ArticlePDF Available

A revision of Herennia (Araneae: Nephilidae: Nephilinae), the Australasian 'coin spiders'

Authors:

Abstract and Figures

The nephilid 'coin spiders' (Herennia Thorell) are known for their arboricolous ladder webs, extreme sexual size dimorphism and peculiar sexual biology. This paper revises Herennia taxonomy, systematics, biology and biogeography. The widespread Asian Herennia multipuncta (Doleschall) (= H. sampitana Karsch, new synonymy; = H. mollis Thorell, new synonymy) is synanthropic and invasive, whereas the other 10 species are narrowly distributed Australasian island endemics: H. agnarssoni, sp. nov. is known from Solomon Islands; H. deelemanae, sp. nov. from northern Borneo; H. etruscilla, sp. nov. from Java; H. gagamba, sp. nov. from the Philippines; H. jernej, sp. nov. from Sumatra; H. milleri, sp. nov. from New Britain; H. oz, sp. nov. from Australia; H. papuana Thorell from New Guinea; H. sonja, sp. nov. from Kalimantan and Sulawesi; and H. tone, sp. nov. from the Philippines. A phylogenetic analysis of seven species of Herennia, six nephilid species and 15 outgroup taxa scored for 190 morphological and behavioural characters resulted in 10 equally parsimonious trees supporting the monophyly of Nephilidae, Herennia, Nephila, Nephilengys and Clitaetra, but the sister-clade to the nephilids is ambiguous. Coin spiders do not fit well established biogeographic lines (Wallace, Huxley) dividing Asian and Australian biotas, but the newly drawn 'Herennia line' suggests an all-Australasian speciation in Herennia. To explain the peculiar male sexual behaviour (palpal mutilation and severance) known in Herennia and Nephilengys, three specific hypotheses based on morphological and behavioural data are proposed: (1) broken embolic conductors function as mating plugs; (2) bulb severance following mutilation is advantageous for the male to avoid hemolymph leakage; and (3) the eunuch protects his parental investment by fighting off rival males.
Content may be subject to copyright.
Introduction
The spider family Nephilidae Simon, 1894, the subject of
numerous biological studies, now contains four genera
according to new phylogenetic analyses (Kuntner, in press;
M. Kuntner, G. Hormiga and J. A. Coddington, unpublished
data): the well known pantropical Nephila Leach, 1815 and
Nephilengys L. Koch, 1872, the lesser known African–
Oriental Clitaetra Simon, 1889 and the Australasian
(sub)tropical genus Herennia Thorell, 1877. Nephila and
Nephilengys are being revised (M. Kuntner unpublished
data; Australasian Nephila also by M. S. Harvey, A. D. Austin
and M. Adams, unpublished data), Clitaetra contains six
species and is sister to all other nephilids (Kuntner, in press)
and Herennia is the subject of this revision. Herennia species
exhibit extreme sexual, in particular size, dimorphism and
ornate appearance (Figs 1, 14, 25–26) as well as fascinating
web architecture, which typically is in the form of a ladder
web closely applied to tree trunks (Figs 14, 15, 25–26). The
sexual behaviour of Herennia is extraordinary. Males break
off parts of their pedipalps and plug female epigyna with
embolic parts (embolus + embolic conductor) (Figs 10EF,
11, 17A, 18C, 22EF). After mating, the males regularly
sever their entire palpal bulb (Fig. 23AE) and continue to
occupy the female web as ‘eunuchs’ (Robinson and
Robinson 1980).
Despite its interesting biology, Herennia has been poorly
studied systematically, behaviourally and morphologically.
Platnick (2005) lists four species of Herennia worldwide:
H. ornatissima (Doleschall, 1859), the type of the genus
ranging from India to China, Malaysia and New Guinea;
H. mollis Thorell, 1887 from Myanmar; H. papuana Thorell,
1881 from New Guinea; and H. sampitana Karsch, 1880 from
Borneo. Of these, after this revision H. papuana remains
valid, H. ornatissima is revalidated and treated as a junior
synonym of H. multipuncta, H. mollis and H. sampitana are
synonyms and nine new species are described. The taxonomy
has proven to be problematic, mainly as a result of scarce
museum material (males with intact pedipalps are rare).
A non-scientific name ‘coin spiders’ seems appropriate for
Herennia, from the Latin meaning of a family of coins.
Biological observations exist for the tropical Asian
H. multipuncta, H. papuana Thorell from Papua New Guinea
Invertebrate Systematics, 2005, 19, 391–436
10.1071/IS05024 1445-5226/05/050391© CSIRO 2005
Matjaž Kuntner
Department of Entomology, National Museum of Natural History, Smithsonian Institution, NHB-105,
PO Box 37012, Washington, DC 20013-7012, USA and Department of Biological Sciences,
George Washington University, 2023 G St. N.W., Washington, DC 20052, USA.
Present address: Institute of Biology, Scientific Research Centre of the Slovenian Academy of Sciences and Arts,
Novi trg 2, PO Box 306, SI-1001 Ljubljana, Slovenia. Email: kuntner@gmail.com
Abstract. The nephilid ‘coin spiders’ (Herennia Thorell) are known for their arboricolous ladder webs, extreme
sexual size dimorphism and peculiar sexual biology. This paper revises Herennia taxonomy, systematics, biology
and biogeography. The widespread Asian Herennia multipuncta (Doleschall) ( = H. sampitana Karsch, new
synonymy; = H. mollis Thorell, new synonymy) is synanthropic and invasive, whereas the other 10 species are
narrowly distributed Australasian island endemics: H. agnarssoni, sp. nov. is known from Solomon Islands;
H. deelemanae, sp. nov. from northern Borneo; H. etruscilla, sp. nov. from Java; H. gagamba, sp. nov. from the
Philippines; H. jernej, sp. nov. from Sumatra; H. milleri, sp. nov. from New Britain; H. oz, sp. nov. from Australia;
H. papuana Thorell from New Guinea; H. sonja, sp. nov. from Kalimantan and Sulawesi; and H. tone, sp. nov. from
the Philippines. A phylogenetic analysis of seven species of Herennia, six nephilid species and 15 outgroup taxa
scored for 190 morphological and behavioural characters resulted in 10 equally parsimonious trees supporting the
monophyly of Nephilidae, Herennia, Nephila, Nephilengys and Clitaetra, but the sister-clade to the nephilids is
ambiguous. Coin spiders do not fit well established biogeographic lines (Wallace, Huxley) dividing Asian and
Australian biotas, but the newly drawn ‘Herennia line’ suggests an all-Australasian speciation in Herennia. To
explain the peculiar male sexual behaviour (palpal mutilation and severance) known in Herennia and Nephilengys,
three specific hypotheses based on morphological and behavioural data are proposed: (1) broken embolic
conductors function as mating plugs; (2) bulb severance following mutilation is advantageous for the male to avoid
hemolymph leakage; and (3) the eunuch protects his parental investment by fighting off rival males.
A revision of Herennia (Araneae:Nephilidae:Nephilinae),
the Australasian ‘coin spiders’
www.publish.csiro.au/journals/is
CSIRO PUBLISHING
M. Kuntner392 Invertebrate Systematics
and H. etruscilla, sp. nov. from Java. Natural history of the
remaining species (in most cases known from only one or a
few individuals) is unknown. Herennia multipuncta appears
native in South Asia and Indochina (these areas apparently
lacking other sympatric Herennia), but also occurs in
Sumatra, Java, Borneo, Sulawesi, the Philippines and the
Moluccas (Ambon). These latter islands are known or pre-
dicted to also harbour narrowly endemic Herennia species.
Although the currently available taxonomic, biogeo-
graphic and ecological data are preliminary, Herennia species
appear to be very narrow endemics and are rare (judging from
availability in museum collections). They are obligate arbori-
coles (see Robinson and Lubin 1979), and appear to be con-
fined to pristine (rain) forests. A notable exception is the
widespread and common Herennia multipuncta (Doleschall),
which is synanthropic, appears to be invasive and could
potentially benefit from human induced habitat destruction.
Herennia deelemanae, sp. nov. is known from only two indi-
viduals from Sabah (Malaysian Borneo). The holotype male
(Fig. 17BC) was discovered as a singleton in extensive
samples of a recent primary rainforest canopy-fogging inven-
tory (evidence of potential rarity in a threatened habitat).
A presumed conspecific female (Fig. 17A) was collected as a
singleton in remote southern Sabah. More canopy-fogging
spider samples are available from Borneo secondary forests,
but they only contain the common H. multipuncta
(Ch. Deeleman, personal communication).
The group Nephilinae Simon, 1894, currently catalogued
in Tetragnathidae (Platnick 2005), has received considerable
systematic attention resulting in a range of classification
schemes (Kuntner 2002, 2003, in press; Kuntner and Hormiga
2002). Hypothesised placements for Nephilinae, Herennia
included, have been in Araneidae (Dahl 1912; Eberhard 1982;
Wunderlich 1986, 2004; Kuntner 2003; Pan et al. 2004) and
in Tetragnathidae (Levi 1980, 1986; Levi and von Eickstedt
1989; Coddington 1990; Hormiga et al. 1995, 2000; Scharff
and Coddington 1997; Griswold et al. 1998; Kuntner 2002;
Kuntner and Hormiga 2002). In the context of Clitaetra and
Nephilengys revisions, Kuntner (in press; unpublished data)
hypothesises the clade (Clitaetra +(Herennia +(Nephila +
Nephilengys))) to represent Nephilidae because the group
falls into neither of the above families. Such placement is also
tested here via phylogenetic analysis of most Herennia
species and selected nephilid and non-nephilid outgroups.
The first species-level phylogeny of Herennia allows prelim-
inary interpretations of its phylogeography.
Materials and methods
Morphological examination
Most morphological observations and illustrations of external struc-
tures were made using a Leica MZ APO dissecting microscope
(www.leica-microsystems.com, verified November 2005) with a
camera lucida. For internal palpal and epigynal anatomy a Leica
DMRM compound microscope with a camera lucida was used.
Microscope images were taken using a Nikon DXM 1200 digital
camera (www.nikon.com, verified November 2005), and assembled
using the Syncroscopy Automontage software (www.syncroscopy.com,
verified November 2005). Digital scanning electron microscope (SEM)
photographs were taken on a Leo 1430VP scanning electron micro-
scope (www.mwrn.com/leo/leo.htm, verified November 2005) at the
Department of Biological Sciences of the George Washington
University. For SEM preparation, specimens were cleaned ultrasoni-
cally for one minute, transferred to 100% ethanol overnight, dissected,
submitted to critical point drying, mounted on rivets using glue and
copper wire, then sputter coated.
Female genitalia and male pedipalps, if dissected, were excised
using scalpels and needles. Male palpal anatomy (trajectory of the
ducts) and internal female genitalic structure were examined by clear-
ing the organs in methyl salicylate (Holm 1979) and mounting them on
a temporary slide (Coddington 1983) and were illustrated under the
compound microscope. In spiders with large and heavily sclerotised
internal genitalia, like nephilids, this technique often produces poor
results. Thus, in order to decipher the trajectory of the copulatory ducts,
the epigyna were exposed to concentrated KOH for the time needed to
completely digest any soft tissues. Cleared genitalia were mostly illus-
trated under a dissecting microscope positioned into the most informa-
tive views or further dissected to expose the details. Although this
technique sometimes makes the spermathecae translucent and eluci-
dates the trajectories within, it is often necessary to interpret the
relationships of the positions where the copulatory and fertilisation
ducts attach to the spermatheca lumen. Male pedipalps were expanded
by exposing them to concentrated KOH for up to an hour, followed by
immersion in distilled water; the process was then repeated as needed.
All measurements are in millimetres, taken using a micrometer eye-
piece. Prosoma and opisthosoma length and height were measured in
lateral view, the width in dorsal view; all measured at widest points
(prosoma height measured at head region). Eye widths are maximum
diameters of the eye lens and eye separations were measured between
lenses. Leg segments were measured in detached legs. Only joints distal
from the trochanter were measured, as the detached leg breaks between
the trochanter and the femur. Maximum lengths of the femur, patella,
tibia, metatarsus and tarsus are reported; the total of these is reported as
total leg length, which corrects for overestimation if all leg joints were
measured. The cheliceral formula consists of three numbers: pro-
marginal cheliceral teeth (PCT, Fig. 6D); retromarginal cheliceral teeth
(RCT, Fig. 6D); and numbers of denticles inbetween the margins in
cheliceral furrow (ChD, Fig. 20E).
Pencil illustrations via camera lucida were kept on acid-free
archival paper. Simple line illustrations were inked on Vellum film
(Denril Multi-Media Vellum, Borden & Riley Paper Co. Inc.,
www.bordenandriley.com, verified November 2005); shaded illustra-
tions were inked on fine-grained coquille board and further rendered
with black, soft Prismacolour pencils (Sanford Corporation, Oak
Brook, IL, USA, www.prismacolour.com, verified November 2005).
All non-digital artwork (photographic slides included) were scanned for
digital enhancement in Adobe Photoshop version 7.0 (Adobe,
http://www.adobe.com/, verified November 2005), where digital
microscope and SEM images were also further manipulated. All image
plates were assembled and labelled in Adobe Illustrator version 10.
The species listed in the Taxonomy section are in alphabetical
order after the type species; such order does not imply phylogenetic
relatedness.
The following anatomic abbreviations are used in text and figures:
AC aciniform gland spigot(s)
AG aggregate gland spigot(s)
ALE anterior lateral eyes
ALS anterior lateral spinneret
AME anterior median eyes
ATA apical tegular apophysis
Invertebrate Systematics 393
BH basal hematodocha
BL book lung(s)
CB cymbium
CD copulatory duct
ChD cheliceral denticles
CO copulatory opening
CY cylindrical gland spigot(s)
E embolus
EA embolic apophysis
EB embolus base
EC embolic conductor
ECF embolic conductor flap
ECG embolic conductor groove
EC-m embolic conductior – membranous part
EH embolus hook
EPC epigynal chamber
ES epigynal septum
ESA epigynal sclerotised arch
ETm embolus-tegulum membrane
F fundus
FD fertilisation duct
Fe femur
FL flagelliform gland spigot(s)
m membrane(ous)
MAP major ampullate gland spigot(s)
mAP minor ampullate gland spigot(s)
Me metatarsus
N nubbin
P paracymbium
Pa patella
PCT promarginal cheliceral teeth
PI piriform gland spigot(s)
PLE posterior lateral eyes
PLS posterior lateral spinneret
PME posterior median eyes
PMS posterior median spinneret
PPS palpal patellar seta(e)
PSL prosomal supracheliceral lobe
RCT retromarginal cheliceral teeth
S spermatheca
Sc scutum
SD sperm duct
ST subtegulum
SU sustentaculum
Ttegulum
Ta tarsus
TCp paired tarsal claw(s)
TCm median tarsal claw
Ti tibia
TO tarsal organ
Tr trichobothrium(a)
Specimen database
The nephilid specimen database was developed in BIOTA 1.6.0
(Colwell 1999). At least one specimen from each sample examined
received a unique specimen code (a required field in BIOTA), which
consisted of the two-letter genus code (‘he’ for Herennia) followed by
the sample number, a letter indicating gender or stage (‘f for a female,
‘m’ for a male, ‘j’ for a juvenile) and the consecutive number within the
sample. These specimen codes, matching those on specimen labels, are
listed in descriptions, behavioural observations and figure captions, in
order to facilitate museum voucher comparisons, if necessary. Other
specimen data in BIOTA included species identification and date,
gender, abundance, type status, museum depository (see museum
abbreviations below) and optional notes. The locality data entered in
BIOTA for each sample, if known, were: a unique locality code; local-
ity name; district; state/province; country; elevation; latitude and longi-
tude; and their accuracy (by convention, the word ‘approx. in this field
means the approximate coordinates were found later through maps or
gazetteers, not measured on the spot). If interpreted, those parts are
added, in square brackets, to the original spellings. The collection data
entered in BIOTA for each sample, if known, were: a unique collection
code; collected by; date collected; site; and source (‘label’ indicating
museum label and ‘notes’as personal notes from the field). The nephilid
specimen database, currently containing more than 4000 entries, is
available from the author. Upon publication of all four nephilid revi-
sions, the entire database will be made available electronically at
http://home.gwu.edu/~kuntner.
Museum abbreviations with names of curators and/or collection
managers are:
AMNH American Museum of Natural History, New York, USA;
Norman I. Platnick, Lou Sorkin.
BMNH British Museum of Natural History, London, UK; Janet
Beccaloni.
CAS California Academy of Sciences, San Francisco, USA;
Charles Griswold, Darrel Ubick.
CD Christa Deeleman-Reinhold’s collection, now in
RMNH, The Netherlands.
MCSNG Museo Civico di Storia Naturale, Genova, Italy;
Giuliano Doria.
MCZ Museum of Comparative Zoology, Harvard University,
Cambridge, USA; Gonzalo Giribet, Laura
Leibensperger.
MHNG Muséum d’Histoire Naturelle de la Ville de Genève,
Switzerland; Peter Schwendinger.
MNHN Muséum National d’Histoire Naturelle, Paris, France;
Christine Rollard.
NHMW Naturhistorisches Museum Wien, Vienna, Austria;
Juergen Gruber.
PMS Prirodoslovni Muzej Slovenije, Ljubljana, Slovenia;
Tomi Trilar.
QM Queensland Museum, Brisbane, Australia; Robert
Raven.
RMNH Rijksmuseum van Natuurlijke Historie, Leiden, The
Netherlands; Erik J. van Nieukerken, Kees van den Berg.
SMF Naturmuseum Senckenberg, Frankfurt, Germany; Peter
Jaeger.
USNM National Museum of Natural History (former United
States National Museum), Smithsonian Institution,
Washington, DC, USA; Jonathan A. Coddington, Scott
Larcher, Dana M. De Roche.
WA MWestern Australian Museum, Perth, Australia; Mark
Harvey.
ZMB Museum für Naturkunde der Humboldt-Universität zu
Berlin, Germany; Jason A. Dunlop.
ZMH Zoologisches Institut und Zoologisches Museum
Universität Hamburg, Germany; Hieronymus Dastych.
ZMUC Zoological Museum, University of Copenhagen,
Denmark; Nikolaj Scharff.
Phylogenetic analysis
Kuntner (in press) provided the first nephilid species-level phylo-
geny emphasising Clitaetra species. Nephilids were represented by all
six known Clitaetra species, a selection of seven Nephila species empha-
sising the Australasian species revised by M. S. Harvey, A. D. Austin and
M. Adams (unpublished data) (the type N. pilipes, N. antipodiana,
N. plumipes, N. edulis, N. clavipes, N. fenestrata, N. clavata), two
Nephilengys species (the type N. malabarensis, N. cruentata) and two
Revision of Herennia
M. Kuntner394 Invertebrate Systematics
Herennia species (the type H. multipuncta, H. papuana). Kuntner (in
press) corroborated the monophyly of all four nephilid genera. Thus, this
study uses only two species of each nephilid genus except Herennia:
Nephila pilipes (type species) and N. fenestrata (which is sister to all
other Nephila species in Kuntner, in press) representing Nephila;
Nephilengys malabarensis (type species) and N. cruentata representing
Nephilengys; Clitaetra episinoides (type species) and C. irenae (this
name is here disclaimed and remains unavailable for nomenclatorial pur-
poses; Kuntner, in press will provide the first description and diagnosis)
re-testing Clitaetra monophyly. All Herennia species known from
quality specimens are in this analysis, but Herennia oz, H. milleri,
H. sonja and H. jernej are omitted owing to limited material. All non-
nephilid outgroups from Kuntner’s (in press) analysis are included here:
the classical nephiline genera Phonognatha and Deliochus; the araneids
Araneus and two Argiope species; the tetragnathid genera Tetragnatha,
Meta and Leucauge; Linyphia (Linyphiidae); Pimoa (Pimoidae);
Steatoda (Theridiidae); Nesticus (Nesticidae); Epeirotypus (Theridio-
somatidae); Uloborus (Uloboridae); and Deinopis (Deinopidae). In
total, 28 taxa were included in this analysis.
Appendix 1 lists and describes the 190 characters and character
states used. The characters and their homologies, all parsimony infor-
mative, will be discussed in more detail elsewhere (M. Kuntner,
G. Hormiga and J. A. Coddington, unpublished data). The phylogenetic
matrix (Appendix 2) in NONA and NEXUS formats is available from
the author and will be submitted to TreeBASE (http://www.treebase.org/,
verified November 2005). The multistate characters were treated as
unordered (Fitch 1971) to avoid unnecessary assumptions of character-
state adjacency. I used NONA version 2.0 (Goloboff 1993) with para-
meters ‘hold 1000’, ‘mult*500’, ‘max*’, and ‘sswap’, under both
‘amb –’ and ‘amb = ‘ for the cladistic analysis and WinClada 1.00.08
(Nixon 2002) to display and manipulate trees and matrices for NONA.
Successive character weighting (Farris 1969) analysis was performed in
NONA with the command ‘run swt.run hold10000 hold/100 mult*100’
(using the macro swt.run). The bootstrap values (Felsenstein 1985)
were calculated in WinClada using default settings (100 replications,
‘mult*10’). Bremer support or decay index values (Bremer 1988, 1994)
were estimated in NONA using the commands ‘hold 10000’and ‘bs10’.
All trees are output from WinClada and their format does not imply
non-monophyly of the group Deinopoidea represented here by the two
primary outgroups, Deinopis and Uloborus.
Classification
I use a combination of Linnean (ICZN 1999) and phylogenetic nomen-
clature (Cantino and de Queiroz 2004). A review of the debate with a
rationale of such combined classification is in preparation (see also
Kuntner, in press). Clade names in this study are consistent with zoo-
logical ranks up to the family level, but are precisely circumscribed fol-
lowing the PhyloCode (PC) articles 7, 9–11 (Cantino and de Queiroz
2004). Phylogenetic definitions (PC Article 9, Note 9.4.1) used here are
node-based where ‘clade (A and B)’ means the least inclusive clade
containing A and B. Because the PhyloCode has not taken effect and
registration database has not been implemented, the names proposed
here are not registered (Article 8).
Biology
The review of Herennia biology (see Discussion) is based on published
accounts of H. papuana (as ornatissima in Robinson 1975, 1982;
Robinson and Robinson 1978, 1980; Robinson and Lubin 1979) and
H. multipuncta (as ornatissima in Masumoto and Okuma 1995;
Murphy and Murphy 2000: 388, figs 12.1–12.2), unpublished data on
H. multipuncta from Thailand (G. Hormiga, personal communication),
as well as personal observations of H. multipuncta in Sri Lanka (1993)
and H. etruscilla in Java (1996). Biology of all other species is
unknown. The synthesis of ‘Herennia ornatissima behaviours by
Eberhard (1982) is based on the above-cited papers for H. papuana.
Additional works on Herennia include anatomy (Roth and Roth 1984),
morphological phylogenetics (Coddington 1990; Hormiga et al. 1995)
and evolution of sexual size dimorphism (Coddington et al. 1997;
Hormiga et al. 2000). Additionally, some behaviours are described here
for the first time.
Taxonomy
Family NEPHILIDAE Simon
Subfamily NEPHILINAE Simon
Genus Herennia Thorell
Coin spiders
Epeira (Argyopes) Doleschall, 1859: 32 (description of E. multi-
puncta and E. ornatissima). – Stoliczka 1869: 236 (description
of E. mammillaris).
Herennia Thorell, 1877: 370 (type species, by original designation,
Epeira multipuncta Doleschall).
Herennia Karsch, 1880: 381 (description of H. sampitana). –
Thorell, 1878: 293; 1881: 77 (description of H. papuana); 1887:
166 (description of H. mollis); 1890; Simon, 1894: 759, figs 828,
835; Roewer, 1942: 925; Bonnet, 1957: 2159; Brignoli, 1983:
241; Platnick, 1989: 306; 1993: 373; 1997: 454; 2005; Murphy
& Murphy, 2000: 388.
Diagnosis
Herennia differs from all non-nephilid spiders by the striated
cheliceral boss in both sexes (Figs 6B, 20AC). Herennia
females differ from all other nephilid genera by the warty
carapace (Figs 5B, DF, 19), wider than long sternum
(except in H. tone and H. gagamba), lobed abdomen
(Figs 1A, CD, 8AC, 9AB, E, 10AD, 14, 16AB, 22AB,
25AB, 26AB, 27A, C, 28A, 29AB, DE, 31AB) and the
epigynum with distinct ventral chambers divided by a
septum (Figs 2AB, 10EF, 11, 16C, G, 17A, 22EF, 27B,
DE, 28BC, 29C, 31CD). Herennia males differ from all
other nephilid genera by the presence of a conspicuous apical
tegular apophysis (Figs 4A, E, 12AB, E, 13BE, 17B,
18AB, 30A, C) and an embolus hook (Figs 4CE, 18A,
30C). They further differ from males of Nephila and
Clitaetra by the large and complex sigmoidally shaped
embolic conductor with an extensive proximal membrane
(Figs 4AC, 12AE, 13, 18, 30, 17BC) and from males of
Nephilengys by the shape of distal embolic conductor part,
which has a flap (Figs 4AB, 11B, F, 12A, C, E, 13CE,
17–18) (absent only in H. papuana, Fig. 30). Webs of
Herennia are built tightly against tree trunks (Figs 14AB,
25, 26A), rocks or walls (Figs 14C, 15), such that the orb
plane follows the substrate shape. The hub-cup (Figs 14AB,
15, 25B), where the spider rests, touches the substrate. The
Herennia web has ‘pseudo radii’ (Robinson and Lubin
1979), which run parallel vertically through the orb
(Figs 25A, 26A) and do not run through the hub.
Invertebrate Systematics 395
Description
Female
General somatic morphology illustrated in H. multi-
puncta (Fig. 1). Live females are cryptic dorsally (Figs 14,
25, 26A), but colourful (red, orange) ventrally (Fig. 26B).
Prosoma. No carapace humps or spines (except very
short ones on top of warts, Fig. 19F). Head region low and
narrow, thoracic region wide (Figs 1AC, 5AC, 14, 19A, D,
20A). Carapace warty, edge ridged (Figs 5BF, 19). Carapace
covered with numerous white and thin, hair-like setae (Figs 5,
19). No tapeta in secondary eyes. Lateral eyes (Figs 1BC,
5AC, 19A, DE) not widely separated from medians. Lateral
eyes on a joint tubercle, slightly separated. PLE equal in size
(H. multipuncta, H. papuana) or larger than PME (H. tone, H.
agnarssoni, H. milleri). Cheliceral boss with numerous striae
(Figs 6B, 20AC). Sternum wider than long (or as wide as
long in H. tone). Sternal humps I to III conspicuous (Fig. 6E),
hump IV inconspicuous, unpaired anterior hump absent.
Cheliceral formula ranges from 3/3/25 (H. papuana) through
3/4/30 (H. milleri) to 4/4/15 (H. agnarssoni).
Appendages. Leg formula 1, 2, 4, 3, except in H. jernej
(1, 2 = 4, 3). Femoral spines I, II short, except in H. jernej
and H. tone. Femur I with a prolateral row of thin hair-like
setae (Figs 5B, D, 19AB, D). Sustentaculum present, hidden
within other setae (Fig. 7AD), or alternatively inconspicu-
ous or absent in H. etruscilla (Fig. 21C), H. papuana and
H. milleri. Prolateral tibiae with a paired lyriform organ
(Fig. 21AB).
Opisthosoma (Figs 1A, CD, 8–9, 10AD, 14, 16AB,
DE, 22AD, 25–26, 27AC, 28A, 29AB, 31AB) dorso-
ventrally flattened, laterally more or less lobed, posteriorly
truncated. Opisthosoma entirely covered with many rows of
large and small sclerotisations. Spinnerets uniformly of the
typical nephiline form (Figs 7EF, 21DF; observed in
H. multipuncta, H. etruscilla, H. papuana) (see also
Hormiga et al. 1995): ALS with ‘normal PI field’ where the
PI spigot base is nearly as long or longer than the shaft
(Griswold et al. 1998: ch. 69, fig. 48B), the major ampullate
spigot and a nubbin; PMS with a sparse aciniform field, and
a nubbin; PLS with the aggregate spigots embracing the flag-
elliform and with the two cylindrical spigots of normal size,
the mesal being peripheral.
Epigynum (Figs 2AB, 10EF, 11, 16C, G, 17A, 22EF,
27B, DE, 28BC, 29C, 31CD) with a pair of ventral
rounded chambers divided by a septum (may be inconspicu-
ous) and laterally positioned copulatory openings. Oval sper-
mathecae attached to the ventral body wall and juxtaposed or
barely separated, with gland pores all over their surface
(Figs 2C–D, 18D, 31E). Copulatory ducts massive, round and
heavily sclerotised, fertilisation ducts short. A heavily sclero-
tised medial arch present in the inner epigynum (Figs 2CD,
18D, 31E), which can extend laterally (Fig. 31E), corre-
sponding to external epigynal sclerotisations.
Male
General somatic morphology illustrated in H. multi-
puncta (Fig. 3) and H. etruscilla (Figs 23–24). Live males are
orange or red (Fig. 25A).
Prosoma (Fig. 3AC) pear-shaped, cephalic region low.
Prosoma highest in the middle. Labium wider than long, as
in Fig. 3D. Both eye rows slightly recurved, eyes roughly
equidistant. Large AME extending anteriorly over clypeus
(Figs 3, 23AE). Lateral eyes on a tubercle, not juxtaposed,
and not widely separated from the medians. Tapeta absent
from secondary eyes. Cheliceral boss striated.
Appendages (Fig. 23CD, F). legs long and slender,
with long spines on femora and tibiae. Leg formula 1,2,4,3.
Opisthosoma (Figs 3, 24) round, dorso-ventrally flat-
tened, with a dorsal scutum. Scutum with two pairs of
apodemes and numerous smaller sclerotisations. Venter with
inconspicuous rows of median and lateral sclerotisations.
Book lung covers not grooved, but with sculptured cuticle
(Fig. 24AB). PLS black, other spinnerets pale. One or two
pairs of white spots lateral to the spinnerets. Epiandrous
gland spigots in a row, roughly divided in two lateral groups
(Fig. 24CD).
Pedipalp (Figs 4, 12–13, 17–18, 30) with a short globular
cymbium, a rectangular paracymbium with an invagination,
a large flat or globular tegulum containing the sperm duct, an
apical tegular apophysis (Figs 4A, E, 12AB, E, 13BE, 17B,
18AB, 30A, C), a prominent and separate subtegulum, and
a massive sigmoidal-shaped embolic conductor enveloping
the embolus (see Diagnosis). The embolus and the embolic
conductor separated from the tegulum and from each other
by an extensive membrane (Fig. 30DE). A single macroseta
present on the distal part of the palpal patella. Embolus base
with an apophysis (Figs 4C, 30CE). Embolus (Figs 4CE,
30CE) long and fairly thin (but not filiform as in Nephila),
with a distal hook.
Monophyly
Herennia monophyly is supported by the following unam-
biguous synapomorphies (see phylogenetic results): ridged
carapace edge (character 3/state 1); carapace warts (8/1);
wider than long sternum (22/1); lobed abdomen (53/1); trun-
cated abdomen tip (55/1); abdominal sigillae (63/1); spotted
dorsum (66/1); lateral epigynal chamber openings (77/1);
apical tegular apophysis (129/1); hooked embolus (155/1);
and presence of pseudoradii (174/1). Ambiguous synapomor-
phies include also (ACCTRAN) short and stout femoral
macrosetae (40/1), oval spermathecae (87/2), rounded apical
paracymbium (121/0), flat embolus tip (157/0), the apparent
absence of body shake (scored only in H. papuana; 186/0) and
(DELTRAN) carapace edge with hair-like setae (4/1), pres-
ence of sternal tubercle IV (29/1), epigynal septum (78/1),
ridged embolic conductor edge (142/1), rectangular orb web
(160/1) and bulbus detachment or eunuch behaviour (185/1).
Revision of Herennia
M. Kuntner396 Invertebrate Systematics
Phylogenetic definition
The genus Herennia is defined as the least inclusive clade
containing the species H. multipuncta (the type), H. papuana,
H. etruscilla and H. deelemanae. This definition avoids the
other newly described Herennia species as specifiers because
they are currently known from females only.
Composition
The genus contains eleven species, of which nine are newly
described: H. agnarssoni, sp. nov. (Solomon Islands);
H. deelemanae, sp. nov. (Malaysia (Sabah)); H. etruscilla,
sp. nov. (Indonesia (Java)); H. gagamba, sp. nov. (Philippines
(Luzon)); H. jernej, sp. nov. (Indonesia (Sumatra));
H. milleri, sp. nov. (Papua New Guinea (New Britain, Rossel
Island)); H. multipuncta (Doleschall) (South and South-east
Asia); H. oz, sp. nov. (Australia (Northern Territory));
H. papuana Thorell (Papua New Guinea); H. sonja, sp. nov.
(Indonesia (Kalimantan, Sulawesi)); and H. tone, sp. nov.
(Philippines (Negros)).
Distribution
Tropical and subtropical Asia and Australasia (Fig. 32).
Taxonomic history
In 1877, Thorell described the new genus Herennia and des-
ignated Epeira multipuncta Doleschall as the type species.
He synonymised H. ornatissima with H. multipuncta and
described a female of this species from Sulawesi, which, he
claimed, was an intermediate form between the Javan
(Epeira multipuncta Dol.) and the Ambonese form
(E. ornatissima Dol.). Thorell (1877: 370) believed Herennia
to be close to the araneid Argiope.
Simon (1894: 757–759, figs 828, 835) erected the
nephiline group Herennieae to contain only Herennia. He
redescribed Herennia (1894: 759), erroneously noting
(contra Thorell 1877) that the type species was H. ornatis-
sima. Simon (1894: 758) agreed with Thorell (1877, 1887,
1890) that Epeira multipuncta Doleschall and E. ornatissima
Doleschall were two forms of the same species. However, he
ignoredThorell’s (1877, 1887) synonymisation of H. ornatis-
sima with H. multipuncta and simply noted (p. 758) that he
preferred the name ornatissima over multipuncta. Thorell
(1877), as the first reviser, designated H. multipuncta as type
species of Herennia.
Simon (1894: 759, figs 828, 835) also redescribed
H. ornatissima. However, the figure of the male pedipalp
(fig. 828) certainly depicts another species (close to
H. papuana) and the figure of the female (fig. 835) cannot
be associated with any species treated here.
Remarks
Simon’s misapplication of Herennia ornatissima versus
H. multipuncta has gone uncorrected in all spider catalogues
(Roewer 1942; Bonnet 1957; Platnick 1989, 1993, 1997,
2005) and has thus misled all Herennia nomenclature since.
Works on South and South-east Asian spiders (e.g. Tikader
1982: 106; Murphy and Murphy 2000: 388) and on spider
systematics (e.g. Hormiga et al. 1995) correctly identify the
species but misapply the name.
Other authors misidentified their organisms. Chrysanthus
(1971: 41) redescribed H. papuana (as H. ornatissima) from
New Guinea (the epigynum figure (fig. 86) was printed
upside down). Likewise, Robinson (1975, 1982), Robinson
and Robinson (1978, 1980) and Robinson and Lubin (1979)
worked on H. papuana, not H. ornatissima. Eberhard (1982)
also probably referred to H. papuana, not H. ornatissima.
In her illustrated guide to the genera of Australian orb-
weavers, Davies (1988: plate 18) provided illustrations of
unidentified Herennia species. The figures of the female
from Australia’s Northern Territory are H. oz, sp. nov. and
those of the male from Papua New Guinea are H. papuana.
However, although all figures faithfully represent the illus-
trated objects (both specimens examined here), the male
pedipalp figures are unfortunate in depicting an expanded
and damaged organ. Unexpanded pedipalps of H. papuana
exhibit a dramatically different morphology. Furthermore,
the labelling of the palpal sclerites, in the same plate, is
incorrect. The sclerite labelled ‘embolus’ is the broken-off
part of the sclerotised embolic conductor, and the sclerite
labelled ‘tegular apophysis’ is in fact the embolus, which had
been broken-off at the tip. Hormiga et al. (1995: fig. 5J)
redrew this damaged pedipalp from Davies (1988).
Etymology
Herennia is feminine in gender (Bonnet 1957: 2159).
However, Thorell’s (1877: 370) original explanation of ety-
mology reads: ‘Herennius, nom. propr. Latinum’. Perhaps
the name refers to the Roman Caesar Herennius Etruscus,
the son of Herennia Etruscilla. Searches on ‘Herennius’or
Herennia consistently point to old Roman coins of the
‘Family Herennia’. Perhaps these Roman coins, which may
resemble the flat and lobed abdomen of the spider Herennia
were the source of Thorell’s name derivation. The non-
scientific name, coin spiders, introduced here, follows this
presumed analogy. See also H. etruscilla etymology.
Key to the species of Herennia (females)
1. First femur with prolateral group of long spines (Figs 16AB,
27C, 31B). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
First femur without prolateral group of long spines. . . . . . . . . 4
2. Epigynum with a prominent anterior border (Fig. 27D);
Philippines. . . . . . . . . . . . . . . . . . . . . . H. gagamba, sp. nov.
Epigynum with a weak anterior border (Figs 16C, 31CD). . . 3
3. Epigynum with thick lateral sclerotised edge (Fig. 16C);
Solomon Islands . . . . . . . . . . . . . . . . H. agnarssoni, sp. nov.
Epigynum with thin lateral sclerotised edge (Fig. 31CD);
Philippines. . . . . . . . . . . . . . . . . . . . . . . . . . H. tone, sp. nov.
4. Carapace with a central yellow V-shaped mark on dark back-
ground (Figs 1C, 14AC, 27A, C). . . . . . . . . . . . . . . . . . . . 5
Invertebrate Systematics 397
Carapace without such mark. . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5. Epigynum with round chambers and thin septum (Figs 2AB,
27B). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Epigynum with oval chambers and strong septum (Fig. 27E);
Sumatra. . . . . . . . . . . . . . . . . . . . . . . . . . . H. jernej, sp. nov.
6. Sclerotised epigynal edge lateral and posterior to the chambers
extensive (Fig. 2AB); South and South-east Asia . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. multipuncta (Dol.)
Sclerotised epigynal edge lateral and posterior to the chambers
thin (Fig. 27B); Kalimantan, Sulawesi. . . . H. sonja, sp. nov.
7. Epigynum round, with weak septum (Fig. 17A); North Borneo . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. deelemanae, sp. nov.
Epigynum wide, with conspicuous septum (Figs 16G, 18C,
28BC, 29C). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Epigynum with oval chambers (Figs 18C, 28BC). . . . . . . . . 9
Epigynum with round chambers (Figs 16G, 29C). . . . . . . . . 10
9. Epigynum with thick lateral sclerotised edge (Fig. 18C); Java. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. etruscilla, sp. nov.
Epigynum with thin lateral sclerotised edge (Fig. 28BC);
Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. oz, sp. nov.
10. Epigynal septum thick (Fig. 29C); Papua New Guinea . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. papuana, Thorell
Epigynal septum thin (Fig. 16G); New Britain . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. milleri, sp. nov.
Key to the species of Herennia (males)
NB: Males of H. agnarssoni, sp. nov., H. gagamba, sp. nov., H. jernej,
sp. nov., H. milleri, sp. nov., H. oz, sp. nov., H. sonja, sp. nov., and
H. tone, sp. nov. are unknown.
1. EC thin (Fig. 30AB); Papua New Guinea. H. papuana, Thorell
EC broad, with a subdistal apophysis (Figs 4AB, 12–13,
17BC, 18AB). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. EC subdistal apophysis prong-like (Fig. 18AB); Java. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. etruscilla, sp. nov.
EC subdistal apophysis flap-like (Figs 4AB, 12–13, 17BC,
18AB). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Distal EC longer than wide (Figs 4AB, 12–13); South and
South-east Asia. . . . . . . . . . . . . . . . . . H. multipuncta (Dol.)
Distal EC wider than long (Fig. 17BC); North Borneo . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. deelemanae, sp. nov.
Herennia multipuncta (Doleschall)
(Figs 1–15)
Epeira (Argyopes) multipuncta Doleschall, 1859: 32, pl. 11, fig. 1,
description of female (from Java).
Epeira (Argyopes) ornatissima Doleschall, 1859: 32, pl. 1, fig. 5,
description of female (from Ambon).
Epeira (Argyopes) mammillaris Stoliczka, 1869: 236, pl. 20, fig. 12,
description of female (from Assam).
Herennia multipuncta Thorell, 1877: 370 (first reviser); 1890: 101.
Herennia sampitana Karsch, 1880: 381, description of female (from
Kalimantan). Syn. nov.
Herennia mollis Thorell, 1887: 166, description of female (from
Myanmar). Syn. nov.
Herennia ornatissima Simon, 1894: 759, figs 828, 835 (description
of male and female). – Roewer, 1942: 925; Bonnet, 1957: 2159,
2160; Tikader, 1982: 106, figs 202–204 (description of female);
Platnick, 1989: 306; 1993: 373; 1997: 454; 2005; Hormiga
et al., 1995: 334, fig. 11A-F; Murphy & Murphy, 2000: 388,
figs 12.1, 12.2.
Material examined
Type material. Type(s) of E. multipuncta do not exist, description
based on artwork, deposited in RMNH.
Presumed holotype female of E. ornatissima in RMNH labelled
‘coll. Doleschall Amboina Epeira ornatissima Dol. from Maluku,
Ambon Island [3°38S, 128°7E] — he077/f1, with numbers 5550 and
1159 on separate labels, examined.
Type(s) of E. mammillaris were not found.
Holotype female of H. sampitana from Sampit, Borneo
[Kalimantan, 2°31S, 112°57E] — he089/f1 in ZMB dry collection,
examined.
Type(s) of H. mollis were not found.
Additional material examined. China: Hainan Island, Sanya
[Yaxian], 18°16N, 109°29E, 5.vi. 1983 — he056/f1 ( USNM).
India: Ootacmund [Udagamandalam], 11°24N, 76°41E (coll.
Hampson) — he093/f1 ( BMNH); Assam, 4 mi W Kochugaon,
26°33N, 90°3E, 50 m, 17.x. 1961 (coll. Ross & Cavagnaro) —
he012/f1 ( CAS); Kerala, Periyar Lake, 9°31N, 77°12E, 900m,
23.iii. 1962 (coll. Ross & Cavagnaro) — he014/f1 (2 CAS),
he014/m1 ( CAS); Trivandrum, 8°30N, 76°56E, ii.1896 — he094/f1
(3 BMNH); Maharashtra, 20 mi N Bombay, 19°15N, 72°50E, 50 m,
21.i.1962 (coll. Roos & Cavagnaro) — he011/f1 ( CAS); Tamil Nadu,
Alagarkoil, 21 km NE Madurai, 10°5N, 78°13E, 27.xii.1989 (coll.
V& B Roth) — he018/f1 ( CAS), he018/j1 (Immature CAS),
he001/m1 ( CAS), he002/m1 ( CAS); 5 km W Rajapalayam,
Ayyanar Falls, 9°27N, 77°30E, 350 m, xi.1979 (coll. W Eberhard) —
he038/f1 ( MCZ), he046/m1 ( MCZ). Indonesia: Java [no loc.
data] — he060/f1 ( MNHN) — he064/f1 ( MNHN) — he067/f1
( MNHN); Java, Tjibodas [Cibodas], 6°43S, 107°2E, 1907 (coll.
T Barbour) — he035/f1 (MCZ); Jawa Barat, Buitenzorg [ = Bogor],
6°35S, 106°47E — he066/f1 ( MNHN) — he092/f1 (10 NHMW),
he092/m1 ( NHMW); (coll. J Barbour) — he033/f1 (3 MCZ),
he045/f1 (2 MCZ); 1909 (coll. O Bryant) — he034/f1 (7 MCZ);
(coll. Palmer & Bryant) — he020/f1 (3 MCZ); 800 ft, 6.iii.1909 (coll.
Palmer & Bryant) — he036/f1 ( MCZ); Propinsi Kalimantan Selatan,
Aranio Dist., Tiwingan Village, 3°33S, 115°2E, 100 m, 9.x.i1996
(coll. M Kuntner) — he048/f1 ( USNM); Sulawesi [no further data],
2°24S, 120°18E — he061/f1 (3MNHN); Nord Celebes [no further
data], 0°45N, 122°0E (coll. Putz) — he084/f1 ( ZMB); Sumatera
Udara, Deli [ = Labuhandeli], 3°44N, 98°40E — he103/f1
( BMNH); Sumatera Utara, Sibaulangit, 3°19N, 98°34E (coll.
L Jackhan) — he087/f1 ( ZMB); Sumatra, 11 km NW Bukittingi,
0°14S, 100°18E, 18.xii.1978 (coll. ES Ross) — he017/f1 (3CAS);
Fort de Kock [Bukittinggi], 0°18S, 100°22E, 920 m (coll. Jacobson)
— he091/f1 ( NHMW); xi.1913 (coll. E Jacobson) — he079/f1
( RMNH); ‘W Coast’, Kalung [Kalang], 1°53N, 98°38E, x.1913
(coll. E Jacobson) — he080/f1 ( RMNH); Kerinci NP, 2°2S, 101°6E,
800 m, nr. river, 30.vii.1988 (coll. S Djojosudharmo) — he076/f1
(CD); Gn. Leuser, Ketambe, 3°48N, 97°32E, camp, 5.ii.1985 (coll.
Suyono) — he071/f1 ( CD); trail 11.4; l. litter, 4.v.1986 (coll.
Suharto) — he072/f1 (2CD), he072/m1 (2 CD); trail 8.1, 8.i.1985
(coll. Sudkro & Sarman) — he075/f1 (CD); ‘Rimba, Panti’ [Panti],
0°21N, 100°3E, on rock outcrop, ii.1985 (coll. P Hillyard) — he108/f1
( BMNH), he108/m1 (2 BMNH); Sebisi, 26.i.1922 — he132/f1
( AMNH); Si Rambe [Sirambas], 0°48N, 99°31E (coll.
Ed Reimoser) — he044/f1 ( MCZ); [no loc. data], 0°0N, 101°0E —
he063/f1 ( MNHN); Sumatra?, ‘Wai Lima Z. Sum. Lampongs’,
xi.1921 (coll. Karny) — he131/f1 ( AMNH). Malaysia: Frasers Hill
[Bukit Fraser], Gap Rest House, 3°42N, 101°44E, wall, viii.1983
(coll. RH Kiew) — he106/f1 (2BMNH); 2 mi. E Gopeng, 4°27N,
101°9E, 100, 21.vi.1962 (coll. Ross & Cavagnaro) — he013/f1
( CAS); 3 mi. SE Ipoh, 4°34N, 101°6E, 50 m (coll. ES Ross) —
he003/m1 ( CAS); Malakka [ = Malayan Pen.] (coll. Jochan) —
he085/f1 ( ZMB); Penang [P. Island], 5°23N, 100°15E (coll.
Revision of Herennia
M. Kuntner398 Invertebrate Systematics
SHower) — he095/f1 ( BMNH); Perak: Senggong (?) [Lenggong],
5°7N, 100°49E, ii.1932 (coll. Burbon) — he068/f1 ( MNHN);
Penang, Tanjung Bungah, Sungei Siru, 3.xii.1963 (coll. HT Pagden) —
he107/f1 ( BMNH); ‘Malay’, Yohore, Mount Austin, 1951 (coll.
CW Franck) — he116/f1 ( ZMUC); Negeri Johor, Kota Tinggi,
1°43N, 103°54E, 19.iii.1948 — he111/f1 ( BMNH); Pahang,
Beserah, 3°51N, 103°22E, tree trunk, 20.x.1901 (coll. Robinson) —
he101/f1 ( BMNH), he101/m1 ( BMNH); Merapoh, Taman Negara,
Kuala Juram, 4°38N, 102°7E, 75–150 m, on webs over a white wall,
12.iii.1999 (coll. Trilar & al.) — he005/f1 (2 PMS), he005/j1 (3 Imm.
PMS), he005/m1 ( PMS); Sabah, Poring Hotsprings, 5°59N,
116°14E, 500, 2.iv.1998 (coll. Deeleman & Zborowski) — he070/m1
( CD). Myanmar: Bhamo, 24°15N, 97°13E (coll. L Fea) —
he120/f1 (ZMUC); 1885–1889 (coll. L Fea) — he081/f1 (ZMH);
Kalaw, 20°37N, 96°34E, 1308 m, large tree trunk, 11 Nov 1978 (coll.
ES Ross) — he016/j1 (5 Imm. CAS), he004/m1 ( CAS); Pegu,
Burmah, 17°19N, 96°29E, xi.1966 (coll. CH Carpenter) — he043/f1
(3 MCZ); Tenasserim, 12°5N, 98°55E (coll. Oates) — he097/f1
( BMNH); Tharrawaddy, 17°38N, 95°47E (coll. EW Oates) —
he096/m1 ( BMNH). Nepal: a few miles W of Beni, on Mayangdi
Khola, 28°205’N, 83°340’E, 914 m, tree trunks, 20.vii.1954 (coll.
KH Hyatt) — he100/f1 (2 BMNH), he100/m1 ( BMNH); Baglung,
Along Kali Gandaki River, 28°160’N, 83°365’E, 914 m, 12.vi.1954
(coll. KH Hyatt) — he099/f1 ( BMNH), he099/m1 ( BMNH).
Philippines: Leyte, Near Dulag, 10°57N, 125°1E, 30 m, 8.iv. 1945
(coll. LW Saylor) — he130/f1 ( AMNH); Philippine Island [no
further data] (coll. Ledyard) — he040/f1 (4 MCZ), he041/f1 (4
MCZ); Philippinen [no exact loc. data] — he105/f1 ( BMNH);
Luzon, Los Banos, 14°10N, 121°10E, vii.1909 (coll. Ledyard) —
he027/m1 ( MCZ); (coll. Ledyard) — he042/f1 (9 MCZ); [Los
Banos], Mt. Makiling, 14°89’N, 121°1124’E (coll. Baker) —
he028/f1 ( MCZ). Singapore: Peirce Reservoir, 1°22N, 103°48E,
tree trunk, edge of secondary forest (coll. Koh) — he006/f1 ( WAM);
Seletar Reservoir, 1°24N, 103°47E, 8 m, forest, 7.xi.1991 (coll.
Andersen et al.) — he118/f1 ( ZMUC), he119/f1 ( ZMUC);
Singapore, 1°19N, 103°50E (coll. HN Ridley) — he110/f1 (2
BMNH); Sungei Seletar, 1°24N, 103°53E (coll. Koh) — he007/m1
(USNM); Ubin Island, 1°24N, 103°58E, 20 m, shrubs along forest
edge and fields, 3.xi.1991 (coll. Andersen et al.) — he117/f1
( ZMUC). Sri lanka: Nähe [near] Aukana Buddha, xii.1981 (coll.
Schmidt) — he112/f1 ( SMF); [Ceylon], 7°27N, 80°44E —
he037/f1 (2 MCZ); Kandy, 7°17N, 80°37E — he069/f1 (25
MNHN), he069/m1 (20 MNHN); Dec 1925 (coll. G Fairchild) —
he039/f1 (2 MCZ); Central Prov, Kandy Dist, Kandy; St. Bridget’s
Guesthouse, 7°17N, 80°37E, around the house at night, 20–22.i.1995
(coll. Kuntner & Antauer) — he008/f1 (5 USNM); Peradeniya
Botanical Gardens, 7°15N, 80°36E — he086/f1 (3 ZMB); 22 Jan
1995 (coll. M Kuntner) — he010/f1 ( USNM); tree trunks, 22 Oct
1982 (coll. Wanless) — he109/f1 (4 BMNH); North-Central Prov,
Anuradhapura Dist, at Anuradhapura water tank near new bus station,
8°18N, 80°24E, taken from a web built against a tree trunk of a bo-tree
at night, 18 Jan 1995 (coll. M Kuntner) — he009/f1 ( USNM).
Taiwan: Kankan, Formosa, vii.1909 (coll. Santer) — he088/f1
(ZMB). Thailand: 20 mi SE Chantaburi, 12°24N, 102°19E, 75 m,
1.viii.1962 (coll. Ross & Cavagnaro) — he015/f1 (CAS); Khao Yai
National Park, 14°20N, 101°30E, savannah, 10 Nov 1987 (coll. CL &
PR Deeleman) — he074/f1 ( CD); Sakhon Nakon, King’s Palace,
17°9N, 104°9E, deciduous forest; on trunk, 7.i.1989 (coll. CL &
PR Deeleman) — he073/f1 (2 CD), he073/m1 ( CD). Vietnam:
Anam [Annam region], 14°15N, 107°50E — he065/f1 (MNHN).
Diagnosis
Females differ from all other Herennia species by the epigy-
nal plate with extensive sclerotisation lateral and posterior to
the chambers (Figs 2AB, 10E, 11A, C, E). Further, they
differ from H. etruscilla, H. oz, H. tone, H. deelemanae,
H. papuana, H. agnarssoni and H. milleri by the orange to
brown carapace with a central yellow V-shaped mark
(Figs 1C, 14), with sparse and weak warts, each bearing a
white long seta (Fig. 5EF), and by the epigynum with round
chambers, weak septum and antero-lateral copulatory open-
ings (Figs 2AB, 10EF, 11). They can be further distin-
guished from H. papuana, H. tone and H. deelemanae by the
flat and round abdomen with conspicuous lobes (Figs 1A,
CD, 8AC, 9AB, E, 10AD). Males differ from H. etrus-
cilla and H. papuana by the short and curved distal embolic
conductor with a broad subdistal flap (Figs 4AB, 12–13)
and from H. deelemanae by the larger size of the pedipalp
and by the relatively smaller embolic conductor flap
(Figs 4A–B, 12–13).
Description
Female (he8/f2, compare with Fig. 1 of he8/f1, both
from Kandy, Sri Lanka)
Total length 12.8.
Prosoma (Figs 1AD, 5–6, 14) 4.9 long, 4.6 wide, 2.5
high. Carapace orange and dark red-brown, yellow laterally
and posteriorly, and with a median V-shaped yellow pattern.
Chelicerae orange. Cheliceral formula 3/4/30. Sternum 1.7
long, 2.1 wide; orange in live animal (white in alcohol).
Maxillae and labium orange-brown. Anterior eye row
recurved, posterior straight. AME diameter 0.23, ALE 0.13,
PME 0.13, PLE 0.13. AME separation 0.22, PME separation
0.32, PME–PLE separation 0.41, AME–ALE separation
0.41, AME–PME separation 0.32, ALE–PLE separation
0.11. Clypeus height 0.38.
Appendages yellow, distal femora and tarsi black.
Femoral spines sparse and short. Leg I length 19.1 (Fe 5.4, Pa
1.6, Ti 4.6, Me 6.0, Ta 1.5).
Opisthosoma (Figs 1A, CD, 8–10, 14) flattened dorso-
ventrally and truncated posteriorly, with four pairs of lateral
lobes; 8.7 long, 8.2 wide (maximum width over abdominal
lobes), 4.1 high. Dorsum white with numerous red-brown
sclerotisations, mostly in depressions, these connected by
dark pigment lines. Lateral opisthosoma with longitudinal
dorso-ventral brown bands and small sclerotisations. Venter
white and yellow, with a distinct broad median band of white
pigment connecting with white pigment patches around
spinnerets and with a central grey area. Four pairs of central
sclerotisations and several rows of lateral smaller sclerotisa-
tions present on venter. Book lung covers and the sclerotisa-
tion around the spiracles red, as are the epigynum and the
sclerotised area anterior to it.
Epigynum as in Figs 2, 10EF, 11 (see also Diagnosis).
Circular epigynal chambers divided by a narrow septum.
Lateral chamber walls heavily sclerotised, black. Inner epig-
ynum as in Fig. 2CD. Copulatory ducts wide and long,
curved.
Invertebrate Systematics 399
Male (he14/m1 from Periyar, India, Figs 1E–G, 3–4,
12–13)
Total length 3.5.
Prosoma 2.0 long, 1.6 wide, 1.1 high; dark orange.
Sternum 0.8 long, 0.8 wide; orange. AME diameter 0.16,
ALE 0.09, PME 0.09, PLE 0.09. AME separation 0.13, PME
separation 0.16, PME–PLE separation 0.13, AME–ALE sep-
aration 0.06, AME–PME separation 0.16, ALE–PLE separa-
tion 0.06. Clypeus height 0.13.
Appendages orange, distal femora and patellae slightly
darker. Leg I length 8.0 (Fe 2.1, Pa 0.6, Ti 1.9, Me 2.4, Ta
1.0).
Opisthosoma 2.0 long, 1.8 wide, 1.1 high. Scutum
orange-grey, venter light grey. A paired white dot present
lateral to the spinnerets.
Pedipalp as in Figs 4, 12–13 (see also Diagnosis). EC sig-
moidal. EC proximally wide and mebraneous, distally flat,
short and broad, sclerotised, with a ridged edge.
Variation
Females: prosoma length ranges from 3.7 to 6.2; total length
from 8.1 to 19.8 (n = 30). The colouration intensity varies, but
the general pattern is as described. Yellow legs of some
females are strikingly dark brown annulated. Most females
have a flattened opisthosoma (Fig. 10AB, but compare with
Figs 1A, 9A, 10D). Very often one or both epigynal openings
are plugged with male embolic conductors (Figs 2B, 10EF,
11). Epigynal shape varies considerably, from round chambers
with a thin but conspicuous septum as described, to almost a
single chamber with weak septum (Fig. 2B, 11A, C). Males:
prosoma length ranges from 1.9 to 2.0; total length from 3.3 to
3.5 (n = 5). Male palpal morphology is as diagnosed.
Revision of Herennia
Fig. 1. Herennia multipuncta somatic morphology and sexual dimorphism. AD, Female
from Sri Lanka (he8/f1): A, lateral; B, frontal; C, dorsal; D, ventral; EG, male from India
(he14/m1); E, lateral; F, dorsal; G, ventral. Note that corresponding female and male views
are in same scale. Scale bars = 1.0 mm.
M. Kuntner400 Invertebrate Systematics
Distribution
Widespread in South and South-east Asia. Records from Sri
Lanka, India, Nepal, Myanmar, Thailand, Malaysia,
Singapore, Indonesia (Sumatra, Western Java, Borneo,
Sulawesi, Ambon), Philippines, Vietnam, China, Taiwan
(Fig. 32).
Natural history
Common on trees and walls close to humans. The species
seems to be the most synanthropic in the genus and appears
to occupy degraded habitats throughout South and South-
east Asia.
Taxonomic history
Doleschall’s (1859) ‘Second Contribution’ to Dutch Indian
(Indonesian) arachnology included descriptions of Epeira
(Argyopes) ornatissima from Amboina (Ambon, Maluku)
and Epeira (Argyopes) multipuncta from Java. Doleschall’s
descriptions, both of females, were in Dutch with Latin diag-
noses and contained female habitus illustrations of each
species. The female syntype of Epeira ornatissima from
Ambon was found in RMNH and is in a very bad condition.
As the description of E. multipuncta was based on an illus-
tration (N.K. = ‘Natuurkundige Komissie’, see Doleschall
1859: 2, 33), the type does not exist. The illustration of
E. multipuncta depicts a V-shaped mark on the carapace,
which is absent in the illustration of E. ornatissima, but the
Dutch description does mention it and it can be seen in the
holotype female. Both Doleschall’s descriptions are of the
same species (widespread in South and South-east Asia) and
thus either of the two names could be used. However, the first
reviser’s choice (Thorell 1877) was multipuncta as the valid
name for this species and ornatissima as its synonym.
Stoliczka (1869) described a new species, Epeira
(Argyopes) mammillaris, from a female collected in Assam.
Stoliczka (1869: 204) gave the type depository as ‘Indian
Museum’. I was unable to locate the type(s) and did not
receive replies to my requests from India (Zoological Survey,
Kolkatta). The description and figures provide no character
that could diagnose the species from the widespread
H. multipuncta so the species is kept a synonym of H. multi-
puncta as in Thorell (1887).
Karsch (1880: 381) described a female from Borneo
(Sampit, Kalimantan) as a new species, Herennia sampitana.
The holotype was available for examination in ZMB, and the
name is here proposed a new synonym of H. multipuncta.
Thorell (1887) synonymised H. mammilaris with
H. multipuncta, confirmed his earlier (1877) synonymy of
H. ornatissima with H. multipuncta and described a new
species based on a female from Myanmar (then Burma),
Herennia mollis Thorell, 1887. I failed to locate the type
series of Herennia mollis, but its depository may be
MCSNG. Having examined only H. multipuncta from
Myanmar, I expect the female holotype of Herennia mollis to
be conspecific. Thorell’s diagnosis mentioned the diagnostic
V-shaped mark on the carapace. Bonnet (1957) synonymised
H. mollis with H. ornatissima ( = H. multipuncta), but the
name was listed as valid by Platnick (2005).
Fig. 2. Herennia multipuncta, female epigynum. AB, Ventral: A, from
Sri Lanka (he8/f1); B, from Singapore (he6/f1), note broken embolic
conductors and a broken embolus. CD, Cleared epigynum anatomy,
from Sri Lanka (he8/f1): C, dorsal; D, ectal. Scale bars = 0.1 mm.
Fig. 3. Herennia multipuncta, male from India (he14/m1): A, lateral;
B, dorsal; C, ventral. Scale bar = 0.5 mm.
Invertebrate Systematics 401
Etymology
The name probably reflects the many dark sclerotised spots
on the female abdomen.
Herennia agnarssoni, sp. nov.
(Fig. 16AC)
Material examined
Holotype. Female (he102/f1 BMNH) from Solomon Islands: Mt.
Austen, 13.xi.1964, P. Greenslade. The locality lies at 9°28S 159°58E
on the island Guadalcanal.
Additional material examined. None.
Diagnosis
Females differ from all other Herennia species except H. tone
and H. gagamba by a group of long conspicuous spines pro-
laterally on femur I (Fig. 16AB). The known female can be
easily distinguished from H. tone and H. gagamba by the
epigynum with extensive posterior and lateral epigynal
chamber sclerotisations (Fig. 16C).
Description
Female (holotype, Fig. 16A–C)
Total length 13.3.
Prosoma (Fig. 16AB) as described in H. papuana,
but relatively longer (length to width ratio 1.4; 1.1 in
H. papuana): 5.5 long, 4.0 wide, 2.9 high. Cheliceral
formula 4/4/15. Sternum 2.0 long, 2.1 wide, bright orange.
PLE size 0.19, PME 0.16 (PLE larger than PME).
Appendages (Fig. 16AB) as described in H. papuana but
legs longer (see also Diagnosis). Leg I length 28.8 (Fe 8.4, Pa
1.7, Ti 6.8, Me 9.4, Ta 2.5). Sustentaculum conspicuous.
Revision of Herennia
Fig. 4. Herennia multipuncta, male pedipalp (AB, E, from India, he14/m1; C, from
Malaysia, he5/m1; D, from Java, he20/m1). A, Ectal; B, mesal; C, embolic division, expanded
and dissected from tegulum (embolus broken twice), note connection of embolus base and
embolic conductor via membrane, ETm (this membrane attaches the embolic division to the
tegulum as illustrated in H. papuana in Fig. 27DE), and additional membrane on EC;
D, broken distal embolus, dissected from female epigynum (note embolus hook); E, bulb
transparent, showing sperm duct, ectal (note position of embolic hook). Scale bars = 0.1 mm.
M. Kuntner402 Invertebrate Systematics
Opisthosoma (Fig. 16AB) barely flattened dorso-
ventrally with four pairs of inconspicuous lateral lobes, trun-
cated posteriorly; 7.5 long, 7.4 wide (maximum width over
abdominal lobes), 4.9 high. Dorsum white with numerous
sigillae. Lateral opisthosoma with small sclerotisations.
Venter white with a distinct broad median band of white
pigment connecting with white pigment patches around
spinnerets. Four pairs of central sclerotisations, and conspic-
uous lateral sclerotisations in many rows. Book lung covers
and the sclerotisation around the spiracles red, as are the
epigynum and the sclerotised area anterior to it. Spinnerets
of the typical form.
Epigynum (Fig. 16C) as diagnosed. Epigynal septum
barely present, weak and deep, chambers thus forming a con-
tinuous area. Lateral walls heavily sclerotised, black.
Male
Unknown.
Distribution
Known only from the type locality in Solomon Islands
(Fig. 32).
Natural history
Unknown.
Etymology
The species epithet is a patronym after my laboratory mate,
Ingi Agnarsson.
Herennia deelemanae, sp. nov.
(Fig. 17)
Material examined
Holotype. Male (he115/m1 RMNH) from [Malaysia, Sabah]
‘N. Borneo: Mt. Kinabalu N.P., Poring Hot Springs, tree 1+2 night,
Fig. 5. Herennia multipuncta, female prosoma, from Sri Lanka (he8/f5): A, frontal; B, dorsal, box delimits
area of image D; C, head region, dorsal; D, carapace edge, dorsal (note ridged edge, arrow); E, same (note
extensive hair-like setae); F, carapace detail (note warty seta base surrounded by hairs). Scale bars = 100 µm.
Invertebrate Systematics 403
6°2N 116°50E, primary forest 500–700m, canopy fogging Aglaia sp.
(Meliaceae), leg. A. Floren 27.iii.97, ex coll. C.L. Deeleman-Reinhold;
2000-704’; the real date is 1998 (C. L. Deeleman, personal
communication).
Paratype. Female (he114/f1 RMNH) from [Malaysia] ‘S Sabah,
Beaufort, 105 km S of: Long Pa Sia area: Payakalaba. 4°25N 115°44E.
12 Apr 1987. Van Tol & Huisman; At light. 18.25–21.00h. Somewhat
disturbed kerangas vegetation. 1000m’.
Additional material examined. None.
Remarks
The female paratype is hypothesised to be conspecific with
the male on the basis of the EC morphology in female copu-
latory opening (Fig. 17A, compare with 17BC) and the rel-
ative proximity of the type localities (both in Sabah).
Diagnosis
Females differ from other known species of Herennia by the
oval epigynum (Fig. 17A) with a weak septum, thin lateral
sclerotised edge and lateral copulatory openings. Males
differ from other known species of Herennia by the smaller
size (prosoma 1.3, total length 3.0) and by the embolic con-
ductor with an extensive flap (Fig. 17BC).
Description
Female (paratype, Fig. 17A)
Total length 8.2.
Prosoma 3.9 long, 3.0 wide, 2.1 high. Carapace brown-
red (no median mark) with conspicuously yellow lateral
edge. Carapace distinctly warty. Chelicerae red-brown.
Sternum 1.6 long, 1.7 wide; orange, with medially protrud-
ing white subcutaneous pigment. Maxillae and labium red-
brown. Anterior eye row recurved, posterior straight. PLE
size 0.15, PME 0.15.
Appendages. Femora and tibiae yellow, other leg joints
brown. Femoral spines fairly sparse, of medium length. Leg
Revision of Herennia
Fig. 6. Herennia multipuncta, female prosoma, from Sri Lanka (he8/f5): A, mouth parts, ventral;
B, cheliceral boss, lateral; C, ventral prosoma, frontal, box delimits area of image D; D, cheliceral detail,
frontal; E, sternum, box delimits area of image F (note paired tubercles (arrows) adjacent to each coxa);
F, sternal slit sensilla. Scale bars = 100 µm, except F = 20 µm.
M. Kuntner404 Invertebrate Systematics
I length 17.0 (Fe 4.9, Pa 1.3, Ti 4.2, Me 5.1, Ta 1.5).
Sustentaculum present, hidden among other setae.
Opisthosoma oblong, flattened dorso-ventrally with
inconspicuous lateral lobes; 4.5 long, 3.7 wide (maximum
width over abdominal lobes), 1.6 high. Dorsum white with
numerous sigillae. Lateral opisthosoma with small scleroti-
sations. Venter white with a distinct broad median band of
white pigment connecting with white pigment patches
around spinnerets. Four pairs of central sclerotisations, and
conspicuous lateral sclerotisations in many rows.
Epigynum (Fig. 17A) as diagnosed.
Male (holotype, Fig. 17B–C)
Total length 3.0.
Prosoma 1.3 long, 1.1 wide, 0.9 high; light orange.
Sternum 0.6 long, 0.6 wide. AME diameter 0.16, ALE 0.08,
PME 0.09, PLE 0.09.
Appendages orange-grey. Leg I length unknown.
Opisthosoma 1.4 long, 1.3 wide, 0.9 high. Scutum brown
(in alcohol). Venter and lateral abdomen grey. A paired white
dot present lateral to the spinnerets.
Pedipalp as diagnosed (Fig. 17BC).
Distribution
Sabah (Malaysian Borneo) (Fig. 32).
Natural history
The male was collected by canopy fogging Aglaia sp.
(Meliaceae) in primary forest at 500–700 m. The female was
collected in ‘somewhat disturbed kerangas vegetation’ at
1000 m.
Etymology
The species epithet is a patronym after Christa Deeleman-
Reinhold, who discovered the species.
Fig. 7. Herennia multipuncta, female from Sri Lanka (AC, he8/f4; DF, he8/f5): A, fourth leg distal
joints; B, fourth tarsus detail, apical (note conspicuous sustentaculum); C, same, lateral (note inconspicuous
sustentaculum); D, fourth tarsus of different female, lateral (note sustentaculum hidden among setae);
E, presumed natural pose of spinnerets, PMS hidden; F, same, detail, showing facing triads (AG and FL
spigots). Scale bars = 100 µm, except F = 10 µm.
Invertebrate Systematics 405
Herennia etruscilla, sp. nov.
(Figs 18–26)
Material examined
Holotype. Male (he58/m1 USNM) from Indonesia: Java,
Yogyakarta, Kaliurang village and adjacent forest, 900–1050 m,
16–17 Oct 1996 (coll. Kuntner & Šereg). The locality lies on the slope
of Mt. Merapi volcano at ~7°35S 110°25E.
Paratypes. Three females (he052/f1 USNM) and two males
(he052/m1 USNM) from Indonesia: W. Java, Cibodas Botanical
Gardens, near Bogor, 1300–1400 m, 24.x.1996 (coll. Kuntner & Šereg).
The locality lies in western Javan highlands at ~6°44S 107°02E. Two
paratype females and a paratype male are also photo vouchers.
Additional material examined. Indonesia: Java [no loc. data] —
he062/f1 (6 MNHN); (coll. H. Jensen) — he121/f1 ( ZMUC); Java,
Yogyakarta, Kaliurang village, 7°35S, 110°25E, 900–1050,
16–17.x.1996 (coll. Kuntner & Šereg) — he058/j1 (Imm. USNM),
Forest at night, 18.x.1996 (coll. Kuntner & Šereg) — he051/f1
( USNM); Tjibodas [Cibodas], 6°43S, 107°2E, 1907 (coll.
T Barbour) — he049/f1 (MCZ); 28.viii.1922 (coll. Th. Mortenssen)
— he122/f1 ( ZMUC), he122/j1 (Imm. ZMUC); Jawa Barat,
Buitenzorg [ = Bogor], 6°35S, 106°47E, 1909 (coll. Palmer & Bryant)
— he053/f1 ( MCZ); Jawa Timur, ‘O. Java’, Tengger Geb. [ = Bromo
Tengger Semeru NP], 8°0S, 113°0E (coll. Fruhstorfer) — he083/f1
(2 ZMB); W Java, Cibodas Botanical Gardens, 6°44S, 107°2E,
1300–1400 m, web on tree trunk, 24 Oct 1996 (coll. Kuntner & Šereg)
—he052/j1 (Imm. USNM).
Diagnosis
Females Herennia etruscilla differ from H. multipuncta,
H. sonja, H. jernej and H. gagamba by the dark red carapace
lacking yellow V-shaped mark (Figs 25–26A), with many con-
spicuous warts, each bearinga short black hook on top(Fig.19)
and by the epigynum with oval chambers, strong septum, and
lateral copulatory openings (Figs 18C,22EF). They can be
distinguished from H. tone, H. deelemanae, H. papuana,
Revision of Herennia
Fig. 8. Herennia multipuncta, female opisthosoma, from Sri Lanka (he8/f4), all ventral view: A, whole
venter, showing four paired opisthosomal lateral lobes (arrows); B, posterior venter, showing truncated
opisthosomal tip (arrow), box delimits area of image C; C, venter detail, showing median apodemes (arrows),
box delimits area of image D; D, venter detail, showing numerous lateral sclerotisations (arrows), box delimits
area of image E; E, lateral venter detail with sclerotisations, box delimits area of image F; F, lateral
sclerotisations detail. Scale bars = 100 µm, except A = 1 mm.
M. Kuntner406 Invertebrate Systematics
H. agnarssoni and H. milleri by a flat and round abdomen with
conspicuous lobes (Figs 22AB, 25–26A), and by the lateral
epigynal chamber sclerotisation, which reaches about halfway
to the anterior epigynal rim (Fig. 18C).The epigynum is larger
and the lateral sclerotisation thicker (Fig. 18C) than in females
of H. oz,and the posterior lateral eyes are the size of posterior
median eyes. Males Herennia etruscilla differ from all other
Herennia species with known males by the extremely long and
curved distal embolic conductor with a subdistal flap in the
shape of a prong (Fig. 18AB).
Description
Female (paratype he52/f2, Fig. 25, compare with
Figs 18C–D, 19–22, 26)
Total length 15.8.
Prosoma (Figs 19–20, 25–26) 5.7 long, 5.6 wide, 2.9
high. Carapace red. Chelicerae red-brown. Cheliceral
formula 3/4/23. Sternum 2.2 long, 2.6 wide; bright red in live
animal (Fig. 26B); in alcohol orange and with medially pro-
truding white subcutaneous pigment. Maxillae and labium
red-brown. Anterior eye row recurved, posterior straight.
AME diameter 0.32, ALE 0.16, PME 0.19, PLE 0.19. AME
separation 0.29, PME separation 0.44, PME–PLE separation
0.49, AME–ALE separation 0.32, AME–PME separation
0.43, ALE–PLE separation 0.18. Clypeus height 0.45.
Appendages (Figs 19AB, 21AC, 25–26). Proximal
femora and tibiae yellow, distal femora and tibiae and other
leg joints black. Femoral spines normal. Leg I length 27.0 (Fe
7.8, Pa 2.0, Ti 6.2, Me 8.7, Ta 2.3). Sustentaculum present,
but may be inconspicuous (e.g. in Fig. 21C).
Opisthosoma (Figs 22AD, 25–26) circular, flattened
dorso-ventrally, with four pairs of conspicuous lateral lobes,
truncated posteriorly; 9.6 long, 10.1 wide (maximum width
over abdominal lobes), 6.8 high. Dorsum with numerous
Fig. 9. Herennia multipuncta, female opisthosoma, from Sri Lanka (he8/f4): A, lateral, showing four paired
opisthosomal lateral lobes (arrows); B, ventral, showing truncated opisthosomal tip (arrow), box delimits area
of image C; C, venter detail, showing median apodemes (arrows), box delimits area of image D; D, venter
detail, showing (medially fused) paired apodeme; E, posterior dorsum; F, dorsum detail, with two sigillae
(arrows). Scale bars = 100 µm, except A = 1 mm.
Invertebrate Systematics 407
apodemes and sigillae (Fig. 22BD). Folium with numerous
orange-brown fields on white (Figs 25–26A). Lateral
opisthosoma with dorso-ventral longitudinal brown bands
and small sclerotisations. Venter in live animal bright orange
with a central black spot (Fig. 26B); venter in alcohol white
and grey, with a distinct broad median band of white pigment
connecting with white pigment patches around spinnerets,
and with a central grey area. Five pairs of central sclerotisa-
tions, and numerous rows of lateral sclerotisations present on
venter. Book lung covers and the sclerotisation around the
spiracles red, as are the epigynum and the sclerotised area
anterior to it.
Epigynum as diagnosed (Figs 18CD, 22EF). Oval epig-
ynal chambers divided by a strong septum. Lateral chamber
walls heavily sclerotised, thick, black. Inner epigynum as in
Fig. 18D. Copulatory ducts wide and short, curved, fertilisa-
tion ducts thin.
Male (holotype, Fig. 18A–B, compare with Figs 23–24,
25A)
Total length 4.7.
Prosoma (Fig. 23, 25A) 2.5 long, 1.9 wide, 1.4 high; light
orange (bright orange in live animal, Fig. 25A). Sternum 1.0
long, 1.0 wide. AME diameter 0.19, ALE 0.09, PME 0.09,
PLE 0.11.
Appendages (Figs 23CD, F, 25A) dark orange. Leg I
length unknown.
Opisthosoma (Figs 24, 25A) 2.6 long, 2.0 wide, 1.1 high.
Scutum whitish (but, orange in live animal, Fig. 25A). Venter
grey, lateral abdomen dark grey. Two paired white dots
present lateral to the spinnerets.
Pedipalp as diagnosed (Fig. 18AB). Conspicuous tegular
apophysis and large embolic conductor present. Embolic con-
ductor proximallywide and membranous, distally thin, sclero-
tised, with a ridged edge and a conspicuous subdistal prong.
Revision of Herennia
Fig. 10. Herennia multipuncta, female opisthosoma, from Sri Lanka (AB, he8/f5; CF, he8/f4): AB, flat
opisthosoma (he8/f5) showing extensive rows of lateral sclerotisations (arrows); A, ventral; B, lateral;
CD, round opisthosoma (he8/f4) showing less conspicuous rows of lateral sclerotisations (arrows);
C,ventral; D, lateral; EF, epigynum, ventral, with embolic conductor (arrow) stuck in copulatory opening.
Scale bars AD = 1 mm, EF = 100 µm.
M. Kuntner408 Invertebrate Systematics
Variation
Females: prosoma length ranges from 4.9 to 6.1; total length
from 12.4 to 15.8 (n = 10). Cheliceral formula 3/4/23 or
3/5/30 (n = 2). Males: prosoma length ranges from 2.0 to 2.5;
total length from 3.7 to 4.7 (n = 2). Live males are bright
orange (Fig. 25A), whereas preserved ones in alcohol are
yellow to grey.
Distribution
Java (Indonesia) (Fig 32).
Natural history
Found on trees in primary forest (Kaliurang) as well as parks
(Cibodas and Bogor). See Biology.
Etymology
Named after Herennia Etruscilla (3rd century AD), wife of
the Roman emperor Trajan Decius, and mother of Roman
emperors Herennius Etruscus and Hostilian. Old Roman
coins with the portrait of Herennia Etruscilla may have been
the source of the generic name derivation (see Etymology of
Herennia). The species epithet is a noun in apposition.
Herennia gagamba, sp. nov.
(Fig. 27CD)
Herennia sp. Hormiga et al., 1995: 334, fig. 11G–J (female geni-
talic illustrations).
Material examined
Holotype. Female (he54/f2 USNM) from Philippines: Luzon;
Nueva Vizcaya. Imugan. Sta. Fe, Smooth-barked tree, farmland.
31.v.1987 (coll. CK Starr). The approximate coordinates of the type
locality are 16°12N 120°51E.
Paratypes. 25 females (he54/f1, USNM) from the same sample as
above.
Additional material examined. Philippines: Luzon, Laguna Prov.,
Los Banos, 14°11N, 121°10E, 18.ii.1945 (coll. B Malkin) — he127/f1
Fig. 11. Herennia multipuncta, female epigyna and male EC plugs: AD, from Singapore (he6/f1);
A,ventral; B, same, right epigynal chamber detail; C, posterior; D, same, right epigynal chamber detail;
EF, from Sri Lanka (he8/f5); E, ventral; F, same, left epigynal chamber detail. Scale bars = 20 µm.
Invertebrate Systematics 409
( AMNH), he127/m1 (AMNH); Nueva Vizcaya. Imugan. Sta. Fe,
16°12N, 120°51E, smooth barked tree, farmland, 31.v.1987 (coll.
CK Starr) —he055/f1 ( USNM); [Luzon], Laguna [de Bay],
Mt. Makiling, above Los Banos, 14°89’N, 121°1124’E, 1984 (coll.
CK Starr) — he128/f1 ( AMNH); 10.iii.1985 (coll. CK Starr) —
he129/f1 ( AMNH).
Diagnosis
Females differ from all other Herennia species by the longer
than wide sternum and from all species, except H. tone and
H. agnarssoni, by a group of long conspicuous spines pro-
laterally on femur I (Fig. 27C). They can be easily distin-
guished from H. tone and H. agnarssoni by the epigynum
shape (Fig. 27D), which is round, has a strong septum and
medially positioned copulatory openings. Although the epig-
ynum shape of Herennia gagamba (Fig. 27D) resembles that
of H. deelemanae, the copulatory openings are positioned
medially and not laterally, and the septum is very pro-
nounced, in the shape of an inverted letter ‘T’.
Description
Female (paratype, he54/f1)
Total length 13.6.
Prosoma as in H. multipuncta, but relatively longer and
narrower (Fig. 27C): 5.6 long, 4.4 wide, 2.5 high. Sternum
2.1 long, 2.0 wide; orange, with medially protruding white
subcutaneous pigment. Maxillae and labium red-brown.
Cheliceral formula 3/3/18 and 3/4/15. Anterior eye row
recurved, posterior straight. PLE size 0.16, PME 0.16.
Appendages (Fig. 27C). femora and tibiae yellow, other
leg joints brown. Femoral spines short, except the diagnostic
prolateral group of long spines on first femur. Leg I length
21.3 (Fe 6.4, Pa 1.6, Ti 4.9, Me 6.5, Ta 1.9). Sustentaculum
present, hidden among other setae.
Opisthosoma (Fig. 27C) oblong, slightly flattened dorso-
ventrally with inconspicuous lateral lobes; 9.3 long, 6.8 wide
(maximum width over abdominal lobes), 5.6 high. Dorsum
white with numerous sigillae, covered with sparse but strong
spines, especially posteriorly. Lateral opisthosoma with
Revision of Herennia
Fig. 12. Herennia multipuncta, male right pedipalp, from India (he14/m1): A, ectal; B, ecto-apical; C, mesal;
D, meso-apical; E, dorso-apical; F, bulb detail, dorso-ectal. Scale bars = 100 µm.
M. Kuntner410 Invertebrate Systematics
small sclerotisations. Venter white with a distinct broad
median band of white pigment connecting with white
pigment patches around spinnerets. Four pairs of central
sclerotisations and conspicuous lateral sclerotisations in
many rows. Spinnerets of the typical nephiline condition.
Epigynum (Fig. 27D) as diagnosed.
Male
The only known male (he127/m1) lacks both pedipalps.
Male pedipalp known only by epigynal plugs consisting of
EC and embolus, found stuck in female copulatory openings
(Fig. 27D); this also illustrated in Hormiga et al. (1995:
fig. 11G–J). EC is distally massive with a subdistal flap
resembling that of H. deelemanae.
Variation
Females: prosoma length ranges from 5.1 to 5.8; total length
from 10.4 to 15.5 (n = 26). Cheliceral formula varies within
the same individual (see above). Opisthosoma almost
unlobed (Fig. 27C) to lobed. Spines on dorsum are very pro-
nounced in some females, and less in others, but are consis-
tently present.
Distribution
Luzon (Philippines) (Fig. 32).
Natural history
Unknown. The collection label (see above) implies the
spiders were collected from tree bark.
Etymology
The species epithet, a noun in apposition, is a Tagalog word
for a spider.
Fig. 13. Herennia multipuncta, male right pedipalp distal sclerites, from India (he14/m1): AB, apical;
C, ectal; DE, dorso-apical; F, meso-apical. Membranous part of embolic conductor (EC-m) is continuous
with the sclerotised part (EC), both attached to embolus base (EB) and tegulum (T) via membrane (compare
with Fig. 4C). Scale bars = 100 µm.
Invertebrate Systematics 411
Herennia jernej, sp. nov.
(Fig. 27E)
Material examined
Holotype. Female (he19/f1 CAS) from Indonesia: ‘Sumatera
Barat; Mangani, mine near Kota Tinggi. 700m. 20 July 1983 Edward S.
Ross‘. The approximate coordinates of the type locality on Sumatra are
00°02S 100°20E.
Paratypes. Tw o immature specimens (he20–21 CAS) from the
same sample as above.
Additional material examined. None.
Diagnosis
Holotype differs from all Herennia species but H. multi-
puncta, H. gagamba and H. sonja by the carapace with a
central yellow V-shaped mark. Epigynum (Fig. 27E) differs
from H. multipuncta, H. gagamba and H. sonja by the oval
chambers, strong septum, and a fairly thin posterior sclero-
tised edge. Leg formula (1, 2 = 4, 3) appears to be unique in
Herennia. Male unknown.
Description
Female (holotype, Fig. 27E)
Total length 10.3.
Prosoma 5.2 long, 4.8 wide, 2.8 high. Sternum 2.0 long,
2.4 wide. Prosoma as described in H. multipuncta except
pale colouration (almost white). Cheliceral formula 3/4/30.
AME diameter 0.23, ALE 0.16, PME 0.14, PLE 0.13. AME
separation 0.32, PME separation 0.38, PME–PLE separation
0.45, AME–ALE separation 0.28, AME–PME separation
0.38, ALE–PLE separation 0.19. Clypeus height 0.40.
Appendages.Leg I length 21.3 (Fe 6.2, Pa 1.4, Ti 4.9,
Me 6.9, Ta 1.9). Femora with long spines.
Opisthosoma as described in H. multipuncta, but longer
than wide and white, 6.2 long, 5.1 wide (maximum width
over abdominal lobes), 3.5 high.
Epigynum as diagnosed (Fig. 27E).
Male
Unknown.
Distribution
Only known from the type locality on the Indonesian island
of Sumatra (Fig. 32).
Revision of Herennia
Fig. 14. Herennia multipuncta, photographs of live spiders from Sri
Lanka: AB, female in her web built tightly against tree trunk, resting at
hub in the shape of a depression (‘hub-cup’, arrow); C, female on a mud
wall. Note diagnostic yellow V-shaped mark on carapace and white and
spotted dorsum.
Fig. 15. Herennia multipuncta, photograph of a female in her web,
from Thailand (by G. Hormiga). Note ‘pseudoradii’ (white arrows),
which do not pass the hub (black arrow).
M. Kuntner412 Invertebrate Systematics
Natural history
Unknown.
Etymology
The species epithet (pronounced ‘yer-ney’), a noun in appo-
sition, is a patronym after my brother, Jernej Kuntner.
Herennia milleri, sp. nov.
(Fig. 16DG)
Material examined
Holotype. Female (he113/f1 ZMUC) from Papua New Guinea:
Bismarck Isl., New Britain, Yalom 1000 m, 10 May 1962, Noona Dan
Exp. 61–62. The locality is approximately at 4°25S 151°44E.
Paratype. Female (he126/f1 AMNH) from Papua New Guinea:
Louisiade Arch., Rossel Island, Abaleti, SSW of Mt. Rossel, 11°21S,
154°10E, 0–50, Camp #12 (Fifth Archbold Exped. to New Guinea:
Papau [sic], 1956 (coll. LJ Brass).
Additional material examined. None.
Diagnosis
Females diagnosed as H. papuana except in having poste-
rior lateral eyes larger than posterior median eyes. Male
embolic conductor (known only from the broken organ in
holotype epigynum, Fig. 16G) has a subdistal prong (unlike
H. papuana), which is longer than in H. etruscilla and
H. oz.
Description
Female (holotype, Fig. 16D–G)
Total length 12.4. Somatic and external genital morphol-
ogy as described in H. papuana but opisthosoma flatter.
Prosoma (Fig. 16DE) 5.6 long, 4.7 wide, 3.1 high.
Cheliceral formula 3/4/30. Sternum 2.1 long, 2.3 wide, bright
orange. PLE size 0.16, PME 0.13 (PLE larger than PME).
Appendages (Fig. 16F) as in H. papuana. Leg I length
23.9 (Fe 6.9, Pa 1.7, Ti 5.6, Me 7.8, Ta 1.9). Sustentaculum
apparently absent.
Opisthosoma (Fig. 16DE) flattened dorso-ventrally with
four pairs of inconspicuous lateral lobes, truncated posteri-
orly; 7.4 long, 5.5 wide (maximum width over abdominal
lobes), 3.7 high.
Epigynum (Fig. 16G) as in H. papuana.
Male
Unknown, except for EC (see Diagnosis).
Fig. 16. Herennia agnarssoni, sp. nov., female holotype from Solomon Islands (AC) and Herennia milleri,
sp. nov., female holotype from New Britain (DG): A, dorsal, note diagnostic long spines on first femur;
B,ventral; C, epigynum, ventral; D, dorsal; E, ventral; F, first femur prolateral (note absence of long spines);
G, epigynum, ventral. Scale bars AB, DF = 1.0 mm, C, G = 0.1 mm.
Invertebrate Systematics 413
Distribution
New Britain and Rossel Island (Papua New Guinea) (Fig. 32).
Natural history
Unknown.
Etymology
The species epithet is a patronym after my laboratory mate,
Jeremy Miller.
Herennia oz, sp. nov.
(Fig. 28)
H. sp. Davies, 1988: fig. 18 (in part), illustration of female habitus
and epigynum.
Material examined
Holotype. Female (he22/f1 QM) from Australia: Northern
Territory: East Alligator River, 12°25S, 132°58E, 16–18 July 1979
‘(RF), G. Monteith & D. Cook’ with additional label ‘Drawn 1985
SR. Monteith’.
Paratype. Female (he23/f1 QM) from Australia: Northern
Territory: West Alligator River, mouth, 12°12S, 132°13E, ‘WA1,
22–24 July 1979 (OF), G. Monteith & D. Cook’.
Additional material examined. Australia: Northern Territory:
Edge Arhemland Res., Ben Hole Billabong, 120 mi SE Darwin,
13°57S, 131°57E, ii.1972 (coll. J Anderson) — ne123/f1 ( AMNH),
ne124/f1 ( AMNH), ne125/f1 ( AMNH).
Diagnosis
Females diagnosed as H. etruscilla, but differ from it by the
smaller epigynum with narrower lateral sclerotisation
(Fig. 28BC) and by the more oblong opisthosoma. Males
unknown, except an embolic conductor in epigynum
(Fig. 28B), which has a flap in the shape of a prong (compare
H. etruscilla).
Revision of Herennia
Fig. 17. Herennia deelemanae, sp. nov., female paratype and male
holotype, from Sabah, Malaysian Borneo: A, epigynum, ventral (note
EC with extensive flap in one copulatory opening and only embolus in
the other); B, male left pedipalp, ectal; C, same, mesal (note extensive
EC flap used as evidence for conspecificity of sexes, each known from
one individual). Scale bar = 0.5 mm.
Fig. 18. Herennia etruscilla, sp. nov. A, B, Male pedipalp, from Java (holotype): A, ectal;
B, mesal. C, D, Female epigynum, from Java (he53): C, ventral; D, dorsal, cleared. Scale bars
= 0.5 mm.
M. Kuntner414 Invertebrate Systematics
Description
Female (holotype, Fig. 28A–B, compare with 28C)
Total length 11.6.
Prosoma (Fig. 28A) as described in H. etruscilla, but with
less warty carapace and with less rounded edges, 5.6 long,
4.6 wide, 2.1 high. Cheliceral formula 3/4/13. Sternum 2.0
long, 2.3 wide, orange with protruding white pigment. PLE
size 0.18, PME 0.16 (PLE larger than PME).
Appendages yellow, tarsi brown. Femora with numerous
short spines. Leg I length 22.0 (Fe 6.2, Pa 1.6, Ti 5.6, Me 6.6,
Ta 2.0). Sustentaculum conspicuous.
Opisthosoma (Fig. 28A) flattened dorso-ventrally with four
pairs of conspicuous lateral lobes, truncated posteriorly; 7.1
long, 6.5 wide (maximum width over abdominal lobes), 2.6
high. Dorsum white with numerous sigillae. Venter white with
a distinct broad median band of white pigment connecting with
white pigment patches around spinnerets. Four pairs of central
sclerotisations and inconspicuous lateral sclerotisations.
Epigynum (Fig. 28B, compare with 28C) as diagnosed.
Epigynal septum strong, chambers oval, copulatory open-
ings lateral. Inner epigynal anatomy as in H. etruscilla.
Male
Unknown, except the EC (see Diagnosis).
Variation
Females: prosoma length ranges from 5.6 to 5.8; total length
from 11.6 to 13.6 (n = 2). Cheliceral formula 3/3/12
(paratype) or 3/4/13 (holotype).
Distribution
Australia’s Northern Territory (Fig. 32).
Fig. 19. Herennia etruscilla, sp. nov., female prosoma, from Java (paratype he52/f1): A, dorsal, boxes
delimiting areas of images B and C; B, lateral carapace detail, note warty cuticle, dense and thin setae
prolaterally on first femur (arrow); C, head region detail with warts and hair-like setae; D, head region;
E,eye region; F, thoracic region detail (note seta (arrow) on top of wart surrounded by hair-like setae). Scale
bars = 100 µm.
Invertebrate Systematics 415
Natural history
Unknown.
Etymology
Named after a nickname for Australia, Oz; the species
epithet is a noun in apposition.
Herennia papuana Thorell
(Figs 29–30)
Herennia papuana Thorell, 1881: 77, description of female (from
New Guinea).
H. ornatissima Chrysanthus, 1971: 41, figs 85–86, description of
female, misidentification.
H. sp. Davies, 1988: fig. 18 (in part), illustration of damaged male
pedipalp, also in Hormiga et al., 1995: fig. 5J.
Material examined
Type material. Not found.
Additional material examined. Papua New Guinea: ‘Pindiu,
Territory New Guinea’, 6°26S, 147°30E, 18.vi.1964 (coll. R Zweifel)
he135/f1 (2AMNH); New Guinea, Regensberg, 12.v.1919 (coll.
Buergers) — he082/f1 (ZMB); Morobe Prov, Wau Ecology Institute
grounds, 7°20S, 146°43E, tree trunks, shrubs, litter, 2.xi.1980 (coll. WA
Shear) he133/f1 (AMNH);11.xi.1980 (coll.WA Shear) he134/f1
( AMNH), he134/m1 ( AMNH); Morobe Province, Wau, 7°20S,
146°43E, 6.iii.1979 (coll. Levi, Lubin, Robinson) he025/f1 (2
MCZ); Jul 1979 (coll. M Robinson) — he024/m1 (5 MCZ); roadside,
19.iii.1979 (coll. H. Levi) he050/f1 ( MCZ), he050/m1 ( MCZ).
Diagnosis
Females differ from H. multipuncta, H. sonja, H. jernej and
H. gagamba by the dark red carapace lacking yellow V-
shaped mark (Fig. 29A), with many conspicuous warts, each
bearing a short black hook on top and by the epigynum
(Fig. 29C) with oval chambers, strong septum, and lateral
Revision of Herennia
Fig. 20. Herennia etruscilla, sp. nov., female prosoma, from Java (paratype he52/f1): A, lateral, box
delimits area of image B; B, cheliceral boss; C, cheliceral boss detail; D, ventral, boxes delimiting areas of
images E and F; E, chelicerae, ventral; F, detail of sternum with slit sensilla (arrows). Scale bars = 100 µm,
except C = 10 µm.
M. Kuntner416 Invertebrate Systematics
copulatory openings. They can be distinguished from H. etr-
uscilla, H. oz, H. tone and H. deelemanae by an oblong
abdomen with inconspicuous lobes (Fig. 29AB), and by the
lateral epigynal chamber sclerotisation, which reaches high
to the anterior epigynal rim (Fig. 29C). They differ from
H. agnarssoni and H. milleri by the stronger epigynal septum
(Fig. 29C). Males differ from all other Herennia species with
known males by the long and thin distal EC, which lacks a
flap or prong (Fig. 30AB).
Description
Female (he25/f1 from Papua New Guinea, Fig. 29)
Total length 13.6.
Prosoma (Fig. 29A) 4.9 long, 4.4 wide, 2.5 high.
Carapace red medially and orange laterally. Chelicerae red-
brown. Cheliceral formula 3/3/25. Sternum 2.0 long, 2.2
wide; orange, with medially protruding white subcutaneous
pigment. Maxillae and labium red-brown. Anterior eye row
recurved, posterior straight. AME diameter 0.28, ALE 0.17,
PME 0.14, PLE 0.14. AME separation 0.22, PME separation
0.35, PME–PLE separation 0.47, AME–ALE separation
0.25, AME–PME separation 0.25, ALE–PLE separation
0.13. Clypeus height 0.28.
Appendages.Femora and tibiae yellow, other leg joints
brown. Femoral spines fairly sparse and short. Leg I length
20.4 (Fe 5.7, Pa 1.5, Ti 4.9, Me 6.4, Ta 1.9). Sustentaculum
apparently absent.
Opisthosoma (Fig. 29AB) barely flattened dorso-
ventrally with three pairs of inconspicuous lateral lobes,
truncated posteriorly; 8.1 long, 5.7 wide (maximum width
over abdominal lobes), 4.8 high. Dorsum white with numer-
ous little sclerotised brown dots. Lateral opisthosoma with
longitudinal dorso-ventral brown bands and small sclerotisa-
tions. Venter white and grey, with a distinct broad median
band of white pigment connecting with white pigment
Fig. 21. Herennia etruscilla, sp. nov., female, from Java (paratype he52/f1): A, fourth tibia, prolateral, box
delimits area of image B; B, same, detail, arrows pointing to paired lyriform organ; C, fourth tarsus, with
apparent absence of sustentaculum; D, spinnerets, apical; E, same, lateral, box delimits area of image F;
F, detail of spinnerets. Scale bars = 100 µm.
Invertebrate Systematics 417
patches around spinnerets and with a central grey area. Four
pairs of central sclerotisations, and a single row of incon-
spicuous lateral sclerotisations present on venter. Book lung
covers and the sclerotisation around the spiracles red, as are
the epigynum and the sclerotised area anterior to it.
Epigynum as diagnosed (Fig. 29C). Oval epigynal cham-
bers divided by a strong septum. Lateral chamber walls
heavily sclerotised, black.
Male (he24/m1 from Papua New Guinea, Fig. 30)
Total length 3.5.
Prosoma 2.0 long, 1.6 wide, 1.1 high; dark orange.
Sternum 0.9 long, 0.8 wide; orange, with wide dark lateral
edges. AME diameter 0.15, ALE 0.09, PME 0.08, PLE 0.09.
AME separation 0.11, PME separation 0.16, PME–PLE sep-
aration 0.13, AME–ALE separation 0.07, AME–PME sepa-
ration 0.16, ALE–PLE separation 0.03. Clypeus height 0.13.
Appendages dark orange. Leg I length 8.0 (Fe 2.2, Pa 0.6,
Ti 1.9, Me 2.3, Ta 1.0).
Opisthosoma 2.1 long, 1.6 wide, 1.0 high. Scutum light
orange, caudally black. Venter grey, lateral abdomen yellow
and dark grey. Two paired white dots present lateral to the
spinnerets.
Pedipalp as in Fig. 30 (see also Diagnosis). EC proxi-
mally wide and membranous, distally thin, sclerotised, and
with a ridged edge, lacking a flap.
Variation
Females: prosoma length ranges from 4.9 to 5.6; total length
from 11.5 to 13.6 (n = 3). Males: prosoma length ranges from
1.6 to 2.0; total length from 3.1 to 3.5 (n = 5). The intensity
of colouration varies. Abdomen colour (in alcohol) ranges
from whitish through orange to dark grey. Distal leg joints of
some males are dark grey.
Revision of Herennia
Fig. 22. Herennia etruscilla, sp. nov., female opisthosoma, from Java (paratype he52/f1): A, lateral;
B, dorsal, box delimits area of image C; C, dorsum detail, box delimits area of image D (note large paired
apodemes); D, detail of dorsal apodeme and another sclerotisation posterio-lateral to it (arrow); E, epigynum,
ventral, box delimits area of image F; F, epigynum detail, note embolic conductors plugging copulatory
openings. Scale bars = AB = 1 mm, CF = 100 µm.
M. Kuntner418 Invertebrate Systematics
Distribution
Papua New Guinea (Fig. 32).
Natural history
Summarised in Biology.
Taxonomic history
Based on a female from New Guinea, Thorell (1881: 77)
described a new species, Herennia papuana. Thorell’s three-
page Latin description treats only two diagnostic characters;
the warty prosoma and the epigynum, which is of a different
form to H. multipuncta. I believe the type depository to be
MCSNG, but the type has remained unavailable for my
examination. As I only know of one species occurring in
New Guinea, I fix the name papuana to that species.
Herennia sonja, sp. nov.
(Fig. 27AB)
Material examined
Holotype. Female (he47/f1 USNM), from Indonesia: South
Kalimantan, Loksado-Malaris villages, E of Kandagan. 250m Elev. 18
Nov 1996 (coll. Kuntner & Šereg). The approximate coordinates are
02°05S 115°25E.
Paratype. Female (he57/f1 USNM), from Indonesia: Propinsi
Kalimantan Selatan, Aranio Dist., Belangian village and surroundings.
100m elev. 10–15 Nov 1996 (coll. Kuntner & Šereg). The approximate
coordinates are 3°36S, 115°4E.
Additional material examined. Indonesia: Celebes [Sulawesi],
Minahassa [peninsular], 1°0N, 124°4E (coll. Kunenethae) —
he104/f1 ( BMNH).
Diagnosis
Females differ from all Herennia species but H. multipuncta,
H. gagamba and H. jernej by the carapace with a central yellow
V-shaped mark (Fig. 27A). Epigynum (Fig. 27B) differs from
Fig. 23. Herennia etruscilla, sp. nov., male ‘eunuch’ prosoma (note both palpal bulbs lacking), from Java
(he52/m1): A, prolateral; B, ventroapical; C, lateral; D, dorsolateral; E, head region, dorsal; F, second tibia,
detail, ectal. Scale bars = 100 µm.
Invertebrate Systematics 419
H. multipuncta, H. gagamba and H. jernej by the elongate
chambers, thin septum and a thin posterior sclerotised edge.
Description
Female (holotype, Fig. 27A–B)
Total length 10.2. Prosoma, opisthosoma, and legs as
described in H. multipuncta.
Prosoma (Fig. 27A) 4.4 long, 3.5 wide, 2.3 high. Sternum
1.4 long, 1.9 wide. AME diameter 0.25, ALE 0.16, PME
0.14, PLE 0.13. AME separation 0.22, PME separation 0.30,
PME–PLE separation 0.38, AME–ALE separation 0.26,
AME–PME separation 0.28, ALE–PLE separation 0.10.
Clypeus height 0.32.
Appendages.Leg I length 19.5 (Fe 5.3, Pa 1.4, Ti 4.4,
Me 6.5, Ta 1.9).
Opisthosoma (Fig. 27A) 5.8 long, 5.8 wide (maximum
width over abdominal lobes), 2.9 high.
Epigynum as diagnosed (Fig. 27B).
Male
Unknown.
Distribution
Indonesia: Southern Kalimantan, Sulawesi (Fig. 32).
Natural history
Unknown.
Etymology
The species epithet (pronounced ‘son-ya’), a noun in apposi-
tion, is a patronym after my mother, Sonja Kuntner.
Revision of Herennia
Fig. 24. Herennia etruscilla, sp. nov., male opisthosoma, from Java (he52/m1): A, ventrolateral; B, book
lung, detail (note ridged surface); C, venter, box delimits area of image D; D, epigastral area with epiandrous
gland spigots; E, anterior venter; F, dorsum and lateral opisthosoma (note numerous scleriotisations on
scutum). Scale bars = 100 µm, except D = 10 µm.
M. Kuntner420 Invertebrate Systematics
Herennia tone, sp. nov.
(Fig. 31)
Material examined
Holotype. Female (he26/f1 MCZ) from Philippines: Dumaquete
Negros 1500’ 6–3-1949 (coll. BB Brues). The approximate coordinates
of the type locality are 09°18N 123°18E. In the same vial a male of
Nephilengys malabarensis was stored, which is now labelled as such.
Paratype. Female (he32/f1 MCZ) from Philippines: Dumaguete,
Negros Or. 3600Ft. x.1942 (coll. JW Chapman). The approximate coor-
dinates are 09°19N 123°08E.
Additional material examined. Philippines: Luzon, Beguios [ =
Baguio], 16°24N, 120°35E, 1933 (coll. JW Chapman) — he031/f1
( MCZ); Negros, Dumaguete, Cuernos Mts. 9°19N, 123°8E, 1097 m,
18.x.1942 (coll. JW Chapman) — he029/f1 ( MCZ), he030/f1
(2 MCZ); Samar Island, [no exact loc. data], 12°0N, 125°0E (coll.
Whitehead) — he098/f1 ( BMNH).
Diagnosis
Females differ from all other Herennia species except
H. gagamba and H. agnarssoni by a group of long conspicu-
ous spines prolaterally on femur I (Fig. 31B). They can be
easily distinguished from H. agnarssoni and H. gagamba by
the shape of epigynum (Fig. 31CD), which is wide with
lateral copulatory openings, a weak septum and thin sclero-
tised edge.
Description
Female (holotype, Fig. 31)
Total length 9.5.
Prosoma (Fig. 31AB) 3.9 long, 3.2 wide, 2.0 high.
Carapace orange-red medially, yellow laterally and frontally,
with few weak warts. Chelicerae orange. Cheliceral formula
3/3/10. Sternum 1.6 long, 1.6 wide; transparent (in alcohol),
with medially protruding white subcutaneous pigment.
Maxillae and labium orange. Anterior eye row recurved, pos-
terior straight. AME diameter 0.22, ALE 0.18, PME 0.13,
PLE 0.16. [PLE larger than PME] AME separation 0.22,
PME separation 0.30, PME–PLE separation 0.35,
AME–ALE separation 0.19, AME–PME separation 0.25,
ALE–PLE separation 0.13. Clypeus height 0.25.
Appendages (Fig. 31AB). Femora and tibiae yellow,
other leg joints brown. Femoral spines fairly sparse and
short, except for the prolateral group of long spines on femur
I. Leg I length 18.6 (Fe 5.2, Pa 1.3, Ti 4.4, Me 5.8, Ta 1.9).
Opisthosoma (Fig. 31AB) slightly flattened dorso-
ventrally with four pairs of inconspicuous lateral lobes,
barely truncated posteriorly; 6.0 long, 5.4 wide (maximum
width over abdominal lobes), 2.8 high. Dorsum white with
Fig. 25. Herennia etruscilla, sp. nov., photographs of live spiders
from Java (paratype female he52/f2 and male he52/m1): A, female at
hub of orb web built tightly against tree trunk and tiny orange ‘eunuch’
male (above, right, see Figs 21–22) cohabiting; note ‘pseudoradii’
(white arrows), which do not pass the hub unlike true radii (black
arrows); B, close up of female resting at hub in the shape of a depression
(‘hub-cup’, arrow); C, same, lateral view (note two-dimensionality of
spider and its web, tightly following tree shape). Note female cryptic
colouration matching tree bark and absence of V-shaped mark on
carapace.
Fig. 26. Herennia etruscilla, sp. nov., photographs of live female from
Java (paratype he52/f3): A, female at hub of her web; note spider cryptic
dorsal colouration, ‘pseudoradii’ in web (white arrows), and egg sac in
web (black arrow); B, close up of female ventral side after removing her
from the web; note bright ventral colouration (blood-red sternum and
orange venter) in striking contrast with cryptic dorsal side (A).
Invertebrate Systematics 421
numerous sclerotised brown dots. Lateral opisthosoma with
longitudinal dorso-ventral brown bands and small sclerotisa-
tions. Venter white, with a central grey area, and with white
pigment patches around spinnerets. Four pairs of central
sclerotisations, and a single row of inconspicuous lateral
sclerotisations present on venter. Book lung covers and the
sclerotisation around the spiracles red, as are the epigynum
and the sclerotised area anterior to it.
Epigynum (Fig. 31CE) wide, with narrow lateral sclero-
tisation and thin black sclerotised edge. Oval epigynal cham-
bers almost connected medially, barely separated with weak
septum. Lateral sclerotised chamber walls thin, posterior
sclerotised chamber wall conspicuously ‘tongued’. Inner
epigynum as in Fig. 31E. Oval spermathecae juxtaposed
medially, copulatory ducts long and massive, connected to
the integument by enveloping the lateral sclerotisation.
Male
Unknown, except for epigynal plugs consisting of
embolic conductors and emboli, found stuck in copulatory
openings of four females (Fig. 31D). The distal EC is
massive and different to the ones described, but seems to
have the subdistal flap.
Variation
Females: prosoma length ranges from 3.9 to 4.6; total length
from 9.3 to 11.5 (n = 6). Carapace colour ranges from orange
to dark red-brown, almost black. Carapace lacks a yellow
Revision of Herennia
Fig. 27. AB, Herennia sonja, sp. nov., female holotype from Kalimantan: A, habitus, dorsal;
B,epigynum, ventral. CD, Herennia gagamba, sp. nov., female paratype (he54/f1) from Luzon:
C, habitus, dorsal (note prolateral long spines on first femur); D, epigynum, ventral. E, Herennia
jernej, sp. nov., epigynum of female holotype from Sumatra, ventral.
M. Kuntner422 Invertebrate Systematics
V- shaped mark entirely (Fig. 31A) or the mark inconspoicu-
ously present (Fig. 31B). Warts on carapace can carry dark
macrosetae. Sternum can be wider than long (he31) or as
wide as long (holotype). Opisthosoma almost unlobed
(Fig. 31A) to lobed (Fig. 31B).
Distribution
Philippine Island Negros (Fig. 32). An additional, question-
able locality is that of the female sample he31, the MCZ
locality label reading: ‘Philippines: Beguio_s 1933;
J.W. Chapman.The closest name to Beguios, which does not
seem to exist, is Baguio on the island of Luzon (see above for
coordinates of Baguio).
Natural history
Unknown.
Etymology
The species epithet (pronounced ‘to-ne’), a noun in apposi-
tion, is a patronym after my father, Tone Kuntner.
Phylogeny
Equally weighted analyses resulted in ten most parsimonious
trees (tree length = 490 steps, CI = 40, RI = 70). In the strict
consensus, four nodes collapse and the length increases to
506 steps (Fig. 33A). Nephilid monophyly (Clitaetra,
(Herennia, (Nephilengys, Nephila))) and the monophyly of
all nephilid genera, Herennia included, is confirmed. As in
previous analysis (Kuntner, in press) the sister-group to
nephilids remains ambiguous. Equally parsimonious trees
group nephilids with: (1) the clade containing all other ara-
neoids; (2) araneids; (3) the clade containing tetragnathids +
reduced piriform clade (here represented by
Theridiosomatidae, Linyphiidae, Pimoidae, Theridiidae and
Nesticidae); (4) Phonognatha, nephilids + Phonognatha, in
turn, are sister to Deliochus, and this clade groups with the
clade containing tetragnathids + reduced piriform clade; and
(5) the clade ((Deliochus + Phonognatha) + (tetragnathids +
reduced piriform clade)).
Successive weighting analysis stabilised after the second
iteration and resulted in a single tree (Fig. 33B) identical to
one of the fundamental cladograms described above as ‘4’.
This tree differs from the successively weighted trees derived
from datasets focusing on Clitaetrinae (Kuntner, in press)
and Nephilengys (M. Kuntner, unpublished data) in the
sister-group relationship of nephilids with Phonognatha.
Such grouping, though very poorly supported (Fig. 33B),
corroborates Hormiga et al.’s (1995) nephiline placement of
Phonognatha, though not within tetragnathids. The unam-
biguous synapomorphies of Phonognatha + Nephilidae are
sternum colouration (character 24/state 3), absence of con-
ductor (133/1), absence of hub bite-out (166/1), presence of
tertiary radii (173/1) and persistence of a non-sticky spiral in
the finished web (176/1). Hormiga et al. (1995) did not
include Deliochus; its grouping with the Phonognatha +
nephilid clade also receives meagre support in the present
study, it is supported by two unambiguous synapomorphies
(chambered epigynum 76/1, presence of the embolic con-
Fig. 28. Herennia oz, sp. nov., female from Australia. AB, Holotype (he22); A, habitus,
dorsal; B, epigynum, ventral (note EC in each copulatory opening). C, Paratype (he23),
epigynum, ventral. Scale bar (BC) = 0.5 mm.
Invertebrate Systematics 423
ductor 136/1), but like the clade Phonognatha + nephilids,
collapses in the strict consensus (Fig. 33A).
Nephilid monophyly sensu Kuntner (in press) is well sup-
ported (23 unambiguous synapomorphies, bootstrap
97/Bremer > 10), and so is the monophyly of Clitaetrinae
(represented by Clitaetra, 99/6), Nephilinae (with Herennia,
Nephila and Nephilengys, 73/4), Nephilengys (90/5), Nephila
(87/4) and Herennia (100/7). The support for Nephila +
Nephilengys is relatively low (50/3). Within Herennia,
H. papuana is sister to all other species (47/1) and within the
latter clade, the group H. multipuncta + H. etruscilla (56/1) is
sister to the clade with the newly described species. Among
these, all but H. deelemanae are known from a single sex,
which may be the reason for shaky branches (Fig. 33B).
Discussion
Classification
The successively weighted tree (Fig. 33B) is the basis for the
proposed taxonomy of Herennia, but it cannot offer a strong
hypothesis for the position of the enigmatic Australian
Phonognatha and Deliochus. Along with the exact position
of Nephilidae within Araneoidea, the relationships of these
two genera will be tested with a broader taxon sample else-
where. However, the results imply that nephilines are not
tetragnathids (contra Hormiga et al. 1995), and their araneid
placement (Wunderlich 1986, 2004; Kuntner 2003; Pan et al.
2004) is supported only by a subset of trees and is thus ques-
tionable. Thus, treating the clade Nephilidae at family level
(as in Kuntner, in press) seems appropriate and such classi-
fication will be retested in future analyses.
Biogeography
Biogeographic lines delimiting regional biotas are usually
derived for one taxon at a time but some well known bound-
aries, like Wallace’s line marking the boundary between the
Oriental andAustralian regions, often coincide with geological
or climatic barriers of the past, which have prevented organism
dispersal (Brown and Lomolino 1998). Wallace’s line corre-
sponds well to the outer limit of the Sunda Shelf, which is
believed to have been part of continental South-east Asia
during the Pleistocene (Brown and Lomolino 1998). Wallace’s
and Huxley’s lines (Fig. 34) run between Bali and Lombok and
between Borneo and Sulawesi, but differ in the inclusion of the
Philippine islands; Wallace’s line marks them as Oriental and
Huxley’s as Australasian (from Brown and Lomolino 1998).
Other lines marking biotic boundaries have been proposed (see
Brown and Lomolino 1998: fig. 10.10) to accommodate spe-
cific taxa, but they mostly run between the islands of eastern
Revision of Herennia
Fig. 29. Herennia papuana, female, from Papua New Guinea (he25): A, habitus, dorsal;
B, opisthosoma, ventral; C, epigynum, ventral (note embolus in copulatory opening). Scale
bars AB = 1 mm, C = 0.1 mm.
M. Kuntner424 Invertebrate Systematics
Indonesia, in the so called Wallacea region.Those eastern lines
may not be relevant for Herennia. The currently understood
Herennia phylogeny is plotted on the geographic distribution
of the species included in the analysis (Fig. 34). The
‘basalmost’ species is H. papuana from New Guinea, and the
most ‘derived’ species are H. agnarssoni from Solomon
Islands and H. tone from the Philippines. Kuntner (in press)
roughly estimates the nephiline clade (with Herennia, Nephila
and Nephilengys)tobeatleast 160 million years old and of
Gondwanan origin.The currently understood Herennia phylo-
geography does not dispute the above, but calls for an expla-
nation of speciation/colonisation of the Laurasian landmasses:
mainland Asia, islands of Indonesia and the Philippines.
Assuming an Australasian origin of Herennia, four dis-
persal events could be postulated over Wallace’s line
(Fig. 34): the clade (H. etruscilla + H. multipuncta),
H. deelemanae, and two dispersals to the Philippines
(H. gagamba, H. tone). The alternative, and more parsi-
monious, explanation may be one dispersal event in the
common ancestor to all species excluding H. papuana and
one to Solomon Islands (H. agnarssoni). Over Huxley’s line
(Fig. 34), at least two dispersal events could be postulated,
the clade (H. etruscilla + H. multipuncta) and H. deelemanae
or, alternatively, one dispersal in the common ancestor to all
species excluding H. papuana and a reversal in the common
ancestor to H. gagamba, H. tone and H. agnarssoni.
However, the newly drawn ‘Herennia line’ (Fig. 34), joining
only Sumatra and Taiwan with SE Asia, is crossed by a single
species, H. multipuncta, suggesting all Australasian specia-
tion events in Herennia.
Fig. 30. Herennia papuana, male left pedipalp, from Papua New Guinea (AB, he24/m1;
CE, he24/m2): A, ectal; B, mesal; C, expanded, embolus pulled out of EC, dorsal; D, distal
sclerites dissected from remaining sclerites; E, further dissection of embolic division. Scale
bar = 0.1 mm. Note that embolus base (EB) and embolic conductor (EC) are attached to
tegulum (T) via joint membrane (ETm) and together form the nephilid embolic division.
Embolic conductor has a sclerotised (EC) and a membranous part (EC-m).
Invertebrate Systematics 425
Such hypothesis of Australasian speciation in Herennia is
consistent with the currently understood ecology of
Herennia, where most species appear to be narrow island
endemics, likely confined to pristine forests, whereas the
widespread H. multipuncta, ranging from India and Sri
Lanka to eastern Indonesia, appears to expand its range
easily with human influence (forest degradation). Indeed,
H. multipuncta appears to be synanthropic and invasive.
Thus, I believe coin spiders have a potential in Australasian
conservation efforts.
Biology
The most well known (ecologically and behaviourally)
species is H. papuana, owing to the research of Robinson
and colleagues (Robinson and Robinson 1978, 1980;
Robinson 1975, 1982; Robinson and Lubin 1979) conducted
at Wau, Papua New Guinea in the 1970s (MCZ vouchers
examined), briefly summarised below.
Habitat preferences
The species is obligately arboricolous. Although the webs
were seen on larger trees of coffee plantations, they were not
documented on rock faces, in contrast to H. multipuncta.
Web architecture
The web is adpressed to the tree surface, is eccentric (hub
in the upper portion) and is generally described as a ‘ladder
web’, meaning it is elongate with parallel sides and more or
less horizontal radii. The so-called ‘hub-cup’ is a retreat of
dense silk that touches the substrate, where the female rests
head-down (compare Figs 14–15, 25–26). Barrier webs and
stabilimenta are absent. The authors explained the side-to-
side curvature of the web plane with the so called pseudoradii
(Fig. 2), which are structurally similar to radii, but differ in
running parallel from top to bottom frame and thus offer
support for the sticky spiral, which follows the basic curvature
Revision of Herennia
Fig. 31. Herennia tone, sp. nov., female from Philippines: A, habitus, dorsal (holotype, he26);
B, same (he31), note prolateral long spines on first femur; C, epigynum, ventral (paratype, he32);
D, same (he31); E, epigynum, cleared, dorsal (paratype, he32). Scale bar E = 0.1 mm.
M. Kuntner426 Invertebrate Systematics
of the trunk. Like the ribs of an umbrella, the authors noted,
the pseudoradii give a curvature to the covering. This archi-
tecture is typical of adult females in their natural environment,
but juvenile spiders and captive adults build a more or less cir-
cular orb web, not necessarily against substrate.
Behaviour
Robinson and coauthors (Robinson and Robinson 1978,
1980; Robinson 1975, 1982; Robinson and Lubin 1979)
documented partial web renewal in H. papuana, a behaviour
typical of all nephilid genera. Web renewal was always done
at night. Egg sacs were attached to the bark of the tree, not in
the web (compare with H. multipuncta). Defence responses
involved raising and flexing of legs and jumping off the web.
The authors discussed the nocturnal habits of Herennia (the
spiders more readily attacked prey at night) as well as their
highly cryptic stance. The female dorsal colour is such that it
blends with the tree bark (and the web does not stand out).
Curiously, the spider’s ventral side is bright red (as for
H. multipuncta and H. etruscilla, which also have bright red
sterna, see Fig. 26B). The authors considered such coloura-
tion function problematical, especially since cage-raised
specimens were yellow (see below for a possible explana-
tion). Small reddish males cohabited in female webs, some-
times as eunuchs (lacking palpal bulbs, see below).
Predation
The authors found no preference for a particular prey
taxon or ecotype (the spiders generally avoided ants). The
predatory repertoire resembled that of Nephila, attacking all
prey with a bite (never attack wrapping). After a biting
attack, the spider either wrapped the prey in situ or pulled it
from the web at capture site. This was followed by the trans-
portation and attachment of prey (either in chelicerae or sus-
pended on silk) to the hub. The spiders sometimes attacked
by shuttling through the web at the hub-cup and approaching
the prey on the inner side of the web, with the dorsal side
facing the bark. Although the frequency or significance of
such ‘backside attack’ was not discussed, it may protect the
spider from predators. A similar ‘side flip’ as predator avoid-
ance is known in Clitaetra (Kuntner, in press), some Nephila
(M. Kuntner, unpublished data) and some Argiope
(M. Kuntner, unpublished data). In female Herennia, the
brightly coloured ventral side, which during backside attack
has to be exposed, could be interpreted as aposomatic.
Mating
Courtship and mating take place on the female web
(Robinson and Robinson 1978, 1980), the behaviour fitting in
a ‘group A type, which, in addition to Herennia, included
Nephila, Nephilengys and certain Argiope species (Robinson
Indian Ocean
Pacific Ocean
etruscilla
oz
papuana
milleri
agnarssoni
multipuncta
jernej
tone
sonja
deelemanae
gagamba
Fig. 32. Herennia species distribution map. The widespread, synanthropic and possibly invasive
H. multipuncta ranges from India and Sri Lanka to eastern Indonesia, but other species appear to be narrow
island endemics, likely confined to pristine forests.
Invertebrate Systematics 427
and Robinson 1978, 1980). Herennia papuana exhibits the
‘suitor phenomenon’ (male cohabiting with immature female)
and ‘eunuchs’. Up to seven males cohabit in the female web.
The only seemingly different behaviour from other nephilids
(e.g. Nephila) is the female access posture, where a receptive
female responds to male stimuli by raising her body partly out
of the hub-cup, thus allowing male access to her venter.
Other species
Web architecture of H. multipuncta (Figs 14–15) and
H. etruscilla (Figs 25, 26A) seems identical to H. papuana:
eccentric ladder webs closely following the transverse cur-
vature of tree trunks (or other substrates), with hub-cups
(Fig. 14AB, 25–26) and pseudoradii (Fig. 25A; the latter are
only known in Herennia). Both H. multipuncta (Fig. 14) and
H. etruscilla (Figs 25, 26A) commonly inhabit disturbed
habitats in addition to forests. The web and its varied location
in Sri Lanka were first described by Simon (1894: 758);
Herennia multipuncta webs can be on walls (Figs 14C, 15),
rock faces (Robinson and Lubin 1979; Murphy and Murphy
2000) as well as on trees (Fig. 14AB). Masumoto and
Okuma (1995) reported a tendency of H. multipuncta to
build on Eucalyptus trees in Malaysia, which supports the
interpretation that the species benefits from primary habitat
destruction. In Java H. etruscilla webs were found on trees in
rain forest (Kaliurang) and on (likely non-native) trees in
Cibodas botanical gardens (Figs 25–26A). According to
available data, both species are sympatric in Bogor, Java.
Egg sacs of H. etruscilla seem to be attached to the web
(Fig. 26A; compare with H. papuana).
Mating plugs and eunuchs
In most Herennia species, broken-off male embolic conduc-
tors were commonly found stuck in female copulatory open-
ings (Figs 10EF, 11, 17A, 18C, 22EF, 28B, 31D). Despite
this, only one male with broken embolic conductors was
found in collections. Instead, the so-called eunuch males are
common, which lack entire palpal bulbs (Fig. 23AE).
Eunuch males are also found in Nephilengys. Although the
function (if any) and frequency of severed bulbs are
unknown, it seems that male embolic conductor mutilation
and bulb amputation are correlated. In fact, Robinson and
Robinson (1980) found the evidence that Nephilengys males
deliberately lose the bulb after mating.
With its peculiar sexual biology, Herennia calls for
attention. The apparently consistent mutilation of male
sexual organs, which plug female genitals, and the bulb sev-
erance (eunuch phenomenon), known in Herennia and
Nephilengys, require explanations. A possible, though at this
point speculative, explanation of these behaviours is pre-
sented as three hypotheses consistent with the currently
available morphological and behavioural data: (1) the broken
embolic conductor may function as a mating plug preventing
future (successful) copulation attempts by other males;
(2) losing/removing the bulb after initial mutilation of the
embolic conductor is advantageous for the (sterile) male in
stopping hemolymph leakage; and (3) a eunuch may protect
his parental investment by fighting off rival males in search
of the female. These hypotheses could be tested with a
variety of approaches: morphological, molecular,
Revision of Herennia
Deinopis
Uloborus
Araneus
Argiope aurantia
Argiope argentata
Linyphia
Pimoa
Steatoda
Nesticus
Epeirotypus
Tetragnatha
Meta
Leucauge
Deliochus
Phonognatha graeffei
Clitaetra episinoides
Clitaetra irenae
Nephila fenestrata
Nephila pilipes
Nephilengys malabarensis
Nephilengys cruentata
Herennia multipuncta
Herennia etruscilla
Herennia deelemanae
Herennia papuana
Herennia tone
Herennia gagamba
Herennia agnarsson
i
97
99
73
50
100
47
56
35
42
50
90
87
28
19
31
58
67
48
64
85
81
60
83
100
90
7
1
1
1
1
-
5
4
3
4
>10
6
-
-
4
3
1
1
1
1
3
4
2
8
-
Deinopis
Uloborus
Araneus
Argiope aurantia
Argiope argentata
Linyphia
Pimoa
Steatoda
Nesticus
Epeirotypus
Tetragnatha
Meta
Leucauge
Deliochus
Phonognatha graeffei
Clitaetra episinoides
Clitaetra irenae
Nephila fenestrata
Nephila pilipes
Nephilengys malabarensis
Nephilengys cruentata
Herennia multipuncta
Herennia etruscilla
Herennia deelemanae
Herennia papuana
Herennia tone
Herennia gagamba
Herennia agnarssoni
A
B
Fig. 33. Phylogeny of the Nephilidae, with particular reference to Herennia. A, Strict consensus of ten shortest cladograms
(L = 490, CI = 40, RI = 70) collapsing four nodes. B, Successively weighted tree identical to one of the fundamental cladograms.
Branch supports given as bootstrap (above, regular type) and Bremer (below, bold).
M. Kuntner428 Invertebrate Systematics
behavioural, experimental and phylogenetic. In this analysis,
the eunuch phenomenon is ambiguously optimised: under
fast optimisation it is a synapomorphy for the clade Herennia
+ (Nephila + Nephilengys) with a reversal in Nephila,
whereas under slow optimisation the feature is hypothesised
to have evolved separately in Nephilengys and Herennia.
Acknowledgments
Views, opinions, interpretations and potential errors in this
paper are my own, not those of who have commented on and
criticised earlier drafts. I thank Jonathan Coddington and
Gustavo Hormiga for advice and help, Ingi Agnarsson and
Jeremy Miller for their daily help, encouragement and com-
ments, Jutta Schneider and Miquel Arnedo for discussing the
ideas and preliminary results from this study, and Marc
Allard, Jim Clark, Diana Lipscomb and Chris Thompson for
their comments on an early draft. The helpful comments of
Camilla Myers, Mark Harvey and Volker Framenau much
improved the paper. Fernando Alvarez-Padilla, Lara
Lopardo, Dana deRoche and Scott Larcher offered assis-
tance and help; Scott Whittaker and Patrick Herendeen pro-
vided SEM help, and Karie Darrow kindly helped with
digital image manipulation. Numerous curators, collection
managers and other biologists have assisted with loans (see
Materials and methods) and by collecting the specimens.
Erik J. van Nieukerken kindly helped translate Doleschall’s
text and find his original artwork. Gustavo Hormiga kindly
provided his unpublished photograph. The fieldwork in
Indonesia was done jointly with Irena Kuntner and Matjaž
Bedjanič. This project was supported by the USA National
Science Foundation (PEET grant DEB-9712353 to Hormiga
and Coddington) and partly by the OTS-STRI-Mellon
Research Exploration Award (to Kuntner and Šereg). I
further acknowledge material and financial support of the
George Washington University, Smithsonian Institution, the
Ministry of Science of the Republic of Slovenia and the
Biological Institute of the Slovenian Academy of Sciences
and Arts. At the latter institution, the support (1999–2001)
was coordinated by Rajko Slapnik, and the final stages
endorsed by Branko Vreš and Oto Luthar. This project would
Wallace
Huxley
Wallace
Huxley
Herennia
Herennia
multipuncta
etruscilla
deelemanae
gagamba
tone
papuana
agnarssoni
Fig. 34. Herennia phylogeography. At least two dispersal events have to be postulated over
Wallace’s and Huxley’s line, respectively, whereas the newly drawn ‘Herennia line’, dividing
Australasia from the core Oriental region, is only crossed by a single species, H. multipuncta. See text
for details.
Invertebrate Systematics 429
have been impossible without the support of my wife Irena,
my parents Sonja and Tone, and my brother Jernej.
References
Bonnet, P. (1957). ‘Bibliographia Araneorum, Vol. 2, Part 3, (F–M).
(Douladoure: Toulouse, France.)
Bremer, K. (1988). The limits of amino acid sequence data in
angiosperm phylogenetic reconstructions. Evolution 42, 795–803.
Bremer, K. (1994). Branch support and tree stability. Cladistics 10,
295–304.
Brignoli, P. M. (1983). ‘A Catalogue of the Araneae Described Between
1940 and 1981. (Manchester University Press in association with
The British Arachnological Society: Manchester, UK.)
Brown, J. H., and Lomolino, M. V. (1998). ‘Biogeography. 2nd edn.
(Sinauer Associates: Sunderland, MA, USA).
Cantino, P. D., and de Queiroz, K. (2004). ‘PhyloCode: A Phylogenetic
Code of Biological Nomenclature, Version 2b. Available online at:
http://www.ohiou.edu/phylocode/ (verified November 2005).
Chrysanthus, F. (1971). Further notes on the spiders of New Guinea I
(Argyopidae). Zoologische Verhandelingen 113, 1–52.
Coddington, J. A. (1983). A temporary slide-mount allowing precise
manipulation of small structures. Verhandlungen des
Naturwissenschaftlischen Vereins in Hamburg, N.S. 26, 291–292.
Coddington, J. A. (1989). Spinneret silk spigot morphology: evidence
for the monophyly of orb-weaving spiders, Cyrtophorinae
(Araneidae), and the group Theridiidae–Nesticidae. Journal of
Arachnology 17, 71–95.
Coddington, J. A. (1990). Ontogeny and homology in the male palpus
of orb weaving spiders and their relatives, with comments on phylo-
geny (Araneoclada: Araneoidea, Deinopoidea). Smithsonian
Contributions to Zoology 496, 1–52.
Coddington, J. A., Hormiga, G., and Scharff, N. (1997). Giant female or
dwarf male spiders? Nature 385, 687–688. doi:10.1038/385687a0
Colwell, R. K. (1999). ‘BIOTA: The Biodiversity Database Manager,
Version 1.6.0. (Sinauer Associates: Sunderland, MA, USA).
Dahl, F. (1912). Seidenspinne und Spinnenseide. Mitteilungen aus dem
Zoologischen Museum in Berlin 6, 1–90.
Davies, V. T. (1988). An illustrated guide to the genera of orb-weaving
spiders in Australia. Memoirs of the Queensland Museum 25,
273–332.
Doleschall, C. L. (1859). Tweede Bijdrage tot de Kennis der Arachniden
van den Indischen Archipel. Verhandelingen der Natuurkundige
Ve r eeniging in Nederlandsch Indië 5, 1–60.
Eberhard, W. G. (1982). Behavioral characters for the higher classi-
fication of orb-weaving spiders. Evolution; International Journal of
Organic Evolution 36, 1067–1095.
Farris, J. S. (1969). A successive approximations approach to character
weighting. Systematic Zoology 18, 374–385.
Felsenstein, J. (1985). Confidence limits on phylogenies: an approach
using the bootstrap. Evolution 39, 783–791.
Fitch, W. M. (1971). Towards defining the course of evolution: minimal
change for a specific tree topology. Systematic Zoology 20,
406–416.
Goloboff, P. A. (1993). ‘NONA, Version 2.0. Available online at:
http://www.cladistics.com/ (verified November 2005).
Griswold, C. E., Coddington, J. A., Hormiga, G., and Scharff, N. (1998).
Phylogeny of the orb-web building spiders (Araneae, Orbiculariae:
Deinopoidea, Araneoidea). Zoological Journal of the Linnean
Society 123, 1–99. doi:10.1006/zjls.1997.0125
Holm, Å. (1979). A taxonomic study of European and East African
species of the genera Pelecopsis and Trichopterna (Araneae,
Linyphiidae) with descriptions of a new genus and two new species
of Pelecopsis from Kenya. Zoologica Scripta 8, 255–278.
Hormiga, G., Eberhard, W. G., and Coddington, J. A. (1995). Web-con-
struction behaviour in Australian Phonognatha and the phylogeny
of nephiline and tetragnathid spiders (Araneae: Tetragnathidae).
Australian Journal of Zoology 43, 313–364. doi:10.1071/
ZO9950313
Hormiga, G., Scharff, N., and Coddington, J. A. (2000). The phylo-
genetic basis of sexual size dimorphism in orb-weaving spiders
(Araneae, Orbiculariae). Systematic Biology 49, 435–462.
doi:10.1080/10635159950127330
International Comission on Zoological Nomenclature (1999).
‘International Code of Zoological Nomenclature. 4th edn.
(International Trust for Zoological Nomenclature: London, UK.)
Karsch, F. (1880). Arachnologische Blätter (Decas I). Zeitschrift für die
Gesammten Naturwissenschaften 53, 373–409.
Kuntner, M. (2002). The placement of Perilla (Araneae, Araneidae)
with comments on araneid phylogeny. The Journal of Arachnology
30, 281–287.
Kuntner, M. (2003). The systematics of nephiline spiders (Araneae,
Tetragnathidae). American Arachnology 66, 9.
Kuntner, M. (in press). Phylogenetic systematics of the Gondwanan
nephilid spider lineage Clitaetrinae (Araneae, Nephilidae).
Zoologica Scripta.
Kuntner, M., and Hormiga, G. (2002). The African spider genus
Singafrotypa (Araneae, Araneidae). The Journal of Arachnology 30,
129–139.
Levi, H. W. (1980). The orb-weaver genus Mecynogea, the subfamily
Metinae and the genera Pachygnatha, Glenognatha and Azilia of the
subfamily Tetragnathinae north of Mexico (Araneae: Araneidae).
Bulletin of the Museum of Comparative Zoology 149, 1–75.
Levi, H. W. (1986). The neotropical orb-weaver genera Chrysometa and
Homalometa (Araneae: Tetragnathidae). Bulletin of the Museum of
Comparative Zoology 151, 91–215.
Levi, H. W., and von Eickstedt, V. R. D. (1989). The Nephilinae spiders
of the Neotropics (Araneae: Tetragnathidae). Memorias do Instituto
Butantan 51, 43–56.
Masumoto, T., and Okuma, C. (1995). Specific web building on euca-
lyptus trees in Herennia ornatissima (Araneae: Tetragnathidae).
Acta Arachnologica 44, 171–172.
Murphy, F., and Murphy, J. (2000). ‘An Introduction to the Spiders of
South East Asia. (Malaysian Nature Society: Kuala Lumpur,
Malaysia.)
Nixon, K. (2002). ‘WinClada, Version 1.00.08. Available online at:
http://www.cladistics.com/ (verified November 2005).
Pan, H. C., Zhou, K. Y., Song, D. X., and Qiu, Y. (2004). Phylogenetic
placement of the spider genus Nephila (Araneae: Araneoidea)
inferred from rRNA and MaSp1 gene sequences. Zoological
Science 21, 343–351. doi:10.2108/zsj.21.343
Platnick, N. I. (1989). ‘Advances in Spider Taxonomy: A Supplement to
Brignoli’s A Catalogue of the Araneae Described between 1940 and
1981. (Manchester University Press: Manchester, UK.)
Platnick, N. I. (1993). ‘Advances in Spider Taxonomy 1988–1991: With
Synonymies and Transfers 1940–1980. (New York Entomological
Society in association with the American Museum of Natural
History: New York, USA.)
Platnick, N. I. (1997). ‘Advances in Spider Taxonomy 1992–1995: With
Redescriptions 1940–1980. (New York Entomological Society in
association with the American Museum of Natural History: New
York, USA.)
Platnick, N. I. (2005). ‘The World Spider Catalog, Version 5.5.
Available online at: research.amnh.org/entomology/spiders/catalog/
(verified November 2005).
Robinson, M. H. (1975). The evolution of predatory behaviour in
araneid spiders. In ‘Function and Evolution in Behavior’.
(Eds G. Baerends, C. Beer and A. Manning.) pp. 292–312.
(Clarendon Press: Oxford, UK.)
Revision of Herennia
M. Kuntner430 Invertebrate Systematics
Robinson, M. H. (1982). Courtship and mating behavior in spiders.
Annual Review of Entomology 27, 1–20. doi:10.1146/annurev.en.
27.010182.000245
Robinson, M. H., and Lubin, Y. D. (1979). Specialists and generalists:
the ecology and behavior of some web-building spiders from Papua
New Guinea, 1. Herennia ornatissima, Argiope ocyaloides and
Arachnura melanura (Araneae: Araneidae). Pacific Insects 21,
97–132.
Robinson, M. H., and Robinson, B. (1978). The evolution of courtship
systems in tropical araneid spiders. Symposia of the Zoological
Society of London 42, 17–29.
Robinson, M. H., and Robinson, B. (1980). Comparative studies of the
courtship and mating behavior of tropical araneid spiders. Pacific
Insects Monograph 36, 35–218.
Roewer, C. F. (1942). ‘Katalog der Araneae von 1758 bis 1940, bzw.
1954., Vol. 1. (Paul Budy: Bremen, Germany.)
Roth, V. D., and Roth, B. M. (1984). A review of appendotomy in
spiders and other arachnids. Bulletin (British Arachnological
Society) 6, 137–146.
Scharff, N., and Coddington, J. A. (1997). A phylogenetic analysis of
the orb-weaving spider family Araneidae (Arachnida, Araneae).
Zoological Journal of the Linnean Society 120, 355–434.
doi:10.1006/zjls.1996.0084
Simon, E. (1894). ‘Histoire Naturelle des Araignées. Vol. 1.
pp. 489–760. (Roret: Paris, France.)
Stoliczka, F. (1869). Contribution towards the knowledge of Indian
Arachnoidea. Journal of the Asiatic Society of Bengal 38, 201–251.
Thorell, T. (1877). Studi sui ragni Malési e Papuani. I. Ragni di Selebes
raccolti nel 1874 dal Dott. O. Beccari. Annali del Museo Civico di
Storia Naturale ‘Giacomo Doria’. Genova 10, 341–634.
Thorell, T. (1878). Studi sui ragni Malési e Papuani. II. Ragni di
Amboina raccolti Prof. O. Beccari. Annali del Museo Civico di
Storia Naturale ‘Giacomo Doria’. Genova 13, 1–317.
Thorell, T. (1881). Studi sui ragni Malési e Papuani. III. Ragni
dell’Austro Malesia e del Capo York, conservati nel Museo civico di
storia naturale di Genova. Annali del Museo Civico di Storia
Naturale,’Giacomo Doria’. Genova 17, 7–27.
Thorell, T. (1887). Viaggio di L. Fea in Birmania e regioni vicine. II.
Primo saggio sui Ragni birmani. Annali del Museo Civico di Storia
Naturale,’Giacomo Doria’. Genova 2, 5–417.
Thorell, T. (1890). Studi sui ragni Malesi e Papuani. IV, 1. Annali del
Museo Civico di Storia Naturale,’Giacomo Doria’. Genova 28,
1–419.
Tikader, B. K. (1982). Spiders: Araneae, Vol. II, Part 1: family
Araneidae (=Argiopidae), typical orbweavers. In ‘The Fauna of
India’. pp. 1–293. (Zoological Survey of India: Calcutta, India.)
Wunderlich, J. (1986). ‘Spinnenfauna gestern und heute. Fossile
Spinnen in Bernstein und ihre heute lebenden Verwandten. (Erich
Bauer, Quelle & Meyer: Wiesbaden, Germany.)
Wunderlich, J. (2004). Fossil spiders in amber and copal. Conclusions,
revisions, new taxa and family diagnoses of fossil and extant taxa.
Beiträge zur Araneologie 3A-B, 1–1908.
Manuscript received 14 June 2005, revised and accepted 23 September
2005.
Invertebrate Systematics 431Revision of Herennia
Appendix 1. List of characters and character states used in the phylogenetic analysis, with description of characters relevant for Herennia
A rigorous homology test of these and additional characters relevant for nephilids will be presented elsewhere (Kuntner, Hormiga and
Coddington, unpublished data)
(1) Female cephalic region: 0, low (Fig. 1A); 1, conspicuously high.
(2) Female carapace: 0, piriform (Fig. 1C); 1, oval with wide head region.
(3) Female carapace edge: 0, smooth; 1, ridged (Figs 1C, 5B, D).
(4) Female carapace edge: 0, glabrous or with few hair-like setae; 1, with an extensive row of hair-like setae (Fig. 5BE).
(5) Female median pair of prosomal tubercles: 0, absent (Figs 1AC, 5B); 1, present.
(6) Fovea (female carapace): 0, inconspicuous (Figs 1A, C, 5B, 19A); 1, pronounced.
(7) Female carapace macrospines: 0, absent; 1, present.
Note: in all Nephilengys species stout erect macrospines aer present on carapace.
(8) Female carapace warts: 0, absent; 1, present (Figs 5F, 19AF).
Note: the carapace cuticle is warty (enlarged functional and non-functional setal bases) in all Herennia species, especially pronounced in H.
etruscilla (Fig. 19).
(9) Female carapace V-mark: 0, absent; 1, present (Figs 1C, 14, 27A, C).
(10) Female carapace hair-like setae: 0, present (Figs 5, 19); 1, absent.
Note: apart from regular setae nephilids (and certain outgroups) have thin short white hair-like setae.
(11) Female median eye region: 0, rounded; 1, median eyes on a tubercle (Figs 1BC, 5C, 19E).
(12) Female lateral eye region: 0, lateral eyes on separate tubercles; 1, lateral eyes on a single tubercle (Figs 1BC, 5C, 19E); 2, rounded.
(13) Female LE separation from ME: 0, not widely separated (Figs 1BC, 5C, 19E); 1, widely separated.
Note: ratio of the distance between the PLE and PME (at its widest point) divided by the width of the PME ocular area (at the widest point).
If the ratio is less than 1, the separation is normal; if the ratio is more than 1, the separation is wide.
(14) Posterior eye row (dorsal view): 0, straight to recurved (Figs 1C, 5BC, 19A, D, E); 1, procurved.
Note: the character is from Scharff and Coddington (1997: character 54).
(15) Female PME: 0, less than one PME diameter apart; 1, one PME diameter or more apart (Figs 1C, 5BC, 19A, D, E).
(16) Female PLE size: 0, equal or less than PME (Fig. 19E); 1, larger than PME.
(17) PME canoe tapetum: 0, absent; 1, full; 2, narrow.
Note: corresponds to character 4 in Hormiga et al. (1995), characters 51 and 52 in Scharff and Coddington (1997) and characters 28 and 29
in Griswold et al. (1998). Herennia lacks eye tapeta.
(18) PLE canoe tapetum: 0, absent; 1, full; 2, narrow.
Note: as above.
(19) Female clypeus height: 0, low (less than three AME diameters); 1, equal or more than three AME diameters.
(20) Endites: 0, very long (> 2 × width); 1, short (length < 2 × width).
(21) Labium and sternum: 0, separate; 1, fused.
(22) Female sternum: 0, longer than wide; 1, as wide as or wider than long.
(23) Sternal slit sensilla: 0, present (Figs 6EF, 20F); 1, absent.
(24) Female sternum colour pattern: 0, inconspicuously coloured; 1, uniformly orange/red (Fig. 26B); 2, medially dark, laterally pale; 3,
medially light, laterally dark; 4, with yellow spots corresponding to tubercles.
(25) Sternal white pigment: 0, absent; 1, present.
(26) Female sternal tubercle I: 0, absent; 1, present.
Note: paired elevations of the female sternum adjacent to coxae I-IV are termed sternal tubercles (Fig. 6E – arrows).
(27) Female sternal tubercle II: 0, absent; 1, present (Fig. 6E – arrows).
(28) Female sternal tubercle III: 0, absent; 1, present (Fig. 6E – arrows).
(29) Female sternal tubercle IV: 0, absent; 1, present (Fig. 6E – arrows).
(30) Female frontal sternal tubercle: 0, absent; 1, present.
(31) Female chilum: 0, absent; 1, present.
Note: The chilum is present as a paired sclerite (Fig. 1B) at the base of chelicerae, just under the clypeus in most nephilids (not in certain
Clitaetra species).
(32)