Content uploaded by Sarah C. Crews
Author content
All content in this area was uploaded by Sarah C. Crews on May 11, 2023
Content may be subject to copyright.
1
Zoological Journal of the Linnean Society, 2023, XX, 1–94. With 76 figures.
A revision of the genus Porteria and the phylogeny and
biogeography of Porteriinae (Araneae: Desidae)
ELIZABETHMORRILL1, SARAHCREWS1, LAURENESPOSITO1, , MARTÍN J.RAMÍREZ2
and CHARLESGRISWOLD1,*
1Department of Entomology, California Academy of Sciences, San Francisco, CA 94118, USA
2Division of Arachnology, Museo Argentino de Ciencias Naturales - CONICET, C1405DJR Buenos Aires,
Argentina
Received 5 January 2022; revised 11 May 2022; accepted for publication 14 October 2022
We revise the Chilean genus Porteria, including the type species, Porteria albopunctata, and 11 new species: Porteria
ajimayo sp. nov., Porteria alopobre sp. nov., Porteria ariasbohartae sp. nov., Porteria bunnyana sp. nov.,
Porteria contulmo sp. nov., Porteria correcaminos sp. nov., Porteria eddardstarki sp. nov., Porteria faberi
sp. nov., Porteria fiura sp. nov., Porteria misbianka sp. nov. and Porteria torobayo sp. nov. A phylogenetic
analysis using six genetic markers confirms the monophyly of Porteriinae, including Baiami and the core porteriines,
here defined to include the ecribellate genera Cambridgea, Corasoides, Nanocambridgea and Porteria. Core
porteriines are diagnosed by a narrowed section of the piriform gland spigot field, the cymbium extended to a narrow
tip and lackof a median apophysis. Porteria and Corasoides are sister taxa, united by the behaviour of running
atop the sheet of a web and by spinning a regular square mesh in the web platform. According to our results, the
diversification of Porteria started about 30 Mya (44–17 Mya). A biogeographic analysis infers that an ancestor of
Porteria reached South America via a founder event from Australia or New Zealand, where their close relatives occur.
ADDITIONAL KEYWORDS: historic biogeography – long-range dispersal – marronoid – RTA clade – spiders.
INTRODUCTION
Porteria Simon, 1904 is a Chilean genus of sheet web
building spiders that range from just north-west of
Santiago to the far south near Punta Arenas in southern
Patagonia, roughly coincident with the distribution
of the southern beech (Nothofagus Blume) forests in
South America. They belong in a group of marronoid
spiders (Wheeler et al., 2017: 2, fig. 5) referred to as the
‘Fused Paracribellar Clade’ by Griswold et al. (1999)
or the ‘Austral cribellates’ (Spagna & Gillespie, 2008).
These groupings were partially corroborated in subse-
quent molecular studies (Wheeler et al., 2017; Crews et
al., 2019) and their members are mainly endemics of
temperate forests in the Southern Hemisphere.
Porteria was first described by Simon (1904) in a study
of late 19th century arachnid collections from Chile. The
genus was placed in the family Agelenidae, subfamily
Cybaeinae, with Porteria albopunctata Simon, 1904
described as the sole species. Half a century later, two
major works revised Porteria. Roth (1967) reviewed
South American agelenids and Lehtinen (1967) pro-
duced a large-scale work on the classification of cribel-
late spiders. Roth (1967) maintained Simon’s family
and subfamily placement, although he mentioned that
distinctions between the subfamilies of what he con-
sidered Agelenidae may be imprecise. Lehtinen (1967)
described the composition of Agelenidae as a large
phenetic lumping: ‘In my opinion, the Agelenidae of
all past and even of recent authors is a mere assem-
blage of three-clawed Araneomorph spiders without
cribellum and without any radical modifications …’ (p.
341). He undertook a major overhaul of Agelenidae, re-
locating many of its previous genera to a diverse array
of families. He moved Porteria into Amaurobiidae, sub-
family Desinae, which now has familial status, and
Desidae is where Porteria now remains. The removal
from Agelenidae was partially based on the unpaired
colulus of Porteria, which Lehtinen showed is paired
in Agelenidae.
*Corresponding author. E-mail: cgriswold@calacademy.org
[Version of record, published online 4 May 2023; urn:lsid:zoo
bank.org:pub:76B9F689-4B90-433C-9837-92E49F1DDE80]
© The Author(s) 2023. Published by Oxford University Press on behalf of The Linnean Society of London.
All rights reserved. For permissions, please e-mail: journals.permissions@oup.com
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
2 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Although Lehtinen (1967) described the limits of
his subfamily Desinae as problematic (‘[t]he limi-
tation of this subfamily is far more difficult than
that of any other subfamily of my Amaurobiidae’: p.
324), novel morphological similarities and recent mo-
lecular work support some of his groupings in the
subfamily, specifically the association of Porteria with
Corasoides Butler, 1929 in his tribe Porteriini. Most
recently, the analysis of Wheeler et al. (2017), using
several genes, confirmed that Porteria is closely re-
lated to the Australian Corasoides, and provided sup-
port for their grouping with the New Zealand genera
Cambridgea L. Koch, 1872 and Nanocambridgea
Forster & Wilton, 1973 and the Australian Baiami
Lehtinen, 1967, erecting the subfamily Porteriinae
for all of them, in the family Desidae. As noted by
Wheeler et al. (2017), the molecular data are con-
gruent with a peculiarly narrowed extension of the
piriform spigot field on the anterior lateral spinnerets
in all porteriines except Baiami. Significant taxo-
nomic work has been done on all the Australasian
relatives of Porteria, with revisions of the genera
Cambridgea and Nanocambridgea (Forster & Wilton,
1973; Walker et al., 2020; Blest et al., 2000), Baiami
(Gray, 1981) and Corasoides (Humphrey, 2017). In this
contribution, we complete the taxonomic coverage of
the Porteriinae by revising Porteria, and we produce
a phylogenetic hypothesis of the subfamily, building
on the previous studies of Wheeler et al. (2017) and
Walker et al. (2020).
The four genera that we identify as core porteriines
are homogeneous in morphology and web architecture
(Porteria, Figs 1, 2; Corasoides, Fig. 3F, G; Cambridgea,
Fig. 3A-C; Nanocambridgea, Fig. 3D, E). We discuss po-
tential synapomorphies in their morphology as well as
their behaviour.
MATERIAL AND METHODS
All measurements are in millimetres unless other-
wise noted and were taken using a reticule in an
Olympus SZH10 stereomicroscope. Habitus lengths
were taken in dorsal view while leg measurements
were taken in lateral view along the dorsal margin.
Spination (macrosetae) patterns are given for dorsal
(d), prolateral (p), ventral (v) and retrolateral I sur-
faces and are listed proximally to distally for each
segment; see Griswold (1987: 6, 7) for a complete ex-
planation of this protocol. Measurements are given
for one specimen of each sex. Variations within a sex
or species are reported separately. Male palpal bulb
measurements are taken from the anterior to posterior
margins of the aveolus (these do not include any pro-
jection of the conductor or embolus). Spinneret morph-
ology is described using the terminology of Coddington
(1989) and Townley & Tillinghast (2003). Female geni-
talia, especially when describing the vulva, is based on
Sierwald’s (1989) terminology. The dictynoid pore as
described in Bennett (2006) is referred to in this paper
as Bennett’s gland (Ramírez, 2014).
The photographs used in this study were taken with
a Leica DFC500 digital camera mounted on a Leica
MZ16A stereomicroscope. Stacks of images were com-
bined using the montage software Helicon Focus.
Dorsal views of female genitalia were taken after soft
tissues were digested with the enzyme pancreatin
(Alvarez-Padilla & Hormiga, 2007). Darker struc-
tures of the vulva were cleared with clove oil or bleach
when necessary. Male palp expansion was carried out
by placing the palps in hot lactic acid until the bulbs
started to expand and shift in orientation and were
then placed in deionized water where further expan-
sion occurred.
Scanning electron micrographs (SEMs) were
taken on a LEO/Zeiss 1450VP SEM at the California
Academy of Sciences (CAS). Specimens were crit-
ical point dried and sputter coated with gold, then
mounted on copper wire with white glue or onto
copper tape. Some spinnerets for electron microscopy
needed to be ‘squeezed’ in order for all spinnerets to
be visible, especially the posterior median spinnerets.
We followed the protocol in Coddington (1989) using
forceps to squeeze the spinnerets out and using a
paper clip to hold the forceps closed in place over-
night while the abdomen hardened in 100% ethanol.
Some images and micrographs were produced for
the project Assembling the Tree of Life—Phylogeny
of Spiders, a multinational project supported by a
grant from the USA National Science Foundation
(NSF—EAR-0228699).
We use the unique identifier ‘CASENT###’
throughout, except where specimens already bear
unique identifiers from their home institutions,
without connoting ownership of specimens. The custo-
dial institution, e.g. ‘AMNH,’ is listed in the ‘Types’ and
‘Other material examined’ sections before the unique
identifier, which is listed in parentheses.
AbbreviAtions
Morphology
AC aciniform gland spigot; AER anterior eye row;
ALE anterior lateral eye; ALS anterior lateral spin-
neret; AME anterior median eye; AVW atrium ven-
tral wall; BG Bennet’s gland; BS1 basal receptacle 1;
BS2 basal receptacle 2; C conductor; CL colulus; CD
copulatory duct; CO copulatory opening; CY cylin-
drical gland spigot; DTA dorsal tibial apophysis; E
embolus; FD fertilization duct; HS spermathecal head;
LRTA long retrolateral apophysis positioned mesad to
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 3
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
RTA; MAP major ampullate gland spigot; MIP minor
ampullate gland spigot; Nu nubbin of gland spigot;
PC paracymbium; PER posterior eye row; PLE pos-
terior lateral eye; PI piriform gland spigot; PLS pos-
terior lateral spinneret; PME posterior median eye;
PMS posterior median spinneret; RTA retrolateral
tibial apophysis; Sc scape; SS spermathecal stalk; Tp
tartipore; VTA ventral tibial apophysis.
Institutions
CAS California Academy of Sciences, San Francisco,
California, USA; AMNH American Museum of
Natural History, New York, USA; FMNH Field
Museum of Natural History, Chicago; FMHD Field
Museum of Natural History, Chicago (Bulk Sample
Collection), USA; MACN-Ar Museo Argentino
de Ciencias Naturales, Buenos Aires, Argentina;
Figure 1. Porteria specimens alive. A, B, P. eddardstarki sp. nov., female from Palmas de Ocoa (MACN-Ar 39150). C, D, P.
eddardstarki sp. nov., male from Palmas de Ocoa (MACN-Ar 39122). E, Porteria sp. (probably P. ajimayo sp. nov.), female
from Pata de Gallina, south of Contulmo (14.II.1992, Platnick, Goloboff, Ramírez, AMNH; photo 791).
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
4 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
MCZ Museum of Comparative Zoology, Harvard
University, Massachusetts, USA; MNHN Muséum
National d’Histoire Naturelle, Paris, France; MHNS
Museo Nacional de Historia Natural, Santiago,
Chile.
Phylogenetic AnAlysis
We obtained sequences of 35 samples from ten of the 12
proposed species of Porteria, plus three immature, un-
identified individuals that were of geographic interest.
Two species, P. alopobre sp. nov. and P. torobayo sp.
Figure 2. Webs of Porteria. A, P. fiura sp. nov., Chiloé. B, Porteria from Volcán Osorno (P. correcaminos sp. nov. or P.
bunnyana sp. nov.). C, P. ajimayo sp. nov. from Parque Pedro del Río Zañartu (Concepción). D, P. eddardstarki sp. nov. from
Zapallar showing glittery appearance of web in sunlight. E, two webs of P. eddardstarki sp. nov. from Quebrada El Tigre,
Zapallar (photo by Jaime Pizarro-Araya). F, web of P. ajimayo sp. nov. MACN-Ar 21074 from Contulmo, inset showing
platform with tangle above.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 5
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 3. A, Cambridgea sp. from Kahurangi National Park, South Island, New Zealand (photo by Peter Michalik). B,
Cambridgea sp. from Mulford Sound, South Island, New Zealand. C, Cambridgea sp. from Punakaiki, South Island, New
Zealand. D, E, Nanocambridgea sp. from Pelorus Bridge, South Island, New Zealand. F, G, Corasoides terania Humphrey,
2017 from Border Ranges National Park, NSW, Australia (photos by Jonas Wolff).
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
6 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
nov., were not included because of the rarity of the spe-
cimens and a lack of proper preservation conducive to
molecular data collection. However, we succeeded in
obtaining fair quality sequences of P. albopunctata,
P. contulmo sp. nov. and P. misbianka sp. nov. for at
least one gene, even though all were preserved in
75% ethanol at room temperature for over a decade.
To these, we added 15 samples of 12 outgroup species
chosen after previous phylogenetic work by Wheeler
et al. (2017) and Kallal et al. (2021). The outgroups
included all genera of Porteriinae, plus representa-
tives of other subfamilies of Desidae: Amphinecta
Simon, 1898 (Amphinectinae), Badumna Thorell,
1890 (Matachiinae), Desis Walckenaer, 1837 (Desinae)
and Metaltella Mello-Leitão, 1931 (Metaltellinae).
The tree was rooted between Desidae and Stiphidion
(Stiphidiidae) based on the phylogenetic arrangement
proposed by Wheeler et al. (2017: fig. 5). The complete
dataset had 50 species; a complete list of vouchers and
sequence identifiers can be found in Table 1.
Sequence data were obtained from the Center for
Comparative Genomics at the California Academy of
Sciences. DNA was extracted from the right four legs
of the specimens using the Qiagen DNeasy Blood and
Tissue Kit. We targeted five genes: the nuclear ribo-
somal genes 18S and 28S, and the coding genes histone
H3 and Actin, in addition to the coding mitochondrial
gene cytochrome c oxidase subunit I (COI). We also
included the mitochondrial ribosomal gene 16S for
outgroup taxa. Details of primers and PCR protocols
are given in the Supporting Information (Appendix
S1).
Raw sequence data were viewed, edited and as-
sembled with SEQUENCHER v.5.4.6 (Gene Codes
Corporation). Alignment of the protein coding genes
was done in the online version of MAFFT v.7 (Katoh &
Standley, 2013) using default parameters. The rDNA
genes 28S, 18S and 16S were aligned according to their
secondary structural conformation using sequences
from previous work as references (Spagna et al., 2010;
Crews et al., 2019). Protein coding genes were exam-
ined in MESQUITE (Maddison & Maddison, 2011) to
ensure that the sequences were free of stop codons.
Gaps were treated as missing data.
The data were concatenated using SequenceMAtrix
(Vaidya et al., 2011) with the following numbers of
characters per gene used in the analyses: 28S = 745;
18S = 1772; Actin = 383; H3 = 347; COI = 665. The data
were partitioned by gene, by codon (Goldman & Yang,
1994; Muse & Gaut, 1994), and by stems and loops for
the rDNA genes (Nylander et al., 2004; Brandley et al.,
2005). The best-fit partitioning schemes and models
were obtained using ModelFinder (Kalyaanamoorthy
et al., 2017) in IQ-Tree v.2.1.1 (Nguyen et al.,
2015; Minh et al., 2020) with the corrected Akaike
Information Criterion (AICc), and the results can be
found in the Supporting Information (Appendix S2).
We conducted two ModelFinder analyses, one for the
IQ-Tree analysis and one for the analysis in MrBAyes
v.3.2.7 (Huelsenbeck & Ronquist, 2001; Ronquist &
Huelsenbeck, 2003) to accommodate that some models
cannot be used in both programmes. All analyses were
conducted on the CIPRES Science Gateway (Miller et
al., 2010).
We conducted both fast maximum likelihood and
Bayesian analyses, the former with IQ-Tree (Nguyen
et al., 2015) and the latter with MrBAyes (Ronquist
& Huelsenbeck, 2003). IQ-Tree was run using 10
000 replicates of ultrafast bootstrapping with the
-bnni parameter engaged to reduce model violations
(Hoang et al., 2018) with partitioning (Chernomor et
al., 2016). First, we conducted three preliminary ana-
lyses varying how substitution models are handled
across partitions (-q, all with same branch lengths; -p,
each partition has the same branch length but its own
evolutionary rate; -Q, each partition with its own set
of branch lengths). We used the results with the best
AICc score as the favoured analysis, and then re-ran
the analysis nine additional times with the favoured
parameters to ensure the search was not getting stuck
on local optima.
The doublet model of nucleotide substitution
(Schöniger & von Haeseler, 1994) was used in MrBAyes
for the stem partitions of the rDNA genes because
this has been shown to improve resolution in com-
bined data analysis (Erpenbeck et al., 2007), including
studies of other marronoid spiders (Spagna et al., 2010;
Crews et al., 2019). MrBAyes was run for 50 million
generations, sampling every 1000th tree using default
parameters. Stationarity was checked using TRACER
(Rambaut et al., 2014). A maximum clade credibility
tree was made using SumTrees (Sukumaran & Holder,
2010, 2015).
Dated phylogeny
We produced a time-calibrated tree with Beast
(BEAST2 on XSEDE). In absence of fossils closely re-
lated to our focal taxa, we used two secondary calibra-
tions taken from the analysis of Magalhaes et al. (2020)
(Fig. 74). One is the divergence between Stiphidiidae
and Desidae [median 73.30 Mya, 95% highest pos-
terior density (HPD) 101.34–48.32 Mya], the second
is the divergence of Desidae (median 56.37 Mya, 95%
HPD 85.01–35.03 Mya). These were implemented as
uniform priors encompassing the 95% HPD, using an
uncorrelated lognormal relaxed molecular clock with
the same partitions used in the MrBAyes analyses. To
simplify the calculations and obtain convergence, we
used a simpler Hasegawa, Kishino and Yano (HKY)
model for all partitions, with a Yule tree prior, uni-
form birth rate, and ucld mean and Gamma ucld with
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 7
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Table 1. Species, voucher numbers and GenBank accessions for spiders used in the molecular analyses. New sequences start with ‘ON’. (-) indicates that there
are no sequence data available for the gene
Species Voucher ID 28S 18S Actin H3 COI 16S Sex/
stage
Locality
Amphinecta luta ARACG000020,
CASENT9019847
KY017023 KY016398 - KY018182 KY017671 KY015820 Female South Island, New
Zealand
Badumna longinqua ARACG000013 FJ607523 KY016401 - KY018185 KY017674 KY015823 Male California, USA
Baiami sp. CG291 ARACG000291,
CASENT9023594
KY017027 KY016403 - - KY017676 KY015825 Female Western Australia
Cambridgea sp. CASENT9055737 ON342792 ON342754 ON352008 ON352047 ON337177 - Juvenile South Island, New
Zealand
Cambridgea reinga ARACG000097 KY017030 KY016406 - KY018188 - KY015827 North Island, New
Zealand
Corasoides sp. CASENT9055506 ON342777 ON342752 ON352009 ON352044 ON337176 Female Queensland, Australia
Corasoides sp. CG294 ARACG000294,
CASENT9023677
KY017033 KY016409 - KY018191 KY017682 KY015830 Juvenile Queensland, Australia
Desis formidabilis ARACG000274,
CASENT9023641
- KY016410 - KY018192 KY017683 KY015831 Male Western Cape, South
Africa
Desis marina DesisNZ HM576644 HM576628 - HM576663 - - New Zealand
Metaltella sp. CG23 ARACG000023,
CASENT9016486
KY017040 KY016416 - KY018198 KY017689 - Female Villarrica, Chile
Metaltella sp. CG60 ARACG000060,
CASENT9020329
KY017042 KY016418 - KY018200 - KY015836 Juvenile Monte Hermoso,
Argentina
Nanocambridgea sp. CASENT9055701 ON342791 ON342753 ON352007 ON352046 ON337174 - Male South Island, New
Zealand
Nanocambridgea sp.
CG203
ARACG000203, CASENT
9021498
KY017045 KY016421 - KY018203. KY017692 KY015837 South Island, New
Zealand
Stiphidion facetum CASENT9023563 ON342767 ON342755 ON352010 ON352045 ON337175 - Male Tasmania, Australia
Stiphidion facetum ARASP000081 KY017385 KY016726 - KY018447 KY017943 KY016135 Male Queensland, Australia
Porteria ajimayo CASENT9055614 ON342783 ON342726 ON352002 ON352035 ON337167 - Female Nahuelbuta, Chile
Porteria ajimayo CASENT9053793-1 ON342771 ON342724 ON352005 ON352021 ON337154 - Juvenile Parque Pedro del Río
Zañartu, Concepción,
Chile
Porteria ajimayo CASENT9053793-2 ON342772 ON342747 ON352003 ON352034 ON337155 - Male Parque Pedro del Río
Zañartu, Concepción,
Chile
Porteria albopunctata CASENT9044702 ON342770 - - - - - Female Laguna San Rafael,
Chile
Porteria ariasbohartae CASENT9026272-f ON342759 ON342734 ON351982 ON352016 ON337145 - Female Oncol Park, Chile
Porteria ariasbohartae CASENT9026272-m ON342768 ON342746 ON351984 ON352017 ON337146 - Male Oncol Park, Chile
Porteria bunnyana CASENT9055665-f ON342786 ON342732 - ON352033 ON337163 - Female Pucón, Chile
Porteria bunnyana CASENT9055695-m ON342787 ON342738 ON351989 ON352027 ON337163 - Male Pucón, Chile
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
8 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Species Voucher ID 28S 18S Actin H3 COI 16S Sex/
stage
Locality
Porteria contulmo MACN-Ar21077 - - - - ON337144 - Male Contulmo, Chile
Porteria contulmo ARAMR000284 KY017050 KY016426 - - KY017696 KY015841 Contulmo, Chile
Porteria correcaminos CASENT9035455 ON342769 ON342733 ON352006 ON352049 ON337147 - Male Puyehue, Chile
Porteria correcaminos CASENT9053812-1 ON342773 ON342727 ON351985 ON352025 ON337148 - Female Lago Chapo, Chile
Porteria correcaminos CASENT9053812-2 ON342774 ON342748 ON351987 ON352022 ON337178 - Female Lago Chapo, Chile
Porteria correcaminos CASENT9053857-1 ON342775 ON342749 ON351991 ON352018 ON337150 - Female Volcán Osorno, Chile
Porteria correcaminos CASENT9053857-2 ON342776 ON342728 ON351988 ON352023 ON337156 - Female Volcán Osorno, Chile
Porteria correcaminos CASENT9055694-1 ON342789 ON342756 ON352013 ON352026 ON337149 - Female Petrohué, Chile
Porteria correcaminos CASENT9055694-2 - ON342725 ON352012 - - - Juvenile Petrohué, Chile
Porteria eddardstarki MACN-Ar27062 ON342760 ON342743 ON351994 ON352039 ON337169 - Female Zapallar, Chile
Porteria eddardstarki MACN-Ar27063 ON342761 ON342744 ON351997 ON352042 ON337171 - Female Zapallar, Chile
Porteria eddardstarki CASENT9055516 ON342780 ON342741 ON351995 ON352040 ON337170 - Female Zapallar, Chile
Porteria eddardstarki CASENT9055690 ON342788 ON342745 ON351996 ON352043 ON337173 - Female Olmué, Chile
Porteria eddardstarki CASENT9055699 ON342790 ON342742 ON351998 ON352041 ON337172 - Male Zapallar, Chile
Porteria eddardstarki CASENT9055512-1 ON342778 ON342757 ON352014 ON352019 ON337157 - Female Cuesta La Dormida,
Chile
Porteria eddardstarki CASENT9055512-2 ON342779 ON342758 ON352015 ON352020 ON337158 - Female Cuesta La Dormida,
Chile
Porteria faberi 35369 ON342763 ON342736 ON351986 ON352028 ON337151 - Juvenile Reserva Costera
Valdiviana, Chile
Porteria faberi 35371 ON342764 ON342737 ON351983 ON352029 ON337153 - Juvenile Reserva Costera
Valdiviana, Chile
Porteria faberi 35399 ON342766 ON342735 ON352004 ON352031 ON337162 - Female Reserva Costera
Valdiviana, Chile
Porteria faberi CASENT9053791 ON342793 ON342729 ON351992 ON352032 ON337152 - Female Reserva Costera
Valdiviana, Chile
Porteria fiura 35305 ON342762 ON342740 ON351999 ON352037 - - Juvenile Chiloé Island, Chile
Porteria fiura CASENT9055611 ON342781 ON342750 ON352001 ON352048 ON337159 - Female Chiloé Island, Chile
Porteria fiura CASENT9055611-B ON342782 ON342739 ON352000 ON352038 ON337160 - Juvenile Chiloé Island, Chile
Porteria misbianka CASENT9044700 - - - - ON337168 - Female Estero Nonguen,
Concepción, Chile
Porteria sp. 35372 ON342765 ON342730 ON351990 ON352030 ON337165 - Juvenile Reserva Costera
Valdiviana, Chile
Porteria sp. CASENT9055663-1 ON342784 ON342731 ON351993 ON352036 ON337161 - Juvenile Nahuelbuta, Chile
Porteria sp. CASENT9055663-2 ON342785 ON342751 ON352011 ON352024 ON337166 - Juvenile Nahuelbuta, Chile
Table 1. Continued
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 9
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
uniform calibrations. Stationarity was reached and no
effective sample size (ESS) < 200.
Biogeographic analysis
With the time-calibrated tree from the BEAST2 ana-
lysis, we used the R package BioGeoBears v.1.1.1
(Matzke, 2013, 2014) to infer ancestral areas. As re-
commended by the developer, the tree was pruned
to include a single representative per species. The
scored areas represent the main landmasses in-
volved in the study: South America (SA), South Africa
(Af), Australia (Au), New Zealand (NZ); Supporting
Information, Table S1). We also considered Antarctica,
a candidate for habitable area in the period of interest,
following the biogeographical model of Estrella et al.
(2019). We added an additional time slice after the
Mid-Pliocene warm period, which represented an in-
habitable Antarctica, following Haywood et al. (2016).
We used three different dispersal multipliers as in
Estrella et al. (2019) representing the connections and
distances of landmasses in the four time slices: 1.0 for
areas directly connected, 0.5 for areas not directly con-
nected but separated by a smaller barrier, and 0.01 for
areas separated by a large oceanic barrier (Supporting
Information, Table S2).
We tested the six biogeographic models imple-
mented in BioGeoBears (DEC, BAYAREALIKE and
DIVALIKE, and their +J variants) and used the AICc
to select the best fitting model. In all cases, we con-
sidered a maximum of three areas occupied by any spe-
cies. Because any dated tree contains considerable age
uncertainty, and groups with low support may not be
monophyletic, the selected model was used for a subse-
quent analysis incorporating the age uncertainty and
topological variants represented in 100 trees randomly
chosen from the stationary Bayesian distribution of
the BEAST2 output. The analyses were replicated on
these 100 trees, calculating ten stochastic mappings
for each tree; the scripts developed by Magalhaes et
al. (2021) were used to automate and summarize these
procedures. Source scripts and input data can be found
in the Supporting Information (Files S1–S3).
comPArAtive mAteriAl exAmined
Baiami brockmani Gray, 1981. Australia: Western
Australia: Big Tingle Tree, Walpole-Nornalup
National Park, 5.29 km 104° ESE Walpole, S34.98329°
E116.78776°, C. Griswold, D. Silva, L.J. Boutin, G.
Hormiga, N. Scharff, February 25–26, 2006, elev.
90 m, general collecting in eucalypt forest, female
CASENT9022915; Crowea, S34º28, E116º10’, S.J.
Curry, gift from WAM, 19 November 1976, pitfall
traps in open (cleared) forest, male CASENT9024809;
Crowea, Creek Site, S34º28’, E116º10’, S.J. Curry, gift
from WAM, 12 December 1977, pitfall traps in open
(cleared) forest, female CASENT9024810. SEM im-
ages taken by Fernando Alvarez-Padilla.
Cambridgea foliata (L. Koch, 1872). New Zealand:
North Island: Northland, Waipoua Forest Trust,
5.73 km ESE Waipoua, S35.69672° E173.58347°, C.
Griswold, D. Silva and H. Wood, 9 February 2005,
elev. 365 m, secondary forest with closed canopy, fe-
males CASENT902130, CASENT9021305, male
CASENT9021304, immature CASENT9023885. SEM
images taken by Meghan Culpepper, Joel Ledford,
Hannah Wood and Nibia Soto Rolón.
Corasoides terania Humphrey, 2017. Australia:
Queensland, O’Reilly’s, Lamington National Park,
S28°13’ E153°08’, C. Griswold, 18–21 July 1992, elev.
935 m, montane forest, immature CASENT9023883;
Main track, O’Reilly’s, J. Gallon, 2 November 1989,
night collecting, male and female QMBS73356.
Binna Burra, Lamington National Park, S28.19398°
E153.18692°, C. Griswold, D. Silva, R. Raven, B. Baehr,
M. Ramírez, 21–23 March 2006, elev. 790 m, rainforest,
general collecting, female CASENT9024561.
Nanocambridgea sp. New Zealand: South Is.,
Marlborough region: Queen Charlotte Track, 14.9 km
057º E Havelock, S41.25230º, E173.94077º, C. Griswold,
D. Silva and H. Wood, 26 February 2005, elev. 0–15 m,
male and female CASENT9023876; Pelorus Bridge
Scenic Res., near Kahikatea Campground, 16.6 km 239º
W Havelock, S41.30131º, E173.57069º, C. Griswold, D.
Silva and H. Wood, 27 February 2005, elev. 40 m, night
collecting, male and female CASENT9023877. SEM
images taken by Joel Ledford and Christopher Vo.
TAXONOMY
PorteriinAe lehtinen, 1967
Porteriini Lehtinen, 1967: 326 (type genus Porteria
Simon, 1904).
Porteriinae Wheeler et al., 2017: 27, 34.
Included genera: Baiami Lehtinen, 1967, Cambridgea
L. Koch, Corasoides Butler, 1929, Nanocambridgea
Forster & Wilton, 1973 and Porteria Simon, 1904.
Diagnosis: The core group of ecribellate porteriines
(Cambridgea, Corasoides, Nanocambridgea, Porteria)
are easily distinguished from other Desidae by having
a narrowed section of the piriform gland spigot field
(Figs 5B, 6B, 7B, D, 8B, 9B, 10B), and the male palpal
cymbium is extended to a narrow tip about 1.5 to 9
times longer than the copulatory bulb and by the
absence of a median apophysis (Figs 12C, F, 26B, 46B).
Baiami are cribellate, the male palp has a moderately
extended cymbium, also lacks a median apophysis, and
has a retrolateral tibial process composed of a basal
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
10 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
lamina surmounted by a curved, spiniform process and
has the piriform gland spigots spread evenly across
the ALS spinning surface, not forming a narrowed
configuration (Fig. 11B, F, G) (see Gray, 1981).
Morphological remarks: The four genera that we
identify as core porteriines are homogeneous in
spinning organ morphology (Figs 5–11). All are
ecribellate, and the colulus is unilobate. The ALS of
the female have two major ampullate gland spigots,
and in the male, one plus a nubbin. The piriform field
extends as a narrow stripe or ‘tail’, with the spigots
gradually smaller as the extension lengthens toward
the posteromedian margin. Porteria has a row of 5 or
6 stout setae just posterior to the spinning field (Figs
5B, 7B). Corasoides and Cambridgea have a brush of
setae in the same place, but they are not so stout nor
aligned as in Porteria (Figs 8B, 9B). Most of the species
examined with SEM have two cylindrical gland spigots
on the PMS and one on the PLS, except for Cambridgea
foliata, with four and three, respectively, probably
related to its larger size. The remaining spigots on the
posterior spinnerets are all similar in shape and size;
these may be all aciniforms, or, alternatively, some are
minor ampullate spigots that are not morphologically
distinct. Nanocambridgea has many more aciniform
spigots than the other species, even though it has the
smallest body size compared to other porteriines (some
Nanocambridgea are as small as 6mm body length,
whereas some Cambridgea and Corasoides may have
body lengths in excess of 12mm). The affiliation of
Baiami to the other porteriines is supported mainly
by the molecular data. Their spinning organs (Fig.
11) show a well-developed, bipartite cribellum,
with strobilate spigots. The ALS have a normal,
approximately circular field of piriform gland spigots,
two major ampullate gland spigots in the female, and
one plus a nubbin in the male. The female PMS have a
single minor ampullate spigot in the anterior position,
eight paracribellar spigots, two cylindrical gland
spigots and five aciniform gland spigots. The male
PMS have anterior minor ampullate and aciniform
gland spigots, but the paracribellars are reduced to
nubbins. The PLS of our exemplars in both sexes are
collapsed; the female has at least one cylindrical gland
spigot, and both sexes have remnants or nubbins of a
triplet, compatible with the common triplet pattern of
a central modified spigot and two paracribellar gland
spigots. The coloration of Porteria and Corasoides is
similar, with white guanine bands laterally on the
opisthosoma (Figs 1B, 13, 37, 60), although the white
bands are less conspicuous in Porteria, and virtually
absent in P. eddardstarki sp. nov.
Behavioural notes: Species of Cambridgea build
extensive sheet webs with a vast snare of knock-down
lines on top and guy lines below the sheet, and the
spiders move on the underside of the sheet (Fig. 3A,
B; Walker et al., 2020). Species of Nanocambridgea
build densely woven sheet webs and also walk below
the sheet. These webs were found in depressions along
tree trunks (Fig. 3D), as well as banks and among tree
roots (J. Wolff, pers. comm.). Species of Corasoides
make a sheet web with a tangle above, guy lines below,
and a depressed retreat, and they walk on top of the
sheet (Figs 3F, 4E-H; see Humphrey, 2021). Their web
platform has layers of parallel threads in a regular
structure, which was described by Butler (1929: 45):
‘To the naked eye it is similar to muslin, having
a definite weft and warp at right angles to each
other; under magnification it becomes irregular,
owing to an overlapping of its layers. One of these
layers isolated exhibits a true square mesh, of a
width of 0.8mm.’
Porteria webs (Fig. 2) are similar to those of
Corasoides, with a regular structure in the platform
(Fig. 4A-D). Most of the crossings of threads in the
square mesh have piriform attachments (Fig. 4C, D, G,
H). Individuals of both Porteria and Corasoides walk
on top of their sheets. Baiami makes cribellate sheet
webs, and the spiders walk below the sheet (Gray,
1981). Of the other representatives in our phylogenetic
analysis, all walk below their webs (Stiphidion Simon,
1902, Badumna and Metaltella) or do not construct
webs (Amphinectaand Desis).
desidAe Pocock, 1895
Porteria simon, 1904
Porteria Simon, 1904: 109 (type species Porteria
albopunctata Simon, 1904, by monotypy).
Diagnosis: Porteria species resemble their relative
porteriines Nanocambridgea, Cambridgea and
Corasoides in the remarkable narrowing of the ALS
piriform gland spigot field (Fig. 5), the cymbium
highly elongated beyond the bulb and the lack of a
median apophysis, but it differs from all of these by
the presence of a thin tibial apophysis (LRTA) arising
near the base of the RTA on the male palpal tibia (Fig.
68), and a line of five to six stout setae just posterior
to the ALS spinning field of both sexes (Fig. 5B) .
Among the Chilean fauna, Porteria species are the
only marronoids with the narrow tip of the cymbium
extending 1.5 to 5 times the length of the copulatory
bulb. The epigyne has a minute to large anterior
median scape (Figs 30C, 53A), and the spermatheca
is bilobed, here referred to as base 1 and base 2 of the
spermatheca. All species except P. eddardstarki have
distinctive dorsal abdominal markings consisting of
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 11
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
white to yellow anterolateral lines and median spots
(Fig. 13). Porteria are like Corasoides but differ from
Nanocambridgea and Cambridgea by walking on the
upper surface of the sheet web instead of hanging
from it.
Description: Small to medium sized spiders with
total length 3.9–9.1. Characteristic markings as
follows (Figs 13, 14): carapace pale yellow to orange-
brown with darkened lateral margins with three
distinct, sometimes diffuse grey lines radiating
outward from thoracic furrow; grey lines outline
cephalic region along cervical groove. Black pigment
surrounding each eye, connecting the lateral eyes and
AMEs. Sternum with dark grey margins enclosing a
pale yellow median region. Paler region varies in size
and shape from large oval to small sliver: this contrast
is faded in older specimens. Endites, labium and
Figure 4. Webs of Porteriinae. A, Porteria ajimayo sp. nov., detail of web platform with cornstarch, showing square mesh.
B, Porteria eddardstarki sp. nov., detail of web platform. C, close-up of (B). D, same, close-up of (C), showing that most
intersections in the square mesh are affixed by silk attachments, except one at the centre. E, Corasoides sp. from Perth,
Western Australia (photo by Kerry Stuart). F, same, detail of platform. G, close-up of a platform area showing squared mesh,
as marked in (F). H, same, another area, note the specks at intersections, probably showing silk attachments. I, Cambridgea
sp. from Milford Sound, New Zealand, detail of platform, showing irregular mesh. J, Nanocambridgea sp. from Pelorus
Bridge, New Zealand, detail of densely woven platform.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
12 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 5. Scanning electron micrographs of spinnerets of female P. faberi sp. nov. from Reserva Costera Valdiviana CASENT
9055712. A, full view. B, ALS showing modified spinning field; arrow to row of modified setae. C, PMS. D, PLS. E, close-up
of major ampullate gland spigots and tartipore on ALS. F, close-up of cylindrical gland spigots on PMS. Abbreviations: ALS,
anterior lateral spinneret; CY, cylindrical gland spigot; MAP, major ampullate gland spigot; Nu, nubbin of gland spigot; PI,
piriform gland spigot; PLS, posterior lateral spinneret; PMS, posterior median spinneret; Tp, tartipore.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 13
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
chelicerae orange-brown in colour; endites and labium
lighter in colour toward tips. Coxae pale yellow with
grey distal margins. Legs with alternating yellow and
grey rings that vary from distinct to subtle, more
obvious in fresh specimens. Dorsum of abdomen dark
grey with white to golden yellow, longitudinal
anterolateral lines converging anteriorly; lines half
the length of abdomen, becoming indistinct spots
posteriorly. Fainter paired median spots sometimes
present. Porteria eddardstarki with anterolateral
lines reduced and dark grey dorsum mottled with
golden yellow (see species descriptions). Venter yellow
to grey, with median grey rectangle and two dark
grey semi-circles anterior to spinnerets. Epigyne
outlined with grey trapezoid with two yellow spots;
trapezoid also present in males between book lungs.
Spinnerets orange-brown to grey. Carapace length
1.20–1.51 times carapace width, height 0.23–0.61
times width. Thoracic fovea 0.07–0.31 times carapace
length, slit like and moderately deep. Posterior eye
row (PER) straight when viewed from above; anterior
eye row (AER) slightly procurved from anterior view
(Fig. 15B). Anterior lateral eyes (ALE) diameter 1.29–
2.00 times anterior median eyes (AME). Clypeus
height 1.67–2.83 times diameter of AME. Posterior
median eyes (PME) about equal to posterior lateral
eyes (PLE) diameter. Secondary eyes with canoe-
shaped tapetum (Fig. 15B). Sternum length 0.94–1.46
times width; labium as long as wide. Chelicera length
5.20–16.5 times clypeus height. Chelicera vertical in
most species, porrect in P. eddardstarki males (Fig.
31). Cheliceral fang margin with escort setae, rake
setae and whisker setae (see Ramírez, 2014).
Retromargin of chelicera with two teeth separated
far apart; promargin typically with five teeth and one
to three small denticles (Fig. 16C); see variations in
species descriptions. Fangs constricted or not;
constriction highly exaggerated in P. albopunctata
Figure 6. Scanning electron micrographs of spinnerets of male P. faberi sp. nov. from Reserva Costera Valdiviana CASENT
9053816. A, full view. B, ALS showing modified spinning field. C, PMS. D, PLS. E, close-up of ALS showing major ampullate
field. Abbreviations: ALS, anterior lateral spinneret; MAP, major ampullate gland spigot; Nu, nubbin of gland spigot; PI,
piriform gland spigot; PLS, posterior lateral spinneret; PMS, posterior median spinneret; TP, tartipore.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
14 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 7. Scanning electron micrographs of spinnerets of female P. correcaminos sp. nov. A, full view. B, ALS, showing
modified spinning field. C, piriform gland spigots in ALS, showing spigots of different sizes. D, ALS spinning field. E, PMS.
F, PLS. (A-E) from Panguipulli, CASENT 9021323, (F) from Volcán Osorno, CASENT 9055582. Abbreviations: ALS, anterior
lateral spinneret; CL, colulus; CY, cylindrical gland spigot; MAP, major ampullate gland spigot; Nu, nubbin of gland spigot;
PI, piriform gland spigot; PLS, posterior lateral spinneret; PMS, posterior median spinneret; Tp, tartipore.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 15
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 8. Scanning electron micrographs of spinnerets of Corasoides terania Humphrey, 2017 from Lamington. A, female
spinnerets, full view. B, female right ALS, inset to major ampullate field. C, female left PMS. D, female right PLS. E,
male, full view. F, male right ALS. G, male right PMS. H, male right PLS. Female CASENT 9024651, male QMBS73356.
Abbreviations: ALS, anterior lateral spinneret; CY, cylindrical gland spigot; MAP, major ampullate gland spigot; Nu, nubbin
of gland spigot; PI, piriform gland spigot; PLS, posterior lateral spinneret; PMS, posterior median spinneret; Tp, tartipore.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
16 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 9. Scanning electron micrographs of spinnerets of Cambridgea foliata (L. Koch, 1872). A, female, full view. B, female
left ALS. C, female left PMS. D, female right PLS. E, male, full view. F, male left ALS. G, male right PMS. H, male right
PLS. Female CASENT 9021303, male CASENT 9021304. Abbreviations: ALS, anterior lateral spinneret; CL, colulus; CY,
cylindrical gland spigot; MAP, major ampullate gland spigot; Nu, nubbin of gland spigot; PI, piriform gland spigot; PLS,
posterior lateral spinneret; PMS, posterior median spinneret; Tp, tartipore.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 17
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 10. Scanning electron micrographs of spinnerets of Nanocambridgea sp. from Pelorus Bridge. A, female, full view.
B, female left ALS. C, female PMS. D, female right PLS. E, male, full view. F, male left ALS. G, male PMS. H, male right
PLS. Male and female CASENT 9023877. Abbreviations: ALS, anterior lateral spinneret; CY, cylindrical gland spigot; MAP,
major ampullate gland spigot; Nu, nubbin of gland spigot; PI, piriform gland spigot; PLS, posterior lateral spinneret; PMS,
posterior median spinneret; Tp, tartipore.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
18 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 11. Scanning electron micrographs of spinnerets of Baiami brockmani Gray, 1981. A, female, full view, inset to
cribellar spigots. B, female left ALS, inset to major ampullate gland spigots. C, female right PMS. D, female right PMS, detail.
E, female right PLS, arrows to apical nubbins. F, male spinnerets. G, male right ALS. H, male left PMS. I, male left PLS,
arrows to apical nubbins. Female CASENT 9024810, male CASENT 9024809. Abbreviations: AC, aciniform gland spigot;
ALS, anterior lateral spinneret; CY, cylindrical gland spigot; MAP, major ampullate gland spigot; MIP, minor ampullate
gland spigot; Nu, nubbin of gland spigot; PI, piriform gland spigot; PLS, posterior lateral spinneret; PMS, posterior median
spinneret.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 19
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 12. Scanning electron micrographs of male palps. A, Cambridgea foliata (L. Koch, 1872), CASENT 9021304, right
palp, retrolateral view. B, same, tibia, ventral view. C, same, copulatory bulb, ventral view. D, Nanocambridgea sp. CASENT
9023877 from Pelorus Bridge, right palp, ventral-retrolateral view. E, same, cymbium and copulatory bulb. F, same, tip of
tibia and copulatory bulb. G, Corasoides terania Humphrey, 2017, CASENT 9024561 from Lamington, right palp ventral-
retrolateral view.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
20 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 13. Dorsal habitus of Porteria females (A-E) and males (F-H). Scale bar = 1mm. A, P. albopunctata Simon, 1904
from Parque Nacional Laguna San Rafael National Park, CASENT9044707. B, H, P. eddardstarki sp. nov. from Viña del
Mar, CASENT9044709. C, P. faberi sp. nov. from Reserva Costera Valdiviana, CASENT9055712. D, P. ajimayo sp. nov. from
Parque Pedro del Río Zañartu, CASENT9053784. E, P. bunnyana sp. nov. from Nahuelbuta National Park, CASENT9055706.
F, P. ajimayo sp. nov. from Monumento Natural Contulmo, MACNAr21081. G, P. ariasbohartae sp. nov. from Parque Oncol,
CASENT9026272. Scale bars = 1mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 21
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 14. Ventral habitus of Porteria females (A-E) and males (F-I). Scale bar = 1mm. A, P. eddardstarki sp. nov. from
Viña del Mar CASENT9044709. B, P. correcaminos sp. nov. from Lago Chapo, CASENT9053925. C, P. correcaminos sp. nov.
from Llanquihue, FMNH98_7. D, P. albopunctata sp. nov. from Laguna San Rafael National Park CASENT9044707. E,
P.fiura sp. nov. from Chiloé, CASENT9055645. F, P. bunnyana sp. nov. from Pucón, CASENT9044667. G, P. contulmo sp. nov.
from Contulmo NM, MACNAr21077. H, P. eddardstarki sp. nov. from Valparaíso, CASENT9044690. I, P. fiura sp. nov. from
Chiloé, CASENT9044672. Scale bars = 1mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
22 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
males (Fig. 23D). Male femur I 1.23–1.63 times
carapace length; female femur I 1.03–1.34 times
carapace length. Leg formula typically 4123,
sometimes 1423; leg I and IV about equal in total
length. General spination as follows (based on P.
eddardstarki, Figs 18–21; see spine map Fig. 22):
Male: palp: femur d1-1-2, patella d1-1, tarsus r0-0-1,
v0-0-1; leg I: femur d1-1-1(p)-1(p)-2, patella d1-1,
tibia d1(r)-0-0, v0-2-2, metatarsus v2-2-3; leg II:
femur d1-3-2-2, patella d1-1, tibia d1(r)-0-0, v0-0-2,
metatarsus p0-1-0-1, r0-0-1, v1(r)-2-2; leg III: femur
d1-3-1(r)-2, patella d1-1, tibia d1(r)-1-0, p0-1-1-0,
r0-0-1-0, v0-0-2, metatarsus d1(r)-2-2, p1-0-0, v2-2-3;
leg IV: femur d1-1-1(r)-1(r)-2, patella d1-1, tibia d1(r)-
1-1, p1-1-0, r0-1-1-0, v0-1(r)-2, metatarsus d1(r)-1-1,
p1-1-1, r0-1-0-1, v2-2-2, tarsus r0-1. Female: palp:
femur d1-1-3, patella d1-1, tibia 1(p)-1(r)-0-1(r),
tarsus d2-1(p)-1(p)-0, p1-0-1-1, r0-1-0-0, v1-1-2; leg I:
femur d1-2-1-1(r)-1(p)-2, p0-1-0, patella d1-1, tibia
d1(r), p0-1-0, v0-2-2, metatarsus v2-2-3; leg II: femur
d2-1-1(p)-2-2-2, patella d1-1, tibia d1(r), v2-2-2,
metatarsus p0-1-1, r0-0-1, v2-2-2; leg III: femur
d3-1(p)-1-1(r)-1(p)-1(r)-2, patella 1-1, tibia d1(r)-1-0,
p0-1-0, r0-1-0, v2-2-2, metatarsus d2-0-0, p0-1-2, r0-1-
1, v2-2-2, tarsus r0-1; leg IV: femur d1-2-1(p)-1(r)-2,
patella 1-1, tibia d1(r)-0-1-0, p0-1-0, r0-1-0, v2-2-2,
metatarsus d2-2-2-0, p0-0-2, r0-0-1, v0-2-2-2, tarsus
r0-1. Spination can vary slightly between and within
Figure 15. Various somatic characters of Porteria. A, face in anterior view, female P. eddardstarki sp. nov. from Pichicuy
MACN-Ar21060. B, eyes in anterior with visible canoe-shaped tapetum in ALE, female P. eddardstarki sp. nov. from Zapallar
CASENT9055699. C, ventral view of right trochanters, III and IV with deeper notch, female P. bunnyana sp. nov. from Pucón
CASENT9020914. D, spinnerets, female P. eddardstarki sp. nov. from Zapallar CASENT9055699.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 23
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Figure 16. Scanning electron micrographs of various somatic characters of Porteria. A, B, D-F, P. correcaminos sp. nov.
CASENT9021323. C, P. faberi sp. nov. CASENT9055712. A, eyes in anterior view. B, carapace in dorsal view. C, chelicera in
posterior view, female. D, tracheae in dorsal view, CASENT9021323. E, pedicel in dorsal view. F, spinnerets in lateral view.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
24 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
species, even between left and right legs of the same
individual. Tibia I with one to three pairs of ventral
spines; spination listed in species descriptions when
possible. Single row of trichobothria on tarsus and
metatarsus, increasing in length distally; few
scattered trichobothria on tibia, present on the male
and female palps; trichobothrial base as in Figure
17E, simple, hood smooth. Tarsi three clawed, with
many teeth on the superior claws and two teeth on
the smaller, inferior claw (Fig. 17A, C), female palp
with single claw with several teeth (Fig. 17D). Tarsal
organ flat, aperture tear shaped, located on dorsal
surface, less than 0.2mm from claw (Fig. 17F).
Trochanters shallowly notched; notch more obvious
on leg IV (Fig. 15C). Spinnerets as follows (Figs 5–7):
anterior spinnerets broad, much wider than long;
bases almost touching. Apical segment small relative
to basal segment. Posterior spinnerets much narrower,
Figure 17. Scanning electron micrographs of various somatic characters. A, P. correcaminos sp. nov. female CASENT9021323,
right claws I, retrolateral view. B, female CASENT9021297, tibia IV chemosensory (centre, bright) and feathery setae. C,
E, F, male P. bunnyana sp. nov. from Flor de Lago, CASENT9044676. D, female P. faberi sp. nov. from Reserva Costera
Valdiviana, CASENT9055712. C, tarsal claws on leg IV. D, claw of left palp. E, trichobothria socket on leg I. F, tarsal organ
on leg IV.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 25
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
with medians small and short and laterals more
elongate. Apical segments smaller, 1/4 to 1/5 the
length of basal segment. Anterior lateral spinnerets
(ALS) with spinning field of males and females with
distinctive ‘tail’ formed by the narrowed piriform
gland spigot field curving inward and posteriorly
towards base from circular main field (Fig. 5B). A line
of five to six stout setae just posterior to spinning
field. Female spinnerets as follows: ALS piriform
gland spigots (PI) larger in the main field and
decreasing in size towards the piriform field tail’s
apex; numerous piriform tartipores (Tp). Two major
ampullate gland spigots (MAP) and major ampullate
tartipore clustered, sunken down and mesad to the
main piriform gland spigot field (Fig. 5C). PMS with
aciniform gland spigots (AC) and two cylindrical
gland spigots (CY) nested within the aciniforms, one
on anterior margin, another on posterior margin.
Posterior lateral spinnerets (PLS) with aciniform
gland spigots, with more elongate shafts than those
on the posterior median spinnerets and a single
anterior cylindrical gland spigot. Spinnerets of males
as follows: ALS with spinning field similarly shaped
to that of females, with piriform gland spigots,
numerous piriform tartipores, and a single MAP
gland spigot with adjacent MAP nubbin (Nu) and
tartipore. PMS with aciniform gland spigots. PLS
with aciniform gland spigots. Colulus (Cl) of both
sexes linguiform with setae on anterior half (Fig. 5F).
Tracheal system (Fig. 16D): spiracle just anteriad of
colulus, less than colulus’ length away; lateral
tracheae unbranched, medians branched, all tracheae
limited to opisthosoma. Male palp with cymbium
greatly elongated distal to bulb (Figs 25E, 28E, 32E,
38E, 45E, 50E, 55D, 58D, 61E, 66D, 71E); cymbium
2.21–5.94 times length of bulb. Prominent bulb
Figure 18. Right legs of male P. eddardstarki sp. nov. from Zapallar, CASENT9055699, prolateral view. A, leg I. B, leg II.
C, leg III. D, leg IV. Scale bars = 1mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
26 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
structures include tegulum with median concavity
present or absent, fleshy conductor, and a slender
embolus of varying lengths; embolus base varies in
shape and is diagnostic in species identification. Tibia
of male palp with three to four apophyses: stout,
acuminate RTA; needle-like LRTA positioned mesad
to RTA; dark, flattened ventral apophysis (VTA);
dorsal tibial apophysis (DTA) present or absent,
shape diagnostic for some species. Knob-like
paracymbium (PC) on retroapical side of alveolus
present or absent. Palpal bulb (Fig. 68B, D) without
median apophysis; embolus (E) with slender tip
converging onto large, fleshy conductor (C). Epigyne
and vulva highly variable; heavily sclerotized or
fleshy, with little to no sclerotization; median scape
(Sc) present in all species but varies in form and size.
Epigyne usually partially obscured by a covering of
long setae. Vulva with short to long copulatory ducts
(CD), spermathecal head (HS) differentiated or not
from rest of spermathecae; pores present on head and
spermathecal stalk (SS); stalk leads to base 1 and
base 2 of the spermatheca. Base 1 of spermatheca can
be distinguished from base 2 by the presence of the
Bennett’s gland (BG) and the attachment of the
fertilization duct (FD). Females of the bunnyana
species group with a central atrium, with ventral
wall flexible, wrinkled (Fig. 64D, F), and a pair of
invaginations on the posterior margin of the epigyne
(Fig. 64A, B; uncertain where they lead).
Biology: The biology that follows is mostly known from
observations made in the field during a January 2013
excursion by the authors Charles Griswold, Elizabeth
Morrill and colleagues Hannah Wood, David Faber and
Figure 19. Right legs of male P. eddardstarki sp. nov. from Zapallar, CASENT9055699, retrolateral view. A, leg I. B, leg II.
C, leg III. D, leg IV.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 27
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
Luke Macaulay, complemented with data from several
excursions by Martín Ramírez. Porteria are sheet web
builders that construct webs in low lying vegetation,
fallen logs and leaf litter (Fig. 2). They are abundant
with many individuals residing in a small area. The
spiders walk or run on top of the sheet. The web itself
is made of a strong, finely woven sheet; above this is
a loosely woven system of knock-down threads that
probably help prevent prey from escaping, and a mess
of supporting lines to secure the sheet to the substrate.
At one edge of the sheet lies the funnelled retreat that
leads into leaf litter, hollow logs or other crevices.
Porteria were often found near austrochilids and seem
to prefer a similar habitat, though austrochilid webs
are generally much higher off the ground than those
of Porteria. The web of Porteria can be distinguished
from often sympatric linyphiid spiders by the sheen
and glittering appearance of the silk in the sunlight;
linyphiid webs often appear softer and duller;
Porteria’s sheet is stronger to the touch than those of
linyphiids. Porteria sheets are not found on artificial
or man-made substrates. The spiders often hide in the
funnel retreats during the day or when their web has
been disturbed. At night, the spiders stand just outside
of the funnel and are much more active pursuing
prey. Many spiders were successfully collected by
mimicking small prey vibrations on the sheet (either
by aspirating small insects or using small twigs) to
draw the spider out of the retreat and then a spoon
was used to cut off access back into the retreat; the
spiders were fast, and if they were able to retreat into
the funnel, they were often lost in the great expanse
Figure 20. Left legs of female P. eddarstarki sp. nov. from Zapallar, CASENT9055699, prolateral view. A, leg I. B, leg II. C,
leg III. D, leg IV.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
28 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
of leaf litter and other debris. It is expected that once
males reach maturity, they discontinue web building
and search for females. Adult males were often found
in penultimate and adult female webs at night. When
males did make small movements towards the female,
she would often move underneath the web nearby and
hang upside down. We know nothing of the eggs or
maternal care of Porteria.
Composition: Twelve species, including the type species
P. albopunctata Simon 1904, and 11 species newly
described here: P. ajimayo, P. alopobre, P. ariasbohartae,
P. bunnyana, P. contulmo, P. correcaminos, P.
eddardstarki, P. faberi, P. fiura, P.misbianka and P.
torobayo.
Distribution: Chile, ranging from IV Región de
Coquimbo at the northern end of their range to Punta
Arenas in Magallanes Province at the southern end of
their range (Fig. 76A-F).
Phylogenetics: Like the other porteriines
Nanocambridgea, Cambridgea and Corasoides, Porteria
have the remarkable narrowing of the ALS piriform
gland spigot field (Fig. 5 ), the cymbium highly elongated
beyond the bulb and lack a median apophysis, but
Porteria differ from all of these by the synapomorphies
of the presence of a thin tibial apophysis (LRTA) arising
near the base of the RTA on the male palpal tibia (Fig.
68) and a line of five to six stout setae just posterior to
the ALS spinning field of both sexes (Fig. 5B).
Figure 21. Left legs of female P. eddarstarki sp. nov. from Zapallar, CASENT9055699, retrolateral view. A, leg I. B, leg II.
C, leg III. D, leg IV.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 29
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
the AlboPunctAtA sPecies grAde
This morphologically distinctive set of species is recog-
nized by the common plesiomorphic genitalic charac-
ters of both males and females. Our phylogeny shows
this set to be paraphyletic, so we refer to this as the
Albopunctata Grade. Most diagnostic are the tegulum
without a median concavity (Figs 26B, 29B, 33B), out-
lined by the dark sperm duct; embolus base elongated
and narrow, not shield like nor heavily sclerotized,
hugging prolateral margin of conductor (Figs 26A, B,
29A, 33A, 39A, B). LRTA projected straight up from
tibia, unbent, in line with frontal plane. Paracymbium
absent (Figs 26C, 29C, 33C, 39C). Epigyne atrium ab-
sent; copulatory openings flush with surface of epigynal
plate (Figs 25F, 28E, 30A, 35A-C, 41A, 43E). Our phyl-
ogeny (Fig. 73) implies those are all plesiomorphic
traits for Porteria.
Included species: Porteria albopunctata and P.
ajimayo, P. eddardstarki, P. fiura and P. misbianka.
Porteria alboPunctata simon, 1904
Figs 23–26
Porteria albopunctata Simon, 1904: 109, figs 7, 8.
Lehtinen, 1967: 440, figs 119, 120. Roth, 1967: 324, pl.
52, figs 6–10.
Types: Lectotype male and paralectotype female
designated by Roth (1967), both from Punta
Arenas, Chile, in MNHN 22323, jar 1933, examined
(CASENT9055543).
Diagnosis: The males of this species can be
distinguished from others in this species grade by
the slender, hook-shaped RTA (Fig. 26C) and the
long spiniform DTA curving retrolaterally (Figs
25E, 26D). Embolus base less bulbous than in P.
fiura (Fig. 29A, B), embolus tip less hooked (Fig.
26A, B). Embolus tip significantly shorter than in
P. eddardstarki (Fig. 33A). Fangs constricted at
midpoint (Fig. 23D). The dark sclerotized middle of
Figure 22. Spine map of left palps and legs of P. eddardstarki sp. nov. for a female (CASENT9055699) from Zapallar and
a male (CASENT9044690) from Valparaíso. Shared spines imply similar spine location between the sexes. Spines slightly
vary within a species and even between right and left legs. Generally, the above map is a good representation for the genus.
See Griswold (1987) for full explanation of spine map (V, ventral; R, retrolateral; D, dorsal; P, prolateral).
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
30 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
key to sPecies oF Porteria
1a. Male ............................................................................................................................................................(2)
1b. Female ...................................................................................................................................................... (12)
2a. Paracymbium absent (Figs 26C, 29C, 33C, 39C); LRTA directed apicad, unbent (Figs 26D, 29D, 39D);
tegulum without median concavity (Figs 26B, 33B), or if present, shallow (Fig. 39C) .......................... (3)
2b. Paracymbium present (Figs 46C, 51C, 59C, 62C, 67C, 72C); LRTA bent, projecting ventrally perpendicu-
lar to frontal plane (Figs 46C, 51C, 56C, 59C, 62C, 67C, 72C); tegulum concave, with median concavity
(Figs 46B, 56B, 59B, 67B, 72B) ..................................................................................................................(6)
3a. Palp with extremely long, hair-like embolus (Figs 39A, B, 40D); embolus base reduced to small
mound on base of bulb on retrolateral side (Fig. 38B); conductor long, deeply grooved and tapering
to a point at apex well past bulb (Fig. 40A, D). Tegulum with shallow (Fig. 39C) or no median concavity
....................................................................................................................................................... P. ajimayo
3b. Embolus of short to medium length, less than length of bulb (Figs 26A, 29A, 33A); embolus base
more elongated, at least 1/3 length of bulb; conductor a straight projection on median line of bulb,
simple and untwisted (Figs 26B, 29B, 33B). Tegulum convex, without median concavity (Figs 26B, 29B,
33B) ...........................................................................................................................................................(4)
4a. DTA absent (Figs 32E, 33C, 34C). Embolus greater than ½ length of bulb; embolus base less than
½ length of bulb (Fig. 32D). RTA short and stout, with thickly pointed apex (Figs 33B, 34A)
............................................................................................................................................... P. eddardstarki
4b. DTA present (Figs 28B, 29D). Embolus less than ½ the length of bulb; embolus base elongated, greater
than ½ length of bulb. RTA bulbous (Fig. 28B) or slender and hook like (Fig. 25A) .............................(5)
5a. DTA small, tooth like (Fig. 29D). RTA round and bulbous, curving ventrad at apex (Figs 28B, D, 29B).
Embolus base long, almost length of bulb, abruptly tapering to thick, hook-like embolus (Fig. 29A, B)
............................................................................................................................................................ P. fiura
5b. DTA elongated and spiniform (Figs 25E, 26D), directed towards retrolateral side. RTA long, slender and
hook like (Figs 25A, 26B). Embolus base shorter and more gradually tapering to embolus (Fig. 26A, B)
...............................................................................................................................................P. albopunctata
6a. DTA serrate (Figs 50D, 51C, 52B) ....................................................................................................P. faberi
6b. DTA smooth, at most with a few grooves, varying forms (Figs 46C, 56C, 62C, 67C, 72D) .....................(7)
7a. Embolus base with notch on anterior margin where embolus starts to differentiate (Figs 46B, 67B),
shallow in some species (Fig. 62B) ...........................................................................................................(8)
7b. Embolus base with anterior margin smooth, embolus differentiating in line with base (Figs 56B, 59B,
72B) ..........................................................................................................................................................(10)
8a. DTA with sharp apex directed retrolaterally, rounded base, arrowhead like in dorsal view, appearing
anvil shaped in lateral views (Figs 61E, 62D, 66D, 67D) ........................................................................(9)
8b. DTA thick and stubby in dorsal view (Figs 46D, 47B), ‘U’-shaped hook in lateral views (Figs 45A, D, 46C,
47C) ............................................................................................................................................P. bunnyana
9a. Apex of DTA highly elongate, spear like (Figs 66D, 67D) ......................................................... P. contulmo
9b. Apex of DTA shorter, tooth like (Figs 61E, 62D) ................................................................. P. correcaminos
10a. DTA thick and thumb like in lateral views (Figs 56C, 59C) ................................................................. (11)
10b. DTA slender, finger like in retrolateral view (Fig. 7I D), not visible in prolateral view (Fig. 72A)
............................................................................................................................................. P. ariasbohartae
11a. Embolus base narrow, approximately ½ width of bulb (Fig. 56B), RTA with long, tapered apex (Fig. 56B,
D) ................................................................................................................................................ ..P. torobayo
11b. Embolus base wide, greater than ½ width of bulb (Fig. 59B), RTA with thickened apex and tooth-like
projection on anterior margin (Fig. 59B, D) ............................................................................... P. alopobre
12a. Epigyne with copulatory openings in atrium (Figs 45E, 48A, 53A, 55E, 61F, 64A-C) ......................... (13)
12b. Epigyne without atrium, copulatory openings flush with surface of epigynal plate (Figs 25F, 28E, 35A-
C, 41A, 43E) .............................................................................................................................................(18)
13a. Atrium with anterior margin less than or equal to width of posterior margin (Figs 48A, 64A); median
septum absent, but fleshy median bulge may be present or absent; lateral margins of atrium ridged
(Figs 45E, 61F, 64C, 69A, 71F) ................................................................................................................ (14)
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 31
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
the epigynal area, as well as the ‘X’ like configuration
of the internal structures visible through the cuticle,
are also diagnostic (Fig. 25F, H). Internally, the short
copulatory ducts (Fig. 25H) separate this species from
all but P. fiura. Vulvae of P. fiura and P. albopunctata
are easily distinguished, with spermatheca stalks
almost touching in P. albopunctata (Fig. 25H) and
conspicuous Bennett’s gland pores in P. fiura, visible
in dorsal view (Fig. 28F).
Description: Male: Based on lectotype. Markings
as in Figure 23A-C, dorsum of abdomen with yellow
anterolateral lines and five or six pairs of median
spots; sternum with pale centre surrounded by dark
grey. Total length 5.70. Carapace length 1.33 times
width. Clypeus height 2.5 times AME diameter. ALE
diameter 2 times AME diameter. Chelicera length
6.2 times clypeus height, fang with constriction
between base and apex (Fig. 23D). Sternum length
13b. Atrium with wide anterior margin forming hood (Figs 53A, 55E); atrium appearing wider anteriorly
than posteriorly due to lateral lobes; smooth, wide median septum retreating into atrium anteriorly;
lateral margins of atrium smooth (Figs 50F, G, 53C, D, 55E) ...............................................................(17)
14a. Atrium with large, fleshy median bulge (Figs 61F, 64A, 71F); scape attached to bulge, small and thumb
like ............................................................................................................................................................ (15)
14b. Atrium without median bulge (Figs 48A, 66F); scape larger and tooth like, attached medially to anterior
margin of atrium ......................................................................................................................................(16)
15a. Large groove between lateral margins and posterior lobes of atrium (Figs 61F, 64A, C); scape on small
bulge on anterior of epigyne (Fig. 64C) ...............................................................................P. correcaminos
15b. Lateral margins of atrium continuous with posterior margin, folded but no large groove; median bulge
extended all the way to posterior margin (Fig. 71F); scape on posterior tip of bulge overlapping posterior
margin ................................................................................................................................. P. ariasbohartae
16a. Vulva (Fig. 66E, G) with atrium ventral wall extended well past head of spermatheca, with series of
ridges; copulatory ducts ‘S’-shaped; right and left Base 1s of spermathecae with small gap between
them............................................................................................................................................. P. contulmo
16b. Vulva (Figs 45G, 48B) with atrium ventral wall extended just beyond spermatheca, smooth, heart
shaped with deep median ‘V’ shape; copulatory ducts straight; right and left Base 1s of spermathecae
almost touching ..........................................................................................................................P. bunnyana
17a. Epigyne with small median scape on anterior hood, finger like (Fig. 53A). Vulva with broad atrium
ventral wall, taller than stalks of spermatheca and wider than base receptacles; left and right stalks of
spermatheca separated by half of their lengths (Figs 50H, 53B) ..................................................P. faberi
17b. Epigyne lacking scape on anterior hood (Fig. 55E). Vulva with narrow and petite atrium ventral wall,
stalks of spermatheca almost reaching anterior margin of atrium ventral wall; left and right stalks of
spermatheca separated by less than 1/3 of their lengths (Fig. 55G) ........................................ P. torobayo
18a. Epigyne with long, sclerotized median strip (Figs 25F, 28E) ................................................................ (19)
18b. Epigyne without median strip (Figs 35A-C, 41A, 43E)..........................................................................(20)
19a. Darker median strip dilated anteriorly surrounding the small but visible scape (Fig. 28E); Bases 1 and
2 of spermatheca forming one large oval; left and right stalks not touching (Fig. 28F) ................ P. fiura
19b. Darker median strip not dilated anteriorly (Fig. 25F); fleshy mound anterior to dark strip but without
noticeable scape; internal structures form X-shape through cuticle (Fig. 25F); Bases 1 and 2 distinct
lobes; stalks touching (Fig. 25H) .......................................................................................... P. albopunctata
20a. Epigyne well sclerotized with two large posterior lobes (Fig. 35A-C); large median scape extended be-
yond posterior lobes (Fig. 36C).............................................................................................P. eddardstarki
20b. Epigyne without posterior lobes (Figs 41A, 43E); scape small (Fig. 43D) ............................................(21)
21a. Epigyne simple with little sclerotization (Fig. 41A); fleshy median septum dividing copulatory open-
ings; two dark circles posterior to openings representing internal receptacles. Tiny nub-like scape on
septum, visible in profile, difficult to see in ventral view. Vulva with extremely long copulatory ducts
making longitudinal loops at sides (Fig. 41B, D) ........................................................................ P. ajimayo
21b. Epigynal plate lightly sclerotized, tapered posteriorly (Fig. 43E). Rims of copulatory openings darkly
sclerotized, slit like. Small but elongated median scape anterior to openings, visible in ventral view
(Fig. 43D). Internal structures visible through cuticle; two round receptacles separated far apart are
connected by copulatory ducts forming a ‘peak’ in the middle of the epigyne (Fig. 43E). Copulatory ducts
‘S’-shaped, elongate but not making longitudinal loops at sides (Fig. 43F) .......................... P. misbianka
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
32 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
1.07 times width. Femur I 1.09 times carapace length.
Leg formula 4123 (for MCZ46441), legs not intact on
type specimen. Cymbium length 2.73 times length
of bulb. Cheliceral promargin with five teeth on left
chelicera and four teeth and one denticle on right. Leg
spination as follows: palp: femur d1-1-3, patella d1-1,
tarsus p0-0-1, r0-0-1; leg I: femur d1-1-1(p)-2, patella
d1-1, tibia d1(r)-0-0, v0-0-2, metatarsus p0-01, r0-0-
1, v2-2-1; leg II (right): femur d2-1(p)-1-2-2, patella
d1-1, tibia d1(r)-0-0, p0-1-1-0, v0-1(r)-2, metatarsus
p0-1-2, v2-2-2; leg III: femur d3-1-1(p)-2, patella d1-1,
tibia d1(r)-0-0, p0-1-1-0, r0-1-1-0, v0-0-2, metatarsus
d2-0-1, p0-1-0-1, r0-1-0-2, v1(r)-2-1, tarsus r0-1, v0-1;
leg IV: femur d1-1-0-2, patella d1-1, tibia d1(r)-0-0,
p0-1-1-0, r0-1-1-0, v1(p)-0-2, metatarsus d1(r)-0-1-02,
p1-1-0, r0-1-0-1, v2-1(r)-2, tarsus r0-2, v0-3. Palp as in
Figures 25A-E and 26A-D, with four tibial apophyses:
VTA flattened and triangular in ventral view, slim,
curved plate in retrolateral view; slender hook-shaped
RTA; LRTA in line with frontal plane; DTA long and
spiniform, directed retrolaterally; LRTA and DTA
appear connected, as in one bifurcating structure.
Tegulum with smooth surface, dark sperm duct
following outer margin of tegulum. Conductor simple,
column like. Embolus base narrow and elongated,
parallel to the conductor; lightly sclerotized. Embolus
tip short, gradually tapered to tip, resting on conductor.
Paracymbium absent.
Leg measurements (male MCZ 46441, Puerto Edén,
right): leg I 11.52 (3.08, 3.95, 2.75, 1.74); leg II 10.05
(2.88, 3.28, 2.41, 1.47); leg III 9.45 (2.61, 2.95, 2.54,
1.34); leg IV 12.19 (3.22, 3.82, 3.55, 1.61); palp 5 (1.95,
1.05, –, 2.00).
Description: Female: based on CASENT9044707,
from Laguna San Rafael National Park. Markings
as in Figure 24A-C, dorsum of abdomen with yellow
anterolateral lines followed by two or three posterior
spots; four or five pairs of median spots; sternum with
pale centre surrounded by dark grey. Total length 5.76.
Carapace length 1.38 times width. Clypeus height
2.6 times AME diameter. ALE diameter 2.4 times
AME diameter. Chelicera length 6.15 times clypeus
height. Sternum length 1.08 times width. Femur I 1.04
times carapace length. Leg formula 4123. Cheliceral
promargin with four teeth and one denticle right,
Figure 23. A-C, male habitus of P. albopunctata sp. nov., lectotype from Punta Arenas, MNHN22323, CASENT9055543.
A, dorsal view. B, ventral view. C, lateral view. D, chelicera and fang, anterior view, showing fang constriction, from Puerto
Edén MCZ46441.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 33
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
five teeth left. Leg spination as follows: (right legs of
CASENT9044702, Laguna San Rafael) palp: femur
d1-1-3, patella d1-1, tibia d1-1, p1-0-0, tarsus d1(r)-0-0,
p1-1, r1-1, v2-2-2; leg I: femur d1-1-0-2, p0-0-1-0, patella
d1-1, tibia d1(r)-0-0, v0-0-2, metatarsus p0-0-1, r0-0-1,
v0-2-1; leg II femur d2-1-1(p)-2, patella d1-1, tibia d1(r)-
0-0, p0-0-1-0, v0-1(r)-2, metatarsus p0-1-1, r0-0-1, v0-2-
1; leg III: femur d3-1-1(p)-2, patella 1-1, tibia d1(r)-0-0,
p0-1-1-0, r0-0-1-0, v0-1(p)-2, metatarsus d0-0-1, p0-1-1,
r0-1-1, v1(p)-2-1; leg IV: femur d1-0-1(p)-2, patella d1-1,
tibia d1(r)-0-0, p1-1-0, r0-0-1-0, v0-1(p)-2, metatarsus
d2-1-1, p0-1-0-1, r0-1-0-1, v0-1(p)-2, tarsus r0-1, v0-1.
Epigyne as in Figure 25F-H, with a small fleshy lip in
the middle of the anterior margin with minuscule fleshy
scape, copulatory openings on either side, inconspicuous.
Darkly sclerotized rectangular band running down
middle of epigyne to epigastric furrow; internal
structures visible through cuticle, dark orange, arranged
in an ‘X’ with posterior arms longer than anterior. Vulva
with short copulatory ducts anteriorly, curved from
middle to meet the stalk of the spermatheca adjacent
to head; head at anterior tip of stalk with pores; stalk
banana shaped, bowed out, with stalks almost touching
in centre. Base 1 large and more elongate than Base 2
which sits anteriorly to Base 1 (with copulatory ducts
on its anterior surface); fertilization ducts positioned on
the posteromedian surface of Base 1. Bennett’s gland
pore not visible in dorsal view.
Leg measurements (right): leg I 10.10 (2.80, 3.50,
2.30, 1.50); leg II 8.60 (2.60, 2.80, 1.95, 1.25); leg III
8.35 (2.40, 2.60, 2.15, 1.20); leg IV 10.95 (3.10, 3.50,
3.00, 1.35); palp 3.60 (1.30, 1.10, –, 1.20).
Variation: (N = 2). Total length 4.75–5.76. Carapace
length 1.27–1.39 times width. Clypeus height 2.4–2.5
times AME diameter. ALE diameter 2 times AME
diameter. Chelicera length 4.93–5.5 times clypeus
height. Sternum length 0.96–1.07 times width. Femur
I 1.17 times carapace length. Cheliceral promargin
with five teeth. Leg III tarsus: v0-3, leg IV tarsus: r0-1,
v0-3.
Distribution: Known only from the far south of Chile
with a northernmost range of Laguna San Rafael
Figure 24. Habitus of female P. albopunctata sp. nov. from Laguna San Rafael National Park; A and B CASENT9044707.
A, dorsal view. B. ventral view. C, lateral view, CASENT9044702. Scale bars = 1mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
34 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
National Park, extending south to Punta Arenas (Fig.
76C).
Other material examined: XI Aysén: Aisén Province,
Laguna San Rafael National Park, May 16–24, 1990,
L.E. Peña, three females, AMNH (CASENT9044702,
9044707). XII Magallanes: Wellington Island, Puerto
Edén c. 49ºS, December 10–13, 1962, P.J. Darlington,
one male (MCZ46441).
Notes: No specimens of P. albopunctata were found
on our Chile expedition in January 2013 and great
Figure 25. Genitalia of P. albopunctata sp. nov., A-E, lectotype from Punta Arenas, MNHN22323, CASENT9055543. A,
ventral view. B, retrolateral view. C, dorsal view. D, prolateral view. E, ventral view with entire cymbium. F-H, female from
Laguna San Rafael National Park, CASENT9044702. F, ventral view. G, lateral view showing small scape. H, dorsal view
showing internal structures. Scale bars = 0.2mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 35
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
effort was made to search several different habitats
within the Punta Arenas Peninsula, including the wet
Nothofagus forests on the southern coast, urban parks,
green median strips along roadways and residential
areas within the city, and the dry rocky shrubland
to the north-west by the shores of Seno Otway. The
locality information for the types is vague; the range
may have shifted farther north since these were
collected more than 100 years ago, e.g. to Laguna
San Rafael, or perhaps the species has become more
isolated along the many islands off the Patagonian
coastline. In places where other Porteria species are
found, they are widespread, and their conspicuous
sheet webs dominate the landscape, as such they are
not easily overlooked.
Porteria fiura sp. nov.
Figs 27–30
Zoobank registration: urn:lsid:zoobank.
org:act:D7676422-0EB3-461E-B74D-9CB5A7C44B99.
Types: Female holotype from Chile, X Región de los
Lagos: Chiloé Province, Chiloé Island, Chiloé National
Park, Sendero Tepual, 2.3 km NNE Cucao, S42º37.039’,
W74º06.129’, elev. 1 m, January 16, 2013, C. Griswold,
E. Morrill and D. Faber, ‘general collecting in bog
forest dominated by Tepualia’ deposited in MHNS
(CASENT9055645). Male paratype from Chiloé Island,
no specific locality, December 15–18, 1985, L.E. Peña,
AMNH (CASENT9044672).
Etymology: Named after La Fiura, an ugly, villainous
woman who lives in the forest and clothes herself in
moss; according to Chilote mythology, she seduces
men and then drives them insane. Chiloé Island is
the type locality, whose culture is rich in mythology.
Sheet webs of P. fiura are often built on mossy
substrates.
Diagnosis: Males can be distinguished from others
in the Albopunctata Grade by the round, stubby
RTA (Figs 28B, 29B, C) and the small tooth-like
DTA (Fig. 29D). The embolus base is wider (Fig.
29B) than that of P. albopunctata and the embolus
tip more hook shaped (Fig. 29A); embolus tip
much shorter than in P. eddardstarki. Females
most closely resemble P. albopunctata but can be
Figure 26. Left male palp of P. albopunctata sp. nov. from Wellington Island, Puerto Edén, MCZ46441. A, prolateral view.
B, ventral view. C, retrolateral view. D, dorsal view. Illustration by Rachel Diaz-Bastin.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
36 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
distinguished by the sclerotized median area being
dilated anteriorly in P. fiura as well as the presence
of a prominent scape (Figs 28E, 30A, B). Internally,
the spermatheca stalks are situated much farther
apart in P. fiura than P. albopunctata; Bennett’s
gland pores large and conspicuous in dorsal view
(Figs 28F, 30B).
Description: Male: based on paratype
CASENT9044672. Markings as in Figure 27A-C,
dorsum of abdomen with yellow anterolateral lines
followed by two posterior spots and two pairs of
median spots; sternum with pale centre surrounded by
dark grey border. Total length 5.63. Carapace length
1.37 times width. Clypeus height 2.29 times AME
Figure 27. Habitus of P. fiura sp. nov. A-C, male from Chiloé Island CASENT9044672. A, dorsal view. B, ventral view. C,
lateral view. D-F, female from Chiloé National Park CASENT9056299 and 9055645. D, dorsal view. E, ventral view. F, lateral
view. Scale bars = 1mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
SYSTEMATICS OF PORTERIA SPIDERS 37
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
diameter. ALE diameter 1.71 times AME diameter.
Chelicera length 6.28 times clypeus height. Sternum
as long as wide. Femur I 1.37 times carapace length.
Leg formula 4123. Cymbium length 3.25 times length
of bulb. Cheliceral promargin with four teeth and one
denticle. Leg spination as follows: palp: femur d1-1-3,
patella d1-1, tarsus p0-0-1, v0-0-1(r); leg I: d1-1-1(p)-2,
patella d1-1, tibia d1(r)-0-0, v0-0-2, metatarsus v0-2-2;
leg II: femur d1-1-1(p)-2, patella d1-1, tibia d1(r)-0-0,
p0-0-1-0, v0-1(r)-2, metatarsus d0-0-1, p1-0-1, r0-0-1,
v0-2-1; leg III: femur d1-1(r)-2, patella d1-1, tibia d1(r)-
0-0, p0-1-1-0, r0-0-1-0, v0-1(r)-2, metatarsus d1(p)-0-1,
p0-1-0-1, r0-1-0-1, v0-2-2, tarsus r0-1-1(v), v0-1; leg IV:
femur d1-1(p)-2, patella d1-1, tibia d1(r)-0-0, p0-1-1-
0, v0-0-2, metatarsus d2-0-1, p0-1-1, r0-1-1-1, v0-0-2,
tarsus r0-1. Palp as in Figures 28 and 29, with four
tibial apophyses: VTA triangular, flattened plate in
ventral view, outline finger like in retrolateral view;
RTA short and apex round, curved, directed ventrally;
LRTA projected straight out of tibia, parallel to frontal
plane. DTA small and tooth like; DTA and LRTA
appear connected at fleshy base (Fig. 28D), like that
of albopunctata. Tegulum without median concavity,
outlined by dark sperm duct. Conductor originating
on the midline of bulb, short and simple projection.
Embolus base elongated, nearly the length of the bulb;
embolus tip short, spiniform, curved like a hook to
meet conductor. Paracymbium absent.
Leg measurements (right): leg I 13.50 (3.75, 4.50,
3.25, 2.00); leg II 11.15 (3.15, 3.50, 2.85, 1.65); leg III
10.55 (2.85, 3.25, 3.00, 1.45); leg IV 13.80 (3.75, 4.30,
4.20, 1.55); palp 4.85 (1.80, 1.10, –, 1.95).
Description: Female: based on CASENT9055645,
from Chiloé National Park. Markings as in Figure
27D-F. Total length 5.80. Carapace length 1.34 times
width. Clypeus height 1.86 times AME diameter. ALE
diameter 1.71 times AME diameter. Chelicera length
6.15 times clypeus height. Sternum as long as wide.
Femur I 1.24 times carapace length. Leg formula 4123.
Cheliceral promargin with five teeth and one denticle.
Leg spination as follows: palp: d1-1-3, patella d1-1, tibia
Figure 28. Genitalia of P. fiura sp. nov. A-D, male from Chiloé Island CASENT9044672. A, left palp, retrolateral view. B,
ventral view. C, prolateral view. D, dorsal view. E, F, female from Chiloé, CASENT9055645. E, ventral view. F, dorsal view,
showing internal structures. Scale bars = 0.2mm.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023
38 E. MORRILL ET AL.
© 2023 The Linnean Society of London, Zoological Journal of the Linnean Society, 2023, XX, 1–94
d2-0-1(r), tarsus d2-0, p0-1-1-0, r0-1-1-0, v2-1(r)-3; leg
I: femur d1-2-1(p)-2, patella d1-1, tibia d1(r)-0-0, v0-2-
2, metatarsus p0-0-1, r0-0-1, v2-2-1; leg II: femur d1-1-
1(p)-2, patella d1-1, tibia d1(r)-0-0, p0-1-1-0, v0-2-2,
metatarsus p0-1-0-1, r0-1-0-1, v1(r)-2-2; leg III: femur
d2-1-1(p)-2, patella d1-1, tibia d1(r)-0-0, p0-1-1-0, r0-0-
1-0, v0-1(r)-2, metatarsus d1(p)-0-2, p0-1-0-1, r0-1-0-
1, v2-2-1, tarsus r0-1, v0-2; leg IV: femur d1-0-1(p)-2,
patella d1-1, tibia d1(r)-0-0, p0-1-1-0, r0-1-1-0, v0-2-2,
metatarsus d2-1(r)-2, p0-1-0-1, r0-1-0-1, v2-2-1, tarsus
r0-2, v0-2. Epigyne as in Figures 28E, F and 30A, C,
lightly sclerotized except for narrow median area which
is dilated anteriorly where the scape originates; scape
small and finger like, pointed posteriad; copulatory
openings inconspicuous, located just anterior to scape,
divided by fleshy septum; internal structures visible
through cuticle, most prominently two large circular
patches on either side of sclerotized area. Vulva (Figs
28F, 30B, D) with anterior, short copulatory ducts
joining the spermathecal stalk just posterior to head
of spermatheca. Head situated at the anterior apex
of stalk, pores present. Stalks banana shaped, bowed
out laterally, pores present. Base 1 large circular
structure with large Bennett’s gland pore in its centre
when viewed dorsally; fertilization duct attached to
Base 1 on median posterior corner. Base 2 joining the
anterior margin of Base 1, somewhat crescent shaped;
copulatory duct resting on anterior surface of Base 2.
In total, the two receptacles look like one large, ovoid
receptacle; scanning electron micrograph shows the
division of the two bases in more detail (Fig. 30B).
Leg measurements: leg I 12.17 (3.32, 4.20, 2.85,
1.80); leg II 10.25 (2.95, 3.40, 2.45, 1.45); leg III 9.70
(2.80, 3.00, 2.55, 1.35); leg IV 12.65 (3.45, 4.00, 3.55,
1.65); palp 3.92 (1.32, 1.32, –, 1.28).
Variation: (N = 3). Total length 3.9–5.76. Carapace
length 1.27–1.37 times width. Clypeus height 1.67–2.2
times AME diameter. ALE diameter 1.67–2.2 times
AME diameter. Chelicera length 5–6.18 times clypeus
height. Sternum length 1–1.04 times width. Femur I
1.08–1.26 times carapace length. Cheliceral promargin
with four teeth and one denticle left, five teeth right.
Distribution: Only known from Chiloé Island and on
the mainland in Palena, 70 km south of Chaitén (Fig.
76C).
Other material examined: X Región de los Lagos:
Chiloé Province, same locality as holotype, nine
females, CAS (CASENT9055645); Palena Province, 70
km S of Chaitén, elev. 500 m, January 16, 1986, N.I.
Figure 29. Left male palp of P. fiura sp. nov. AMNH paratype, CASENT9044672, Chiloé Island. A, prolateral view. B,
ventral view. C, retrolateral view. D, tibia, dorsal view. Illustration by Rachel Diaz-Bastin.
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlac093/7152280 by guest on 11 May 2023