carapace and legs.
Natural history
At Porto Velho, Rondônia, specimens were captured in the
understorey of dense ombrophilous forest at 50, 1050 and
2050 m from the Madeira River. In Peru, specimens were
collected in primary and secondary floodplain forests, in webs
in the understorey next to rivers (100–200 m). In Tambopata,
a male was found in a female’s web.
Distribution
This species is known from the Amazon, from French Guiana and
Pará, Brazil, to Peru. The records from Colombia by Levi (1985)
were misidentified and are of M. woytkowskii (see below).
Comparative M. cucharas material examined (all identified
by Levi 1985)
PERU: Huánuco: Cucharas, Huallaga Valley, 2 <,2,(CAS 9058771); Santa
Tereza, Huallaga Valley, 1 <(CAS 9058775); Tingo Maria, 3 <,6,(CAS
9058772, 9058773, 9058774, 9058776, 9058777, 9058778).
Remarks
We have re-examined a male Levi (1985) apparently used for
his description of M. cornuta, but it was stored in the same vial as
a female of M. woytkowskii (MCZ 91700). On the other hand,
females of M. cornuta are frequently collected in the same
areas as undescribed male #5 (Porto Velho, Rondônia, Brazil;
Tambopata, Madre de Dios, Peru; French Guiana: see Dierkens
2011). In the above-referenced 4-year inventory at Porto Velho,
four specimens of undescribed male #5 and 64 females of
M. cornuta were collected, and on two occasions both were
collected in the same sampling unit (one hour of nocturnal
hand searching). However, not a single male of M. cornuta
(sensu Levi 1985) was collected in the same area.
Furthermore, the genetic distance between undescribed male
#5 and M. cornuta females is among the smallest of the genus
–only 0.8% (Fig. 4). These findings strongly suggest undescribed
male #5 is, in fact, that of M. cornuta. As a consequence, the male
described as M. cornuta by Levi (1985) is, in fact, M. woytkowskii
(see below). The true male of M. cornuta was first illustrated by
Dierkens (2011:figs 38, 47).
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 31
Micrathena woytkowskii (Levi)
Chaetacis woytkowskii Levi, 1985: 608, figs 821–826 (female holotype
and 10 female paratypes from Cucharas, Huallaga Valley, Huánuco,
Peru (MCZ), not examined).
Chaetacis cornuta: Levi, 1985: 608, figs 809–811 (misidentified in part,
only males).
Micrathena woytkowskii: Magalhães & Santos, 2012: 31.
Material examined
COLOMBIA: Putumayo: Puerto Asis, Río Putumayo, 1 <,2,(MCZ 91700,
91725).
Diagnosis
Male diagnosed by Levi (1985: 604) under the name Chaetacis
cornuta:‘The paracymbium of the male has a diagnostic hook
(fig. 811)’. Female diagnosed by Levi (1985: 608): ‘Chaetacis
woytkowskii differs from C. cornuta by having a longer abdomen,
the second and fourth spines subequal in size (figs 821, 822), and
by having the epigynum more cone-shaped in profile (figs 825,
826); it differs from M. cucharas by having spines rather than
tubercles behind the eyes (figs 821, 822)’.
Description
Male described by Levi (1985) under the name Chaetacis cornuta
(see his figs 809–811). Female described by Levi (1985).
Distribution
This species is known from Peru and Colombia.
Remarks
Levi (1985) examined males and females of this species from
Colombia and misidentified them as M. cornuta. We have re-
examined some of these specimens seen by him, and the females
fit with his diagnosis of M. woytkowskii, while the males fit his
diagnosis of M. cornuta. We here describe the true male of
(B)
(A)
(C)
Fig. 14. Micrathena spinulata F.O. Pickard-Cambridge, 1904, male from Pico de Orizaba, Veracruz,
Mexico (IBSP). (A) Habitus, dorsal; (B) palpal bulb (right, mirrored), prolateral; (C) same, retrolateral. Scale
bars = 1 mm (A) and 0.1 mm (B,C). Abbreviations: BP, basal projection of the median apophysis;
C, conductor; CL, conductor lobe; Cy, cymbium; DP, digitiform projection of the median apophysis;
E, embolus; P, paracymbium; R, radix; ST, subtegulum; TA, terminal apophysis.
32 Invertebrate Systematics I. L. F. Magalhaes et al.
Fig. 15. Geographic distribution records of the species with newly matched males and females. Circles represent previously known records and stars
represent new records. Scale bars = 500 km.
Matching sexes in dimorphic Micrathena spiders Invertebrate Systematics 33
M. cornuta (see above), thus the male described by Levi as
M. cornuta becomes unpaired. As it has been collected with
females of M. woytkowskii, it seems more parsimonious that it
belongs with the latter species.
Micrathena spinulata F.O. Pickard-Cambridge
(Figs 3–5,14,15)
Micrathena spinulata F.O. Pickard-Cambridge, 1904: 530, pl. 50, fig. 7
(four female syntypes from Amula, Guerrero, Mexico, and one
syntype from Omilteme, Guerrero, WSW of Chilpancingo, 2400 m
alt., deposited at the British Museum of Natural History, not
examined); Chickering, 1961: 459, figs 181–184; Levi, 1985: 592,
594, figs 751–755.
Material examined
MEXICO: Veracruz: Pico de Orizaba Volcano, Atotonilco de Calcahuaco, 2
<,2,(IBSP 166385–166388).
Diagnosis
Males differ from those of M. gracilis (Walckenaer, 1805) and
M. horrida (Taczanowski, 1873) by having the posterior end of
the abdomen entire (Fig. 14A), rather than pseudosegmented.
They are most similar to those of M. forcipata (Thorell, 1859)
due to the enlarged and curved conductor and the broad and
blunt paracymbium; they can be distinguished by the strongly
curved embolus, visible just below the terminal apophysis
(Fig. 14B; E), and by the more curved digitiform projection of
the median apophysis (Fig. 14B;DP). Female diagnosed by Levi
(1985: 594): ‘Micrathena spinulata differs from M. striata and
M. margerita by having all six spines the same size (figs 751,
752)’.
Description
Male from Pico de Orizaba Volcano, Veracruz, Mexico (IBSP
166386). Carapace light brown, with yellowish white pars
cephalica. Sternum and labium dark brown. Endites light
brown. Chelicerae light brown, darker at apex. Legs yellowish
brown. Abdomen white, with light-brown median markings.
Carapace suboval, widest at pars thoracica, with subsquarish
pars cephalica and round thoracic groove. First coxa without
hook and second femur without groove. Legs without
macrosetae, except for a line of 3 or 4 macrosetae on the
dorsum of femora I and II. Abdomen rectangular, with
posterior end not pseudosegmented and with a pair of tiny
tubercles at the posterior end (Fig. 14A). Total length 2.81;
carapace 1.3 long, 0.94 wide at its widest point; abdomen 1.7
long; length of femur I 1.3, patella 0.43, tibia 0.99, metatarsus
0.86, tarsus 0.48; femur II 1.17, patella 0.38, tibia 0.79; femur III
0.71, patella 0.23, tibia 0.43; femur IV 1.17, patella 0.33, tibia
0.76, metatarsus 0.79, tarsus 0.38. Palp (Fig. 14B) with short,
bifid terminal apophysis with subsquarish apex. Embolus
sinuous, thicker near base, partially fused to terminal apophysis.
Conductor projecting anteriorly and with sclerotised apex pointing
posteriorly. Conductor lobe present, subsquarish and pointing
dorsally. Paramedian apophysis absent. Median apophysis
reduced to basal projection, which bears a long, curved and
unsclerotised digitiform projection. Tegular projection wide,
subsquarish. Paracymbium enlarged, but simple (Fig. 14C).
For the female description, see Levi (1985).
Intraspecific variation
Females (n= 2) vary in carapace length from 1.78 to 1.88. Males
(n= 2) vary in total length from 2.81 to 2.90 and in carapace length
from 1.27 to 1.30.
Distribution
This species is known from Central Mexico.
Natural history
Label data state that specimens were collected in Quercus forest
2300 m above sea level using the ‘looking-up’method.
Remarks
We first became aware that the undescribed male had been
collected through the Araneomorphae of Mexico Digital
Images Catalogue (Alvarez-Padilla Laboratory 2014), which
reports males and females repeatedly collected together in the
same inventory in Veracruz, Mexico. The morphological
phylogeny suggests that both the putative male and the female
fall within the gracilis-group (Fig. 3). Also, they do bear some
morphological resemblance to each other, especially regarding
the shape and colour of carapace and legs. Finally, COI sequences
are almost identical in males and females (Fig. 4).
New records
During the course of this study, we found some specimens
representing new species records for Argentina and Peru. They
are detailed below.
Micrathena bandeirante (Magalhães & Santos)
Chaetacis bandeirante Magalhães & Santos, 2011:49–53, figs 20–40
(holotype male from Usina Hidrelétrica Engenheiro Sérgio Motta,
Presidente Epitácio [21450S52
050W, 310 m], São Paulo, Brazil,
Equipe IBSP, 16.i–13.ii.1999, deposited in IBSP 23255. Paratypes:
female from the same locality (IBSP 160897); two males and
four females from the same locality (IBSP 160898); one male and
two females from Base de Pesquisa do Instituto Brasileiro de
Desenvolvimento Florestal, Poconé [16150S56
370W, 142 m],
Mato Grosso, Brazil, U. A. Drumond, 16.ii.1984 (MZSP 11443);
two males and two females from Passo do Lontra, Corumbá [1900S
57390W, 118 m], Mato Grosso do Sul, Brazil, J. Raizer et al. iv.1998
(UFMG 4889); one female from Nhecolândia [19140S57
020S, 86 m],
S. Haris, 11.xi.1987 (MNRJ 14587); all examined).
Material examined
ARGENTINA: Formosa: Pirané, 2 ,(MACN 32539).
Micrathena annulata Reimoser
Micrathenaannulata Reimoser, 1917: 149, 150, pl. 9, fig. 31 (nine females
and one juvenile syntypes from Santa Catarina, Brazil, deposited in
Naturhistorisches Museum, Wien, not examined; see World Spider
Catalog (2016) for complete taxonomic citation history).
Material examined
ARGENTINA: Misiones: Cainguás, Parque Provincial Salto Encantado, 1 ,
(MACN 32548).
34 Invertebrate Systematics I. L. F. Magalhaes et al.
Micrathena lata Chickering
Micrathena lata Chickering, 1960:6,figs 8–12 (female holotype from
Teresópolis, Rio de Janeiro, Brazil, deposited in MCZ, not examined;
see World Spider Catalog (2016) for complete taxonomic citation
history).
Material examined
ARGENTINA: Misiones: San Javier, 1 ,(MACN 33832).
Micrathena sanctispiritus Brignoli
Micrathena sanctispiritus Brignoli, 1983: 249 (new name for M. parallela
Mello-Leitão, 1940 preoccupied by Micrathena parallela O. Pickard-
Cambridge,1890) (male holotypefrom Colatina, Espírito Santo, Brazil,
M. Rosa, 1936–1937, in MNRJ, not examined; see World Spider
Catalog (2016) for complete taxonomic citation history).
Material examined
ARGENTINA: Misiones: San Pedro, Parque Provincial Cruce Caballero, 1 ,
(MACN 32552).
Micrathena shealsi Chickering
(Fig. 2D)
Micrathenashealsi Chickering, 1960:8,figs13–17 (female holotype from
Sunchal, Argentina, deposited in MCZ, not examined; see World
Spider Catalog (2016) for complete taxonomic citation history).
Material examined
PERU: Cusco: Paucartambo, Kosñipat, road to Manu, 1 ,(MUSM-ENT
510061).
Acknowledgements
We would like to thank C. Griswold and A. Carmichael (CAS), L. S. Carvalho
(CHNUFPI), A. D. Brescovit (IBSP), M. Oliveira (INPA), M. J. Ramírez and
C. Scioscia (MACN), G. Giribet and L. Leibensperger (MCZ), A. B. Kury
(MNRJ), A. B. Bonaldo (MPEG), D. Silva (MUSM-ENT), and R. Pinto-da-
Rocha and S. Outeda-Jorge (MZSP) for making specimens available. Many
thanks are due to A. Anker, F. U. Yamamoto and G. F. B. Pereira, who
collected specimens essential for this study and allowed publication of
their gorgeous photos of living specimens. We are very much indebted to
M. Servin-Pastor and F. Álvarez-Padilla for sorting and lending specimens
of M. spinulata collected within the research project funded by the Program
UNAM-DGAPA-PAPIIT code IN213612. L. S. Carvalho provided some
specimens, and he and L. M. Moras helped with some of the sequence data
acquisition. R. Ott provided information on some specimens deposited at
the Museo de Ciências Naturais spider collection in Porto Alegre, Brazil. The
permissionfor collecting atParque Estadual Sete Passagens has been issued by
SEMA-Bahia, and fieldwork there was kindly supported by the reserve staff.
AAN thanks PPGEco-INPA, IBAMA/ICMBio and PARNA Pico da Neblina
for collecting permits, FUNAI and the Ayrca, a local Yanomami association,
for their reception at the Yanomami Indigenous Land, and several colleagues
for their help with the fieldwork. Molecular work has been carried out in
the Laboratório de Malacologia e Sistemática Molecular and Laboratório de
Biodiversidade e Evolu¸cão Molecular at the Universidade Federal de Minas
Gerais. We thank the Willi Hennig Society for sponsoring and allowing free
use of TNT. Earlier versions of the manuscript were improved by critical
readings by R. Botero-Trujillo and by a thorough review by M. Harvey,
M. G. Rix and an anonymous referee. This study was supported by grants from
CNPq and CONICET to ILFM (372279/2013–9), from CNPq, BECA-IEB/
Moore Foundation (B/2007/01/BDP/01) and Wildlife Conservation Society
(WCS) to AAN, from Programa Pronoturno (FUMP/UFMG) to PHM and
from FAPEMIG (APQ-01991–09; PPM-00553–11, PPM-00335–13), CNPq
(474680/2010–0; 475179/2012–9, 407288/2013–9, 308072/2012–0),
FAPESP (2011/50689–0) and Instituto Nacional de Ciência e Tecnologia
dos Hymenoptera Parasitóides da Região Sudeste Brasileira (http:/www.
hympar.ufscar.br/) to AJS.
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