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Arachnologische Mitteilungen / Arachnology Letters 64: 83-92 Karlsruhe, Dezember 2022
e Italian spider fauna is amongst the most diverse in all
of Europe (Nentwig et al. 2022). However, the description
of multiple new taxa and the recurring identication of pre-
viously unrecorded species keep enriching knowledge of the
known biodiversity. Currently, 53 spider families are repre-
sented in Italy, for a total of 440 genera and 1700 species
(Pantini & Isaia 2019). Among these, Gnaphosidae is the
second most diverse family in the country, with a total of 169
recorded species and 31 genera.
Anagraphis Simon, 1893 is a small genus represented
by only seven known species distributed in the Palaearc-
tic and Afrotropical regions of the world (World Spi-
der Catalog 2022). Several of these taxa are quite ob-
scure, having been supercially described in the rst
half of the 20th century and largely ignored ever since.
Anagraphis ochracea (L. Koch, 1867) is a poorly studied and
rarely observed species, originally described as Liocranum
ochraceum from Corfu, Greece, and currently recorded also for
Albania, Northern Macedonia, Turkey, the Russian Caucasus
and Sardinia (World Spider Catalog 2022).
Until now, studies on the genus Anagraphis and, more spe-
cically, on A. ochracea, have focused exclusively on taxonomy
and distribution. Most of the more recent contributions have
concentrated only on Aegean species (Chatzaki et al. 2002a,
2002b), and have centered their work on preserved museum
material and dead specimens collected in the eld. No author
has ever provided behavioural or ecological notes for A. ochra-
cea or any similar, congeneric species.
Myrmecophily, or the tendency of certain species to live
within or near ant nests in strong symbiotic association with
ants (Donisthorpe 1927), has been observed in only 13 of
the 132 known spider families, for a total of 41 dierent spe-
cies (Cushing 1997, 2012). Fourteen of these are currently
recorded for Italy (Pantini & Isaia 2019). Although several
authors have noted and studied this peculiar behaviour with-
in the Araneae (Hölldobler & Wilson 1990, Cushing 1997,
2012, Witte et al. 2008, Nelson & Jackson 2009), most have
primarily focused on myrmecomorphy (mimcry of ants) and
myrmecophagy (the consumption of ants). As of today, very
few contributions have discussed myrmecophily and the so-
cial integration of spiders in ant colonies (Mendonça et al.
2019). Several myrmecophile spider species not only resort
to active, evasive manoeuvres to avoid direct interaction with
the ant, but have also developed the ability to chemically hide.
Some absorb, biosynthesise or imitate the cuticular hydrocar-
bon prole of the hosts in order to survive unscathed within
the ant colony (Cushing 2012). Others reduce the emis-
sion of cuticular hydrocarbons in order to remain unnoticed
(Parmentier et al. 2017). Given the very limited number of
contributions focusing on the subject (e.g., Castellucci et al.
2022), the study of myrmecophile spiders in Italy is a largely
unexplored eld.
Here, we record and report the presence of three distinct
populations of A. ochracea in south-central Italy. Field sam-
pling and extensive observations were carried out, both in the
First records of Anagraphis ochracea (Araneae: Gnaphosidae) for continental Italy
and Sicily with new observations on its myrmecophilous lifestyle
Luigi Lenzini, Filippo Castellucci, Mattia Poso, Alessandro Kulczycki, Enrico Simeon,
Gabriele Greco, Andrea Piccinini & Carlo Maria Legittimo
doi: 10.30963/aramit6410
Abstract. In the present study we describe and discuss for the rst time the peculiar myrmecophilous habits of Anagraphis ochracea
(L.Koch, 1867) and its strong association with the ant species Messor ibericus Santschi, 1931. The study is based on behavioural observa-
tions carried out both in the eld and in captivity, and sheds light on the lifestyle of this poorly studied and rarely observed species. We
also record the presence of A. ochracea on continental Italy and Sicily for the rst time, provide a brief overview of its taxonomic history
and present photographs of adult and juvenile specimens, the egg sac and the copulatory organs of both sexes. Finally, we provide a
DNA-barcode (COI) for A. ochracea, which is the rst for the genus Anagraphis as well.
Keywords: ant, ant association, Arachnida, Messor ibericus, myrmecophily, spider, symbiosis
Zusammenfassung. Erster Nachweis von Anagraphis ochracea (Araneae: Gnaphosidae) für das italienische Festland und Sizilien,
mit neuen Beobachtungen zum myrmekophilen Lebensstil der Art. In der vorliegenden Studie wird zum ersten Mal das besondere
myrmekophile Verhalten von Anagraphis ochracea (L. Koch, 1867) und ihre enge Bindung an die Ameisenart Messor ibericus Santschi,
1931 beschrieben und diskutiert. Die Studie basiert auf Beobachtungen zum Verhalten im natürlichen Lebensraum wie auch im Labor
und gibt Aufschluss über die Lebensweise dieser wenig erforschten und selten gefundenen Art. Ebenso wird der Erstnachweis von
A.ochracea für das italienische Festland und Sizilien erbracht, sowie eine Zusammenfassung der taxonomischen Historie, Bilder adulter
wie auch juveniler Tiere, der Kopulationsorgane beider Geschlechter und des Eikokons präsentiert. Zusätzlich wird erstmals der geneti-
sche Barcode (COI) der Art, und auch der Gattung Anagraphis, beschrieben.
Luigi LENZINI, Alessandro KULCZYCKI, Enrico SIMEON, Carlo Maria LEGITTIMO,
Aracnolia - Associazione Italiana di Aracnologia, via Gramsci 29, 33052 Cervignano
del Friuli, Italy; E-mail: luigilenzini@fastwebnet.it, alessandro.kulczycki@gmail.com,
enricosimeon@gmail.com, carlomarialegittimo@yahoo.it
Filippo CASTELLUCCI, Department of Biological, Geological and Environmental
Sciences, University of Bologna, via Selmi 3, 40126 Bologna, Italy; Zoology Section,
Natural History Museum of Denmark, University of Copenhagen, Universitetsparken
15, 2100, Copenhagen, Denmark; Aracnolia - Associazione Italiana di Aracnologia,
via Gramsci 29, 33052 Cervignano del Friuli, Italy;
E-mail: lippo.castellucci2@unibo.it
Mattia POSO , Museo di Storia Naturale del Salento, S.P. Calimera-Borgagne, km 1,
I-73021 Calimera, Italy; Aracnolia - Associazione Italiana di Aracnologia, via
Gramsci 29, 33052 Cervignano del Friuli, Italy; E-mail: posomattia@gmail.com
Andrea PICCININI, Department of Biological, Geological and Environmental
Sciences, University of Bologna, via Selmi 3, 40126 Bologna, Italy; Aracnolia –
Associazione Italiana di Aracnologia, via Gramsci 29, 33052 Cervignano del Friuli,
Italy; E-mail: andrea.piccio97@gmail.com
Gabriele GRECO, Department of Anatomy, Physiology and Biochemistry, Swedish
University of Agricultural, Sciences, Box 7011, 750 07 Uppsala, Sweden; Laboratory
of Bio-Inspired, Bionic, Nano, Meta, Materials & Mechanics, Department of Civil,
Environmental and Mechanical Engineering, University of Trento, Via Mesiano, 77,
38123 Trento, Italy; Aracnolia - Associazione Italiana di Aracnologia, via Gramsci 29,
33052 Cervignano del Friuli, Italy; E-mail: gabriele.greco@slu.se
Academic Editor: Tobias Bauer
Submitted: 16.11.2022, accepted 20.12.2022, online 28.12.2022
84 L. Lenzini, F. Castellucci, M. Poso, A. Kulczycki, E. Simeon, G. Greco, A. Piccinini, C. M. Legittimo
eld and in the lab, to thoroughly investigate its behaviour.
Signicant and previously unrecorded behavioural informa-
tion was obtained. We report for the rst time that A. ochra-
cea is strongly associated with the ant species Messor ibericus
Santschi, 1931, and lives unscathed and unnoticed within
their colonies.
e focus of this work is thus to present a series of obser-
vations that describe Anagraphis’s unreported myrmecophil-
ous lifestyle and the presence of a new species in continental
Italy and Sicily. Furthermore, we provide high quality photos
of the habitus and genitalia to further facilitate its identica-
tion, publishing the very rst contribution towards the ge-
netic characterization of both the target species and the genus
Anagraphis as a whole.
Materials and methods
During the course of this study, more than 40 dierent live
specimens were observed and analysed, both in the wild and
in the lab. e very rst hint of the strong myrmecophilous
habit of A. ochracea was observed and reported by Luigi Len-
zini in populations from the Lazio region.
Subsequently, similar observations were carried out by
Mattia Poso in southern Italy, specically in the area around
Lecce. All specimens were preserved in either 75% or 96%
ethanol. Over the years, several specimens encountered in
the Caarella Valley were collected from the eld and held
in captivity by L. Lenzini for additional behavioural obser-
vations. e spiders were raised or kept individually in 5cm
wide, squared glass enclosures covered by an opaque liq. e
setup was simple and comprised a layer of mixed terrain in-
tended to recreate the natural substrate found in the eld near
ant colonies. Enclosures were kept in a dimly lit environment
while water was sporadically sprinkled to simulate a rainy day
and to provide a degree of humidity to the substrate.
Specimen identication and morphological analysis was
carried out utilizing both a Zeiss Stereomicroscope II and a
Leica MZ16. e photos of preserved specimens were ob-
tained by connecting an Olympus E-M1 or an Olympus
E-M5mkII to the microscopes. Photos of the habitus and
of live specimens in the eld were obtained using the fol-
lowing cameras: Panasonic Lumix FZ28, FZ48, FZ200+
Raynox DCR250, Olympus E-M1 or Olympus E-M5 mkII
equipped with a Zuiko 60mm f2.8 + Raynox DCR250. Pho-
tos were stacked with Helicon Focus (Version 7.0.2) and pro-
cessed in Adobe Photoshop CC 2018.
e articial formicarium used to simulate a natural set-
ting in the lab was purposely constructed in plexiglass by
Valerio Dolci. e structure consists of a wide outside arena
and a series of interconnected and sub horizontal, underly-
ing chambers. After having observed and conrmed a clear
degree of mutual tolerance between A. ochracea and M. iberi-
cus under lab conditions, several other species of spiders were
introduced to test whether the same ant colony would react
dierently to these spiders. A various number of specimens
of Filistata insidiatrix (Forsskål, 1775) (Filistatidae), Ly-
cosoides coarctata (Dufour, 1831) (Agelenidae), Steatoda nobi-
lis (orell, 1875) (eriidiidae), and Scotophaeus blackwalli
(orell, 1871) (Gnaphosidae) were utilized. ese were se-
lected among non-myrmecophilic species that are commonly
found in the same area as A. ochracea in the Caarella Park
and being of comparable dimensions to A. ochracea.
Genomic DNA was extracted from two legs of each
specimen using the NucleoSpin® DNA Insect kit (Mach-
erey-Nagel) and following the manufacturer’s instructions. A
partial segment of gene locus cytochrome oxidase subunit I
(COI) was targeted for PCR amplication using the primer
pair LCO1490-HCO2198 (Folmer et al. 1994) as per proto-
col established by Wheeler et al. (2017). PCR products were
utilized for DNA electrophoresis on a 1% agarose gel and
puried, prior to sequencing, using the ExoSAP-IT Prod-
uct Cleanup Reagent (ermo Fisher Scientic). Sanger se-
quencing for both forward and reverse reads was performed
by Macrogen Europe (Amsterdam, Netherlands). Chroma-
tograms were read using the SeqTrace v.0.9.0 software, while
a BLASTn (Zhang & Madden 1997) search was run on the
NCBI database to test for possible contamination. e result-
ing sequence was submitted to GenBank.
All information pertaining to the annual weather and rain-
fall of the sampled localities was obtained from the World-
Clim 2.1 database (https://www.worldclim.org/). Specimens
observed during this study are preserved and deposited in the
following institutions: MNHT, Museum of Natural History
of Trieste, Trieste, Italy; MSNS, Natural History Museum of
the Salento, Calimera, Italy; NHMB, Natural History Mu-
seum, London, UK; ZMUA, Zoological Museum of the Uni-
versity of Athens, Greece; NHMC, Natural History Museum
of Crete, Greece.
Taxonomy and distribution
Anagraphis ochracea (L. Koch, 1867) (Figs 1, 3)
For complete taxonomic references, see World Spider Cata-
log (2022)
Type material. Holotype ( of Liocranum ochraceum from
Corfu (GREECE): NHMB (MB b842); examined. Well
preserved and in good condition.
Holotype ), paratype ( of Talanites pallidus from Attica
(Attiki), Penteli-Dionysos and Parnis-Phyli (GREECE):
Coll. Hadjissarantos (ZMUA). Not examined. e type se-
ries could not be found in the ZMUA museum collection or
in any other Greek collection (C. Georgiadis, E. Valakos, M.
Chatzaki pers. comm.) and is presumed lost.
Examined material. ITALY: Lazio: Rome (RM), Parco Re-
gionale dell’Appia Antica, (41.862681°N, 12.514195°E): 1 (,
25. Feb. 2012, L. Lenzini (MNHT); 1 ), 14. Jul. 2012, L. Len-
zini (MNHT); 1 ), 1 (, 18. Apr. 2018, L. Lenzini (MNHT);
3 )), 3 ((, 15. May 2018, L. Lenzini (MNHT); 2 )), 11. Jun.
2018, C. M. Legittimo (MNHT); 1 (, 20. Feb. 2019, L. Len-
zini (MNHT); 2 )), Feb. 2019, L. Lenzini (MNHT); 1 ),
15. Jun. 2019, L. Lenzini (MNHT). Apulia: Lecce (LE), fraz.
Solicara, (40.429889°N, 18.173889°E): 1 ( subadult, 16. Aug.
2018, M. Poso (MNHT); 1 (, Mar. 2019, M. Poso (MNHT);
1 ), 12. Apr. 2019, M. Poso (MNHT); 1 ), 19. Apr. 2019,
M. Poso (MSNS); Sicily: Buccheri (SR), (37.124667°N,
14.848194°E): 1 ), 29. Sep. 2020, G. Romagna (MNHT).
GREECE: Trikala: Antichasia: 1 (, May – Jun. 2011,
(NHMC FC 11602); Pieria: Litochoro: 1 ), 19. Jun. – 9.
Aug. 2014, (NHMC FC 8798); Magnesia: Alikes: 1 ), 23 .
Apr. – 27. Jun. 2014, (NHMC FC 9276); Portaria to Chania:
1 ), 24. Apr. – 28. Jun. 2014, (NHMC FC 9281); Afetes: 2
)), 1(, 25. Apr. – 29. Jun. 2014, (NHMC FC 9283); Genati:
1 (, 25. Apr. – 30. Jun. 2014, (NHMC FC 9284); Samos
New records for the myrmecophilous Anagraphis ochracea 85
island: Psili Ammos: 1 ), 6. Jul. – 30. Jul. 2006, (NHMC FC
9214); Corfu: Corfu: 1 (, Josef Erber, Holotype (NHMB
BM b842).
Additional material examined
Anagraphis pallens Simon, 1893
GREECE: Karpathos, Pigadia-Aperi: 5 )), 5 ((, 12. May –
23. Aug. 2001 (NHMC FC 1774); Pyles-Volada: 2 )), 3 ((,
12. May – 23. Aug. 2001 (NHMC FC 1772); Avlona: 1),
12. Oct. 2013 (NHMC FC 15973); Achordea: 2 )), 2 ((,
12. Oct. 2013 (NHMC FC 15974); Rhodes: Afantou-Psin-
thos: 3 )), 1 (, 14. May – 10. Jul. 2006 (NHMC FC 8461).
Taxonomic history and distribution
In 1867, L. Koch described Liocranum ochraceum from an
adult female collected on the Greek island of Corfu. In 1940,
Hadjissarantos described Talanites pallides from two type lo-
calities in the Attica region, not far from Athens: Penteli and
Fyli. Decades later, Chatzaki et al. (2002b) provided a modern
redescription of T. pallides, transfered the species to the genus
Anagraphis and added the Greek isle of Antikythera as a new
sampled locality. Lastly, Bosmans (2014) identied Liocranum
ochraceum as a senior synonym of Anagraphis pallida and pro-
posed their synonymy.
e authors of the present work could not locate the type
series of A. pallida. No specimen was found in the ZMUA
collection, and the series is presumed lost. As such, it is not
possible to either conrm or refute the validity of the syn-
onymy proposed by Bosmans (2014). However, the holoty-
pe of A. ochracea and the description provided by Chatzaki
et al. (2002b) for Talanites pallides do indeed share multiple
striking similarities. Bosmans (2014) also proposed the syn-
onymy of Anagraphis ochraceum with Macedoniella karamani
Drensky, 1935, a species originally described from Northern
Macedonia. Additional, more recent records of A. ochraceum
have been made in Turkey (Demircan & Topçu 2015) and
Sardinia (Caria et al. 2021). A single female record from
the Russian Caucasus (Ponomarev & Shmatko 2020) needs
further conrmation (Nentwig et al. 2022).
During this study, three distinct Italian populations of
Anagraphis ochracea were observed and recorded. One, the
rst, inside the metropolitan area of the capital city of Rome,
Lazio; one in the northeastern area surrounding Lecce, Apu-
lia; one near Buccheri (SR) on the island of Sicily (Fig. 2).
Numerous specimens were collected and identied, but more
than 40 dierent individuals were observed and photogra-
phed in the eld (Tab. 1).
Anagraphis ochracea is a small to medium sized spider di-
stinguished by its homogeneous cream-coloured appearance
Fig. 1: Anagraphis ochracea. a. habitus of adult female; b. habitus of adult male; c. preserved female under microscopic light; d. closeup of distinctive
abundance of long setae
86 L. Lenzini, F. Castellucci, M. Poso, A. Kulczycki, E. Simeon, G. Greco, A. Piccinini, C. M. Legittimo
and a complete absence of a dorsal pattern. A detailed descrip-
tion of the species was provided by Chatzaki et al. (2002b).
One distinctive feature that visually sets it apart from other
Italian gnaphosids is the thick layering of setae covering the
entire body. is confers to the spider a distinctive, velvety
look. Hair and setae are abundant over the entire opisthoso-
ma, especially in its frontal region, over all segments of legs
and pedipalps and on the prosoma where, in the ocular region
and on the chelicerae, long forward-projecting hairs can also
be discerned (Fig. 1d).
No signicant morphological dierences were observed
between the three Italian populations. Both genitalia and
Fig. 2: Characteristics of localities where Anagraphis ochracea was found in Italy. a. The three distinct, Italian populations identied and reported in the
present work; b. Temperature and precipitation at the three sampled localities
Tab. 1: A detailed list of the total number of specimens observed in the wild from 2009 to 2022, with the associated ant species. Abbreviations: LL, L.Len-
zini; CML, C. M. Legittimo; GM, G. Mascia; MP, M. Poso; GR, G. Romagna
N° Locality Date Obs. Ant species N° Locality Date Obs. Ant species
1 Roma, Caarella 20. Jan. 2009 LL --- 27 Roma, Caarella 08. Nov. 2020 LL Messor ibericus
2 Roma, Caarella 29. Nov. 2009 LL Messor ibericus 28 Roma, Caarella 21. Dec. 2020 LL Messor ibericus
3 Roma, Caarella 25. Feb. 2012 LL Messor ibericus 29 Roma, Caarella 14. Jan. 2021 LL Messor ibericus
4 Roma, Caarella 29. Mar. 2012 LL Messor ibericus 30 Roma, Caarella 25. Apr. 2021 LL Messor ibericus
5 Roma, Caarella 03. May 2012 LL Messor ibericus 31 Roma, Caarella 19. May 2021 LL Messor wasmanni
6 Roma, Caarella 14. Jun. 2012 LL Messor ibericus 32 Roma, Caarella 04. Oct. 2021 LL Messor ibericus
7 Roma, Caarella 29. Nov. 2012 LL Messor ibericus 33 Roma, Caarella 05. Oct. 2021 LL Messor ibericus
8 Roma, Ostia Antica 23. Dec. 2012 LL Messor ibericus 34 Roma, Malagrotta 18. Oct. 2020 LL Messor ibericus
9 Roma, Caarella 18. Mar. 2014 LL Messor ibericus 35 Roma, Tor Marancia 31. Oct. 2020 LL Messor ibericus
10 Fiumicino, Torrinpietra 28. Oct. 2017 LL Messor ibericus 36 Roma, Caarella 17. Mar. 2022 LL Messor ibericus
11 Roma, Caarella 12. Nov. 2017 LL Messor ibericus 37 Roma, Caarella 05. Apr. 2022 LL ---
12 Roma, Caarella 19. Nov. 2017 LL Messor ibericus 38 Roma, Caarella 08. Apr. 2022 LL Messor ibericus
13 Roma, Caarella 16. Mar. 2018 LL Messor wasmanni 39 Tivoli, Monte Catillo 30. Apr. 2022 LL Messor ibericus
14 Roma, Caarella 17. Mar. 2018 LL Messor ibericus 40 Lecce, Solicara Jun. 2018 GM ---
15 Roma, Caarella 18. Apr. 2018 LL Messor ibericus 41 Lecce, Solicara 24. Jun. 2018 MP Messor ibericus
16 Roma, Caarella 15. May 2018 LL Messor ibericus 42 Lecce, Solicara 16. Aug. 2018 MP Messor ibericus
17 Roma, Caarella 16. May 2018 LL Messor ibericus 43 Lecce, Solicara 16. Aug. 2018 MP Messor ibericus
18 Roma, Caarella 11. Jun. 2018 CML Messor ibericus 44 Lecce, Solicara Sept. 2018 MP Messor ibericus
19 Roma, Caarella 11. Jun. 2018 CML --- 45 Lecce, Solicara Mar. 2019 MP Messor ibericus
20 Roma, Malagrotta 09. Dec. 2018 LL Messor ibericus 46 Lecce, Solicara 12. Apr. 2019 MP Messor ibericus
21 Roma, Caarella 20. Feb. 2019 LL Messor ibericus 47 Lecce, Solicara 19. Apr. 2019 MP ---
22 Roma, Caarella 15. Jun. 2019 LL Messor ibericus 48 Lecce, Solicara 26. Apr. 2019 MP Messor ibericus
23 Roma, Caarella 23. Feb. 2020 LL Camponotus aethiops 49 Lecce, Solicara 26. Apr. 2019 MP Messor ibericus
24 Roma, Caarella 23. Feb. 2020 LL Messor ibericus 50 Lecce, Solicara 05. May 2019 MP Messor ibericus
25 Roma, Caarella 29. Sep. 2020 LL Messor ibericus 51 Lecce, Solicara 05. May 2019 MP Messor ibericus
26 Roma, Caarella 21. Oct. 2020 LL Messor ibericus 52 Buccheri (SR) 29. Sep. 2020 GR ---
New records for the myrmecophilous Anagraphis ochracea 87
size variability were found to be very consistent in adults,
with body length ranging between 6–9 mm in females and
5–6.5mm in males. e analysis of female epigynes and male
palpal organs revealed little to no intraspecic variation be-
tween the three sampled localities (Fig. 3).
All Italian specimens were found to morphologically
coincide with individuals of A. ochracea used for comparison,
collected in Greece (essaly, Central Macedonia, Samos) by
Prof. Maria Chatzaki (see list of materials). ey also pro-
ved to be perfectly consistent with the holotype of A. ochracea.
Conversely, and despite a supercial resemblance in general
appearance, all Italian specimens were found to possess very
dierent genitalia compared to individuals of the closely rela-
ted Anagraphis pallens Simon, 1893. e latter were collected
on the island of Karpathos and Rhodes and, once again, pro-
vided to us by Prof. Chatzaki.
Lastly, a morphological character worth mentioning that
seems to have been ignored until now, is the presence of di-
stinctly long, thin, often distally recurved setae on the dorsal
side of most leg segments (Fig. 4).
DNA barcode
We have analyzed an A. ochracea specimen collected in Lecce,
Solicara. Sanger sequencing produced a nal sequence of 693
base pairs relative to a fragment of gene locus COI (accession
number OP871103), which is the standard, DNA barcoding
region used for metazoans, established by the International
Barcode of Life Consortium (https://ibol.org). No frame
shifts or stop codons were identied following translation
of the nucleotide sequence to an amino acid sequence. e
resulting DNA barcode is the very rst to be made pub-
licly available for both A. ochracea and the genus Anagraphis.
Ecological notes and behavioural observation in the wild
populations in Lazio
e presence of A. ochracea in Italy and its myrmecophilic
lifestyle were rst recorded by Luigi Lenzini starting from
November 2009. Specimens were found in the Appian Way
Regional Park (Rome), more specically in the section oc-
cupied by the Caarella Park (20m a.s.l) (Fig. 2a). e Caf-
farella Valley preserves the typical scenery of the Roman
countryside and is an integral portion of the vast Regional
Park. Agropastoral activities, such as extensive agriculture and
ovine breeding, survive to this day and coexist naturally with
the inhabitants of the surrounding neighbourhoods that use
the park for their own recreational activities. e valley is sub-
ject to a predominantly Mediterranean climate, and we report
mean annual temperature and rainfall (Fig. 2b). e areas of
Fig. 3: Anagraphis ochracea. a. epigyne (scale bar: 0.1 mm); b. male left palp in ventral view; c. male left palp in retrolateral view (scale bar: 0.2 mm)
Fig. 4: Anagraphis ochracea, long
setae on the dorsal side of legs.
a. patella and tibia, b. close-up of
long and distally recurved seta on
joint of patella/tibia
88 L. Lenzini, F. Castellucci, M. Poso, A. Kulczycki, E. Simeon, G. Greco, A. Piccinini, C. M. Legittimo
the valley where specimens of A. ochracea were observed and
collected are uncultivated pastures, grasslands and periodi-
cally mowed meadows, all with little to no tree cover.
e rst two specimens observed were found after lifting
a at rock that covered a Messor ibericus ant colony. e area
beneath the rock hosted several dozen worker ants and two
spiders were observed moving among them without alarming
the colony, never attempting to ee outside the area like most
other spiders would do and never being attacked by the alar-
med colony. Over the course of the following years, more than
30 similar observations were carried out in several other zo-
nes of the Caarella Park (Tab. 1). Following more targeted
searches, additional specimens were also found and observed
in more distant localities around the city of Rome: in Ostia
Antica (5m a.s.l) in December 2012, in Torrinpietra (60m
a.s.l), Fiumicino, in October 2017, in Malagrotta (20m a.s.l)
in December 2018 and October 2020 and, lastly, in Mount
Catillo (330m a.s.l.), Tivoli, in April 2022. ese additional
locations dier little in habitat and vegetation cover to the
areas of the Caarella Park where the rst specimens were
observed. Exceptions include Ostia Antica, where the patch
of land in which the specimen was found is used for horti-
culture and has higher average humidity levels, and Tivoli, a
location of higher altitude compared to the rest, characterized
by calcareous terrain and abundant rocky outcrops.
Over the course of 13 years, approximately 40 dierent,
adult and juvenile specimens were observed and photogra-
phed in Rome and its immediate surroundings. Almost all
individuals were found living amongst Messor ibericus colonies
(Fig. 5). Only three instances involved dierent ant species,
namely Messor wasmanni Krausse, 1910 in two occasions and
Camponotus aethiops (Latreille, 1798) in one.
Specimens were not always found alone, and on several
occasions, multiple A. ochracea were found living within the
same colony, even at dierent developmental stages. In one
instance, ve dierent individuals were found beneath the
same rock covering a Messor ibericus ant nest.
e dates in which specimens were observed or collected
are essentially random and follow no regular pattern. ey
were, rather, directly aected by the seasonal activity of ants
and by the periods of favourable conditions of the terrain.
Most observations were, thus, carried out in spring and in au-
tumn, when ant colonies are active, the vegetation of pastures
and grasslands in the urban park is contained and the rocks
that cover ant colonies are readily visible and easily accessible.
In most of the observations carried out in the Lazio re-
gion, A. ochracea was the sole arthropod species, besides the
ants, found beneath the rock. Only on rare occasions were
individuals found together with other myrmecophilous or
soil-dwelling arthropods. ese included orthopterans of the
genus Myrmecophilus, several dierent isopod species (see
Fig.5) and centipedes of the genus Lithobius.
Population in Apulia
e number of observations and the unique behavioural na-
ture of the ndings in central Italy sparked interest and lead
to targeted searches in other areas of the peninsula. Ultimate-
ly, between 2018 and 2019, Anagraphis ochreacea was found
also in southern Italy, specically in the region of Apulia,
near Solicara, north-east of city of Lecce (Fig. 2a). Several
specimens were observed and collected in a rural zone that
featured an abundance of Messor ibericus colonies. is area
is characterized by a dry Mediterranean steppe environment
and is dominated by xerophilous herbs and grasses. Several
plots of land have been dedicated to the cultivation of wheat
and sunower. e general lack of shrubbery, tree cover and
shade means that solar radiation is high and, accordingly, soil
temperatures reach extreme levels during the summer season
(Fig. 2b). e location where specimens were collected is a
dry area bordered by a 200-meter-long rock mound covered
in brambles and other vegetation. Here, numerous, large, well
developed Messor ibericus colonies were found coexisting with
several, smaller colonies of M. wasmanni.
In accordance with the observations carried out in cen-
tral Italy, and except for a single wandering adult male, all
specimens of A. ochreacea encountered in Apulia were found
within or in immediate proximity to Messor ibericus ant nests.
Most of these specimens were found dwelling unscathed
within the supercial area of the colony beneath the rocks,
surrounded by unalarmed M. ibericus workers. In accordance
with our previous observations from the Lazio region, several
specimens, disturbed by the removal of the overlaying rock,
were often seen quickly escaping inside the colony utilizing
the busy entrance tunnels. On one occasion, ten small, juve-
Fig. 5: Anagraphis ochracea. The vast majority of the specimens were found together with Messor ibericus ants. a. female together with worker ants in
Caarella Park (Rome), b. ditto
New records for the myrmecophilous Anagraphis ochracea 89
nile specimens were found resting together inside a 15 cm
deep chamber of the ant nest.
e frequency of the encounters in southern Italy was,
for the most part, directly correlated with the periods of ma-
ximum activity of M. ibericus. As a result, most individuals
were observed during spring and autumn when colonies reach
peak activity levels, while very few were found in summer and
winter, when surface activity of the ants is at its lowest. Du-
ring the hottest months of the year, when maximum daily
temperatures reach 38-39 °C, specimens of A. ochreacea were
observed only during the twilight hours (between 19.30 and
20.00). is is when the ants abandon the protection of the
thermally insulated depths and return to the supercial areas
of the nest, directly underneath the rocks. All sampling and
observations were carried out in dry conditions, never during
rainfall or with wet terrain. Finally, in Apulia A. ochreacea was
found coexisting on multiple occasions with soil dwelling iso-
pods and, dierently to what was observed in central Italy,
myrmecophilous Zygentoma species.
Observations in Sicily
e specimen from Sicily was collected under drastically
dierent environmental conditions. is single adult male,
presumably wandering in the search of females, was found,
surrounded by trapped dust, on the bathroom oor of a medi-
cal clinic. As a result, no observation on myrmecophilous be-
haviour and ecological information on the preferred natural
habitat can be provided for the Sicilian population. However,
the small town of Buccheri where the specimen was found,
is located at 820 m a.s.l on the northern slope of Mount
Lauro (986m a.s.l), in the province of Syracuse (Fig. 2a). e
general area is characterized by a semi-continental, temper-
ate climate. In the winter season, low clouds and foggy days
are frequent and weak to moderate frosts can occur (Fig. 2b).
e vegetation surrounding the town is mainly composed of
young, Mediterranean, coniferous woodlands planted in the
mid-1900s. At greater altitudes, around 920 m a.s.l, the trees
give way to open, barren clearings mostly used for pasture.
Behavioural observations in controlled conditions
All specimens from Caarella Valley observed in captivity
proved to be highly adaptable to the captive space provided,
showing no signs of erratic behaviour or stress. None ever at-
tempted to escape the enclosure by climbing the slippery glass
surfaces, nor did any ever try to dig a retreat in the substrate.
Spiders generally stayed idle and remained motionless on the
terrain, showing sudden bursts of movement only when dis-
turbed or when prey items were dropped in the enclosure.
Captive specimens were relatively long-lived, with a notable
adult female still thriving after 19 months.
During captive breeding, several dierent prey items were
oered to each spider to test the feeding habits of the spe-
cies. Messor ibericus workers were entirely ignored on every
occasion, and so were the larvae and pupae, suggesting that
A. ochracea does not prey on ants. Isopods belonging to both
Porcellionidae and the Platyarthridae, common in large ant
colonies, were also always neglected by the spider. On the
other hand, all specimens proved to be extremely responsive
towards several Zygentoma species (Lepismatidae - Tricho-
lepisma sp.), as well as other more generic species such as the
common ermobia domestica Packard, 1873. eir presence
always elicited a rapid reaction from the spider that promptly
pounced on the prey and rmly grasped it with all its legs.
None of the A. ochracea were ever observed utilizing silk dur-
ing prey capture.
e level of predatory responsiveness shown towards
lepismatids was quite remarkable when compared to other
prey items. Interestingly, spiders aggressively hunted every
lepismatid in the enclosure even when multiple specimens
were dropped inside at the same time. e spiders promptly
killed every specimen, and on many occasions did not even
attempt to eat the multiple deceased prey.
Ten years of acquired experience of captive breeding
have shown that A. ochracea is a spider that eats sporadically
and can tolerate long periods of time without food. Hunted
prey is often only partially consumed before being discarded.
Lepismatid prey items were generally oered once every one
or two months without ever observing a physical state of food
deprivation in any of the spiders.
Two adult females built multiple egg sacs during captive
breeding. e rst laid eggs in the enclosure shortly after cap-
ture. e second female, captured on 8. Nov. 2020, laid four
dierent egg sacs during captivity: one after 7 months (18.
Jun. 2021), another on 1. Aug. 2021, a third egg sac after 12
months (21. Oct. 2021) and a nal fourth after 18 months
(31. May 2022). e egg sac of A. ochracea is circular, 5mm
wide, dome-shaped, attened on one side, whitish in colour
(Fig. 6) and fastened to at surfaces. On numerous occasions
during eld sampling, identical egg sacs were observed at-
tached to the underside of the rocks that covered ant colonies
known to host specimens of A. ochracea.
At least one of these egg sacs was fertile and hatched
approximately one month after deposition (26. Aug. 2021).
ough a tiny exit hole was observed on their surface, the
hatching process was not immediately noticed as rst instar
juveniles are particularly small and can easily blend with the
substrate. Only one of the hatched spiderlings was spotted in
the enclosure (Fig. 6), so an average number of eggs laid by
this species cannot be reliably provided.
An articial ant nest of Messor ibericus was setup in an
attempt to replicate, under visible lab conditions, the peculiar
symbiosis observed in the wild between the two species. Due
Fig. 6: Anagraphis ochracea. Adult female with egg sac and a small juvenile
(arrow)
90 L. Lenzini, F. Castellucci, M. Poso, A. Kulczycki, E. Simeon, G. Greco, A. Piccinini, C. M. Legittimo
to the lack of any reliable scientic source in literature, no
specic protocols or methodologies were followed during the
setup of the formicarium. e authors attempted to ensure
that the most optimal conditions were provided to guarantee
reliable behavioural observations. A healthy and very popu-
lous, mature colony of approximately 2000 individuals was in-
troduced in the articial nest and fully settled over a period of
more than 12 months, occupying the overlaying arena and the
numerous galleries and secondary chambers of the system.
Following the complete establishment of the colony, and
during its most crowded phase, a single adult female of A.
ochracea was introduced into the formicarium. After three
full days of cohabitation, the spider was still unharmed and
was observed calmly wandering around the main arena and
the uppermost chambers, often walking directly on top of
the ants (Fig. 7). e ants themselves appeared to tolerate its
presence: workers would either briey probe the spider with
their antennae before continuing their work undisturbed or
casually walk alongside it ignoring its presence entirely. On
very rare occasions, some workers were observed trying to bite
one of its legs, an attempt that regularly failed when the spi-
der briey sprinted o a short distance away. ese sporadic
attempts by individual ants never propagated into a collective
alarmed or defensive state of the colony. ey may, however,
explain why on three separate occasions in the eld, speci-
mens were found to be either missing a limb or possessing a
freshly regenerated appendage. us, it is plausible to assume
that in certain rare instances, the spider may encounter brief
acts of hostilities by its ant symbionts.
Several days later, a second adult female was introduced
to the colony, once again in the arena of the formicarium. e
same interspecic interactions were observed: the spider mo-
ved freely within the colony and the ants appeared to tolerate
its presence entirely, carrying on with their regular activities
undisturbed and unalarmed. is second female peacefully
cohabited with the ants for one full day before being removed
from the nest.
After having observed and conrmed a clear degree of
mutual tolerance between A. ochracea and M. ibericus even
in lab conditions, several other non-myrmecophilic species
(Steatoda nobilis, Lycosoides coarctata, Scotophaeus blackwalli
and Filistata insidiatrix) of spiders were introduced to test
whether the same colony would react dierently. Shortly af-
ter their introduction to the formicarium, each one of these
species provoked clear, collective alarm and agitation, before
being promptly attacked, overwhelmed and cut to pieces by
the workers. is is not unusual as despite being primarily
granivores, M. ibericus will occasionally feed on live or dead
invertebrates such as earthworms, insect larvae and other arth-
ropods as well as defend its nest voraciously against intruders.
Myrmecophily of Anagraphis ochracea
Numerous, repeated observations carried out in the wild in
both Lazio and Apulia, clearly demonstrate the remarkable
habit of A. ochracea to live, undisturbed, within ant colonies.
Except for three isolated cases that involved Messor wasmanni
and Camponotus aethiops, the majority of the specimens were
found in association with Messor ibericus. Interestingly, when
disturbed by the removal of the overlying rock, rather than
running away from the alarmed ants like most other spider
species do, A. ochracea always remained among the startled
colony and often escaped directly inside the busy entrance
tunnels of the nest. Additional lab observations unequivocally
conrmed that A. ochracea is capable of blending within and
being accepted even by a newly encountered ant colony. is
is in stark contrast to what was observed with other spider
species instead, all of which were promptly overwhelmed by
workers and disposed of shortly after their introduction to the
formicarium.
Ant colonies can be considered complex ecosystems on
their own. Each ant fulls a specic role to help the colony
survive. Defence and protection are of utmost importance:
the worker caste will staunchly defend the colony from po-
tential predators, and any external intruder is generally con-
sidered a threat (Hölldobler & Wilson 1990). Nevertheless, a
great number of arthropod species, both arachnid and insect,
have developed ways to live unharmed within the colonies, in
a symbiotic relationship with ants. Ant nests, in fact, provide
a signicant number of ecological benets to would-be hosts,
including a relatively stable microclimate, a constant abun-
dance of food sources and a considerable level of protection
from their own natural predators (Cushing 1997, 2012, Witte
et al. 2008, Nelson & Jackson 2009). Evidently, for such a
risky evolutionary adaption to be successful, the benets must
Fig.7: Anagraphis ochracea. The female specimens used in the experiment moved freely among workers in the arena of the articial formicarium. a. adult
female surrounded by unalarmed Messor ibericus worker, b. ditto, in a debris-free area of the arena
New records for the myrmecophilous Anagraphis ochracea 91
greatly outweigh the drawbacks of living inside a thriving ant
colony (Ceccarelli 2013). To establish a stable symbiotic re-
lationship with ants, these arthropods, generically known as
myrmecophiles, have developed ways of penetrating the bio-
logical barriers of the colony in order to survive unharmed or
unseen. e means through which myrmecophiles manage to
blend in such a risky environment are varied and can dier
substantially between species, and may be of morphological,
behavioural, and chemical nature (Cushing 1997, 2012).
e observational scope of this study did not allow us
to condently arm through which mechanism A. ochracea
successfully manages to survive unscathed within M. iberi-
cus colonies. Whether it be through mechanical or chemical
processes is unclear, and further, more thorough analysis on
cuticular hydrocarbons of both species will be carried out in
the future.
Anagraphis ochracea is not a myrmecomorph, and lab ob-
servations indicate that it is in no way myrmecophagous eit-
her. us, it is easy to assume that it utilizes Messor ibericus
colonies to nd refuge, protection from external threats and
easy access to a wide range of varied, potential food sources.
In fact, observations in the eld have revealed that Italian M.
ibericus colonies harbour a wide range of dierent myrmeco-
philous arthropods. We recorded the following:
– Isopoda: Porcellionidae and Platyarthridae;
– Zygentoma: Lepismatidae (Tricholepisma and Neoaste-
rolepisma);
– Orthoptera: Myrmecophilus;
– Chilopoda: Lithobiidae (Lithobius romanus (Marzio Zap-
paroli pers. comm.))
– Coleoptera: Oochrotus e Cholovocera;
– Acari: Laelapidae and Scutacaridae (Massimo Plumari
pers. comm.)
– Collembola: Cyphoderidae
With the exception of lithobid centipedes, powerful pre-
dators on their own, and the minute sized Acari and Collem-
bola, which could possibly serve as prey items for the newly
hatched spiderlings, the more likely food source for adult
A.ochracea include myrmecophilous Zygentoma and Ortho-
ptera.
Testing in captivity showed that when introduced to myr-
mecophilous lepismatids, A. ochracea always reacted very ag-
gressively, promptly subduing the prey even when multiple
Zygentoma were oered at the same time. No experiments
were carried out utilizing orthopterans of the genus Myrme-
cophilus.
Years of experience in the captive breeding of this species
have indicated that A. ochracea is not a voracious feeder and
possibly possesses a slower metabolism compared to other
Gnaphosidae. It appears to be a slow-growing species capa-
ble of living multiple years. e fact that both small juveniles
and adults of both sexes were observed living inside healthy,
thriving M. ibericus colonies leads us to believe that A. ochracea
likely completes its entire life cycle in symbiosis with ants.
Observations carried out in the lab also suggest that adult
females lay multiple egg sacs and can preserve male sperm for
several months after mating.
Conclusions
e present work provides the rst records of Anagraphis
ochracea for both continental Italy and the island of Sicily,
with two distinct populations found in the former. Observa-
tions in the eld and experimentation in captivity unequivo-
cally conrm that this species has a myrmecophilous lifestyle
and lives as a symbiont inside colonies of Messor ibericus. In
the wild, specimens of A. ochracea were commonly observed
calmly resting even while surrounded by dozens of alerted
worker ants and were often seen escaping directly inside the
crowded tunnels of the alarmed colony. e use of an articial
formicarium of M. ibericus further conrmed that, as opposed
to other spider species tested, A. ochracea is readily accepted
by the ants and will survive unscathed for days. In conclu-
sion, this species can live undisturbed and unalarmed inside
M. ibericus colonies and does so possibly using some form of
chemical mimicry.
As a result, we demonstrate for the very rst time that, at
least in the areas we studied, this species lives as a myrmeco-
phile inside ant colonies. However, we cannot yet determine
and describe which mechanism is in play during this symbi-
osis, be it mechanical, chemical or of other nature. Further
research on the cuticular hydrocarbons of both symbionts will
be carried out by the authors. Ultimately, the present work
shows how a targeted sampling of obscure, overlooked, hardly
accessible and often neglected microhabitats, such as an ant
colony, can lead to interesting new discoveries from a biogeo-
graphical, behavioural, and ecological point of view.
Acknowledgements
We wish to extend our sincere gratitude to Valerio Dolci for kindly
providing us with both his Messor ibericus formicarium and his sugge-
stions regarding the behaviour of this ant species; Giovanni Bertazzoli
for his unparalleled experience with Messor spp. colonies; Mattia
Menchetti for conrming the identications of all the ant species
collected in the two main localities; Janet Beccaloni of the Natural
History Museum, London, for having kindly provided the holotype
of A. ochracea and for her renowned kindness and availability; Christos
Georgiadis and Valakos of the Zoological Museum of the University
of Athens for their assistance during our search for the holotype of
A. pallida; Maria Chatzaki from Crete for having provided numerous
specimens used for comparison; Giorgio Romagna for having collected
and sent the specimen from Sicily; Giuseppe Mascia for his help
during eld work. Finally, we would like to thank the two reviewers
for their helpful suggestions. is work was conducted and nanced
by Aracnolia – Italian Arachnological Society (www.aracnolia.org).
Authors’ contributions Conceptualization: LL, CML. Field work:
2009-2018 LL; 2018-2022 LL, MP. Taxonomic and morphological
study: CML, AK, ES. Behavioural observation in the Laboratory:
LL. DNA Barcoding: FC, AP. Proof reading: LL, FC, MP, AK, ES,
GG, AP, CML. Writing: CML, LL, FC, AK.
References
Bosmans R 2014 On the identity of the genera Anagraphis Simon,
1893 and Macedoniella Drensky, 1935 with two new synonyms
(Araneae: Gnaphosidae). – Arachnologische Mitteilungen 48:
38-41 – doi: 10.5431/aramit4807
Caria M, Pantini P, Alamanni F, Ancona C, Cillo D & Bazzato E
2021 New records and interesting data for the Sardinian spider
fauna (Arachnida: Araneae). – Fragmenta Entomologica 53: 321-
331 – doi: 10.13133/2284-4880/555
Castellucci F, Schifani E, Luchetti A, Schar N 2022 New asso-
ciation between red wood ant species (Formica rufa group) and
the myrmecophilic spiders Mastigusa arietina and yreosthenius
biovatus. – Bulletin of Insectology 75: 231-238
Ceccarelli FS 2013 Ant-mimicking spiders: strategies for living with
social insects. – Psyche: A Journal of Entomology 2013 (839181):
1-6 – doi: 10.1155/2013/839181
92 L. Lenzini, F. Castellucci, M. Poso, A. Kulczycki, E. Simeon, G. Greco, A. Piccinini, C. M. Legittimo
Chatzaki M, aler K & Mylonas M 2002a Ground spiders (Gna-
phosidae; Araneae) of Crete (Greece). Taxonomy and distribution.
I. – Revue Suisse de Zoologie 109: 559-601 – doi: 10.5962/bhl.
part.79611
Chatzaki M, aler K & Mylonas M 2002b Ground spiders (Gnapho-
sidae, Araneae) of Crete and adjacent areas of Greece. Taxonomy
and distribution. II. – Revue Suisse de Zoologie 109: 603-633 – doi:
10.5962/bhl.part.79612
Cushing PE 1997 Myrmecomorphy and myrmecophily in spi-
ders: a review. – Florida Entomologist 80: 165-193 – doi:
10.2307/3495552
Cushing PE 2012 Spider-ant associations: an updated review of
myrmecomorphy, myrmecophily, and myrmecophagy in spiders.
– Psyche: A Journal of Entomology 2012 (151989): 1-23 – doi:
10.1155/2012/151989
Demircan N & Topçu A 2015 A contribution to the spider fauna
of the European part of Turkey (Araneae). – Serket 14: 176-183
Donisthorpe H 1927 e guests of British ants, their habits and life
histories. Routledge and Sons, London, UK. 244 pp.
Folmer O, Black M, Hoeh W, Lutz R & Vrijenhoek R 1994 DNA
primers for amplication of mitochondrial cytochrome c oxidase
subunit I from diverse metazoan invertebrates. – Molecular Marine
Biology and Biotechnology 3: 294-299
Hadjissarantos H 1940 Les araignées de l’Attique. Athens. 132 pp.
Hölldobler B & Wilson EO 1990 e ants. Harvard University Press,
Cambridge/Mass. 746 pp.
Koch L 1867 Zur Arachniden und Myriapoden-Fauna Süd-Europas.
– Verhandlungen der Kaiserlich-Königlichen Zoologisch-Bota-
nischen Gesellschaft in Wien 17: 857-900
Mendonça CAF, Pesquero MA, Carvalho RDSD & Arruda FV de
2019 Myrmecophily and Myrmecophagy of Attacobius lavape
(Araneae: Corinnidae) on Solenopsis saevissima (Hymenoptera:
Myrmicinae). – Sociobiology 66: 545-550 – doi: 10.13102/socio-
biology.v66i4.4431
Nelson XJ & Jackson RR 2009 e inuence of ants on the ma-
ting strategy of a myrmecophilic jumping spider (Araneae,
Salticidae). – Journal of Natural History 43: 713–735. – doi:
10.1080/00222930802610469
Nentwig W, Blick T, Bosmans R, Gloor D, Hänggi A & Kropf C
2022 Spiders of Europe. Version 3.2022. – Internet: https://www.
araneae.nmbe.ch (1. Mar. 2022) – doi: 10.24436/1
Pantini P & Isaia M 2019 Araneae.it: the online Catalog of Italian
spiders with addenda on other Arachnid Orders occurring in Italy
(Arachnida: Araneae, Opiliones, Palpigradi, Pseudoscorpionida,
Scorpiones, Solifugae). – Fragmenta Entomologica 51: 127-152
– Internet: www.araneae.it (1. Mar 2022)
Parmentier T, Dekoninck W & Wenseleers T 2017 Arthropods as-
sociate with their red wood ant host without matching nestmate
recognition cues. – Journal of Chemical Ecology 43: 644–661 – doi:
10.1007/s10886-017-0868-2
Ponomarev AV & Shmatko VY 2020 New species and new records of
spiders (Aranei) in the south of Russia. – Caucasian Entomological
Bulletin 16: 299–309 – doi: 10.23885/181433262020162-299309
Wheeler WC, Coddington JA, Crowley LM, Dimitrov D, Golobo
PA, Griswold CE, Hormiga G, Prendini L, Ramírez MJ, Sierwald
P, Almeida‐Silva L, Alvarez‐Padilla F, Arnedo MA, Benavides
Silva LR, Benjamin SP, Bond JE, Grismado CJ, Hasan E, He-
din M, Izquierdo MA, Labarque FM, Ledford J, Lopardo L,
Maddison WP, Miller JA, Piacentini LN, Platnick NI, Polotow
D, Silva‐Dávila D, Schar N, Szűts T, Ubick D, Vink CJ, Wood
HM & Zhang J 2017 e spider tree of life: phylogeny of Araneae
based on target‐gene analyses from an extensive taxon sampling.
– Cladistics 33: 574–616 – doi: 10.1111/cla.12182
Witte V, Leingärtner A, Sabaß L, Hashim R & Foitzik S 2008 Sym-
biont microcosm in an ant society and the diversity of interspecic
interactions. – Animal Behaviour 76: 1477–1486 – doi: 10.1016/j.
anbehav.2008.05.010
World Spider Catalog 2022 World spider catalog. Version 23.0.
Natural History Museum Bern. – Internet: http://wsc.nmbe.ch
(1. Mar 2022) – doi: 10.24436/2
Zhang J & Madden TL 1997 PowerBLAST: A new network BLAST
application for interactive or automated sequence analysis and an-
notation. – Genome Research 7: 649–656 – doi: 10.1101/gr.7.6.649