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smithsonian contributions to zoology • number 636

Systematics of the Spider Family 

Deinopidae with a Revision

of the Genus Menneus

Jonathan A. Coddington, 

Matjaž Kuntner, and 

Brent D. Opell
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smithsonian contributions to zoology • number 636

Systematics of the Spider Family 

Deinopidae with a Revision 

of the Genus Menneus

Jonathan A. Coddington, 

Matjaž Kuntner, and 

Brent D. Opell

washington d.c.

2012
ABSTRACT

Coddington, Jonathan A., Matjaž Kuntner, and Brent D. Opell. Systematics of the spider family Deinopidae with a 

revision of the genus Menneus. Smithsonian Contributions to Zoology, number 636, iv + 61 pages, 32 figures, 1 table, 

2012. The enigmatic and rare spiders of the family Deinopidae are known for their extremely large ocelli and their 

unique habit of casting their small cribellate webs over nocturnal pedestrian or aerial prey. Although the monophyly 

of the family has not been controversial, deinopid descriptive systematics has been neglected since the original species 

descriptions. The first goal of this monograph is to test the monophyly of Deinopidae and its genera and thus to establish a phylogenetically informed taxonomy of the species. We provide the first phylogeny of this family on the basis 

of 53 morphological and 3 behavioral characters scored for 17 deinopid species and 2 outgroups. Extant deinopid 

spiders belong to two genera: the larger pantropical Deinopis MacLeay, 1839, diagnosed by the extreme size of their 

posterior median eyes, and the smaller Menneus Simon, 1876, of the Old World (sub)tropics with normally sized eyes. 

Avella Pickard-Cambridge, 1877, and Avellopsis Purcell, 1904, are junior synonyms of Menneus as their type species 

are nested throughout Menneus phylogeny. The second goal is to revise the non-Deinopis species of Deinopidae. Here, 

we recognize, diagnose, illustrate, describe, and phylogenetically place the 14 known Menneus species. Africa currently 

has six species: Menneus camelus Pocock, 1902, from South Africa; M. capensis (Purcell, 1904) new combination from 

Western Cape, South Africa; M. darwini new species from Tanzania; M. dromedarius Purcell, 1904 (removed from 

synonymy of M. camelus), from South Africa and Madagascar; M. samperi new species from Kenya, Tanzania, and 

Uganda; and M. tetragnathoides Simon, 1876 (M. affinis Tullgren, 1910 new synonym), from Angola, Malawi, and 

Tanzania. Australasia currently has eight species: M. aussie new species from eastern Australia and New Caledonia; M. 

bituberculatus new species from Queensland and Indonesian West Papua; M. superciliosus (Thorell, 1881) from eastern 

Australia; M. nemesio new species from New South Wales; M. neocaledonicus (Simon, 1889) from New Caledonia, 

M. quasimodo new species from Western Australia; M. trinodosus Rainbow, 1920 (Deinopis insularis Rainbow, 1920, 

new synonym), from Queensland and New South Wales; and M. wa, new species, from Western Australia. We propose 

Avella angulata L. Koch, 1878, Avella despiciens O. P.-Cambridge, 1877, and Avella unifasciata L. Koch, 1878, as 

nomina dubia. Paleomicromenneus lebanensis Penney, 2003, the earliest known fossil, is similar to Menneus. Menneus

is phylogenetically sister to Deinopis, and within Menneus we recover a grade of Australasian species that includes the 

African M. capensis, followed by a distal clade of African species with peculiar asymmetric somatic morphology.

Cover images, from left to right: Menneus camelus female with web from South Africa (photo by M. Kuntner); Deinopis spinosa female from Florida, USA (photo by J. A. Coddington); Menneus capensis female with web from South 

Africa, (photo by M. Kuntner).

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012

www.scholarlypress.si.edu

Compilation copyright © 2012 Smithsonian Institution

Figures 3–32 are in the public domain. The rights to all other text and images in this publication, including cover and interior designs, 

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Library of Congress Cataloging-in-Publication Data

Coddington, Jonathan A.

Systematics of the spider family Deinopidae, with a revision of the genus Menneus / Jonathan A. Coddington, Matjaž Kuntner, and 

Brent D. Opell.

 p. cm. — (Smithsonian contributions to zoology ; no. 636)

Includes bibliographical references.

1. Deinopidae—Classification. 2. Menneus—Classification. I. Kuntner, Matjaž, 1971– II. Opell, Brent D. III. Title. 

QL458.42.D55C63 2012

595.4’4—dc23

2011023133

ISSN: 0081-0282 (print); 1943-6696 (online)

The paper used in this publication meets the minimum requirements of the American National Standard for Permanence 

of Paper for Printed Library Materials Z39.48–1992.
Contents

INTRODUCTION 1

METHODS AND MATERIALS 3

Abbreviations 3

Acknowledgments 4

Taxonomy 4

Illustrations and Imaging 5

Phylogenetic Methods 5

MORPHOLOGY 6

PHYLOGENY 8

Character Descriptions 8

 Female Somatic Morphology 8

 Female Genital Morphology 9

 Male Somatic Morphology 9

 Male Genital Morphology 9

 Behavior 10

RESULTS 11

TAXONOMY 11

Deinopidae C. L. Koch, 1850 11

Deinopis MacLeay, 1839 11

Paleomicromenneus Penney, 2003 12

Menneus Simon, 1876 12

KEY TO MENNEUS SPECIES 13

African Species 14

 Menneus camelus Pocock, 1902 14

 Menneus capensis (Purcell, 1904), new combination 15

 Menneus darwini, new species 16

 Menneus dromedarius Purcell, 1904 17

 Menneus samperi, new species 18

 Menneus tetragnathoides Simon, 1876 18
i v • smithsonian contributions to zoology

Australasian Species 19

 Menneus aussie, new species 19

 Menneus bituberculatus, new species 20

 Menneus superciliosus (Thorell, 1881) 20

 Menneus nemesio, new species 22

 Menneus neocaledonicus (Simon, 1889) 22

 Menneus quasimodo, new species 23

 Menneus trinodosus Rainbow, 1920 24

 Menneus wa, new species 25

 Menneus pietrzeniukae Wunderlich, 2004 25

Nomina dubia 26

 Avella angulata L. Koch, 1878 26

 Avella despiciens O. P.-Cambridge, 1877 26

 Avella unifasciata L. Koch, 1878 26

FIGURES 27

REFERENCES 59
Systematics of the Spider Family Deinopidae 

with a Revision of the Genus Menneus

INTRODUCTION

The “ogre-faced” spiders of the family Deinopidae are known for their unusual morphology, natural history, and rarity. MacLeay personally discovered 

and described the first deinopid in 1839, writing (p. 10), “I found Deinopis, 

with the last mentioned genus [Ctenus] and Dolomedes, under stones in the 

island of Cuba. It must be assigned to the Wolfspinnen of Koch, but it is very 

unlike any of them hitherto known.” MacLeay must have meant the lower faces 

of large boulders, where it would not be unusual to find Ctenus, Dolomedes, 

or, indeed, Deinopis spinning a web positioned to envelope prey walking on the 

stone surface. However, to temperate biologists used to turning over small rocks 

in search of cursorial spiders, MacLeay’s description probably conveyed entirely 

the wrong impression (or perhaps MacLeay entirely misremembered the circumstances). Lacking any further biological data, several generations of arachnologists continued to speculate that deinopids were a branch of cursorial hunting 

spiders (e.g., O. P. Cambridge, 1877; Thorell, 1881, 1886). Doleschall (1859) 

reported that Deinopis were collected from a “large irregular web spun between 

trees, in the center of which the spider sits with the anterior legs extended along 

the body axis” (as paraphrased by Simon, 1892:209). Marx (1889: 341) added 

to the confusion by reporting that the discoverer of D. spinosa, W. S. Barnard, 

remembered it as being “on a large horizontal web, which resembled very much 

that of Agalena, but he could not find a tubular retreat attached to the web, as 

is the case with Agalena.”

Akerman (1926), studying Menneus, was the first to describe the 1–2 cm 

sized web (Figures 1a,d, 2b–d) and the remarkable attack behavior in which the 

animal grasps the corners of the web with the anterior tarsi and lunges (actually drops) toward prey to envelop it with the extremely extensible web. Since 

Akerman, Baum (1937), Roberts (1954), Clyne (1967, 1968), Mascord (1970), 

Robinson and Robinson (1971), Theuer (1954), Austin and Blest (1979), 

Jonathan A. Coddington, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 

20013-7012, USA; Matjaž Kuntner, Institute of 

Biology, Scientific Research Centre, Slovenian 

Academy of Sciences and Arts, Novi trg 2, P.O. 

Box 306, SI-1001, Ljubljana, Slovenia; Brent D. 

Opell, Department of Biological Sciences, Virginia Polytechnic Institute and State University, 

Blacksburg, Virginia 24061, USA. Correspondence: coddington@si.edu. Manuscript received 

3 December 2009; accepted 21 April 2011.
2 • smithsonian contributions to zoology

Coddington (1986a, 1986b), Coddington and Sobrevila 

(1987), Larsen (1992), and Getty and Coyle (1996) all observed the same specialized architecture and attack behavior. Theuer (1954: 37) studied D. spinosa extensively and 

saw the same web and behavior in animals that he felt sure 

were “no more than 3–4 days old.” However, Mascord 

(1982:5) wrote, “I photographed recently a small net at 

my home which was only 5mm wide by 3.4mm deep, and, 

although it was similar in appearance, it does not have 

the cribellate strands of an adult net,” and Raven (1992) 

offered further details on newborn Deinopis biology. 

The web intrigued biologists since its discovery, but not 

until Eberhard (1982) clarified the basic ethological apomorphies defining the orb web did Coddington (1986a) 

confirm that the deinopid architecture was plausibly homologous (Figure 2d). Since then deinopids have figured 

prominently in studies of spinnerets (Peters, 1982; Coddington, 1989), the comparative morphology of silks (Eberhard and Pereira, 1993), and exemplars in phylogenetics 

(reviewed in Coddington et al., 2004; Coddington, 2005).

Deinopid morphology was similarly enigmatic to classical authors. C. L. Koch (1850:41) erected a new family 

for Deinopis, but that, of course, did not solve the problem 

of their relationships (although he suggested Salticidae). 

Subsequent authors mostly suggested affinities with other, 

more or less large-eyed families such as lycosids, pisaurids, 

oxyopids, or even salticids, or cribellates such as hypochilids, eresids, or agelenids. Simon described Menneus in 

1876 and immediately realized from the distinctive palpal 

morphology that Deinopis was closely related but made no 

comment concerning the placement of the pair. O. P. Cambridge (1877:558) said, “Dinopides, including the genus 

Avella described below, contains now three genera and is 

an exceedingly remarkable group; and I confess that its systematic position is, as yet, a puzzle to me.” Walckenaer, as 

well as MacLeay, who founded it, placed “Dinopis in the 

family Lycosides, between which and the Salticides I have 

at present left it, though that place will probably not long 

retain it. I cannot, however, agree with Dr. L. Koch, who 

places Dinopis in the family Eresides. . . . I would rather 

suggest that its place is near to the Agelenides, though in 

the absence of any knowledge of its habits it is impossible 

to speak with certainty upon this point.” Convinced of 

the monophyly of the Cribellatae, Simon (1892:205–210) 

placed deinopids as a subfamily of Uloboridae, but his reasoning was essentially phenetic rather than phylogenetic. 

Berland (1932:373) followed Simon, and based on Akerman’s observations directly related the deinopid web to the 

uloborid orb. Petrunkevitch (1923, 1928, 1933) disagreed 

only as to rank, restoring Dinopinae [sic] to its own family. 

Millot (1933) strongly supported Dinopidae because of the 

many differences in internal anatomy with Uloboridae, and 

since then the group has usually been regarded as a distinct 

family (Platnick, 2010).

Deinopis, of course, is well known for its huge posterior median eyes (Figure 3a). In photographic terms, 

the eyes are about 0.77 mm f/0.58 lenses, thus extremely 

wide angle, and extremely “fast,” or sensitive to light 

(Blest and Land, 1977). They considered deinopid posterior median ocelli to be the largest of any invertebrate: 

the largest they measured in D. subrufus were 1.40 mm 

in diameter, and subrufus is not a particularly large species. They estimated that Deinopis posterior median eyes 

gathered proportionately about 2000× the amount of light 

as a diurnal eye, either human or salticid. The physiology 

of the posterior median eyes is as impressive as their form. 

Blest and coauthors published a series of papers (Blest and 

Land, 1977; Blest, 1978; Blest et al., 1978a, 1978b, 1979, 

1980; Laughlin et al., 1980; Blest and Price, 1981) that 

showed that Deinopis destroyed most of the photosensitive membrane in these eyes at dawn and resynthesized it 

at dusk, saying, “Dinopis might almost be said to make a 

new retina every day” (Blest, 1978:479). Clearly, the enlarged eyes in Deinopis facilitate visual predation under 

dark conditions.

Deinopids also deserve to be better known for their 

extraordinary genitalia. Many authors have remarked 

on the distinctive male round tegula, round alveoli, and 

spiraled emboli (Figure 4a–e). Females are comparably 

equipped (see Figure 9d). As we explain further below, 

these apomorphies apparently enable extreme rotation 

of the bulb during copulation. Although not illustrated 

here, some Deinopis females have more than 20 twists or 

coils in the copulatory ducts, and their males may have 

correspondingly spiraled emboli (see, e.g., Kuntner et al., 

2008: fig. 20). Although no one has studied the detailed 

sexual biology of any species, the central median apophysis probably anchors the bulb in the epigynum and does 

not rotate. Neither, obviously, does the cymbium. Instead 

the tough, highly spiraled median apophysis membrane 

and basal hematodocha must unwind as many times as 

the male embolus rotates itself into the female copulatory 

ducts. Multicellular animals famously have never evolved 

true axles, wheels, or even repetitive rotation of body 

parts, but Deinopis may hold the world record for trying.

Despite their circumtropical distribution and notoriety, 

deinopids are unusually rare in collections and apparently 

in the field, considering that they are large (body length 

1–2 cm), free-living animals favoring habitats that naturalists and collectors intensely sample (Figures 1, 2). Judging 
number 636 • 3

from collections borrowed for this study, the current global 

total for Deinopidae is about 300 adult specimens of only 

57 nominal species (Platnick, 2010). The 14 species of the 

genus Menneus, the main subject of this paper, are known 

from only 135 adult animals from 106 distinct localities 

worldwide. By the beginning of the twentieth century, only 

two Menneus adults had been collected, and, by 1950, 

only 15. From 1950 until the present, however, 122 adults 

have been collected, possibly because naturalists now realize the importance of collecting at night (see Figure 8b). If 

one supposes that all species with viable populations have 

ranges large enough so that taxonomists would regard 

them as occupying at least two “localities,” one may apply 

“Chao2” estimators (Colwell, 2008) to obtain a rough 

lower-bound estimate of the actual richness of a clade or 

lineage. Four of the 14 Menneus species are known from 

single localities, and two are known from only two, yielding an estimate of clade richness of 30 species, or roughly 

double the currently known total. Of course, such statistical estimators assume random constant sampling effort, 

which is clearly false (Figure 8b), but the method at least 

provides an approximation of the magnitude of undiscovered Menneus diversity.

As a family, Deinopidae is comparably rare. Average 

abundance in quantitative samples from either the Old 

World or New World tropics (Cameroon, Madagascar, 

Thailand: Coddington, unpublished data; Tanzania: Sorensen et al., 2002; Peru: Silva and Coddington, 1996; Bolivia: Coddington et al., 1991, 1996; Guyana: Coddington 

et al., 2009; Tobago: Coddington, unpublished data) is 

0.9 adults per 1,000 individuals—among the lowest local 

abundances of any families from these inventories.

Because Deinopidae are cribellate and spin orb webs, 

the family has featured prominently in the controversy 

over the monophyly of the orb web (Opell, 1979; Coddington, 1986a, 1986b; Coddington and Levi, 1991; 

Bond and Opell, 1998; Coddington et al., 2004). Ethological and morphological evidence to date places Deinopidae consistently as the sister group of Uloboridae, forming 

Deinopoidea, and the latter as sister to the ecribellate, 

primitively orb-spinning Araneoidea (Coddington, 1986a, 

1986b, 1990a, 1990b; Scharff and Coddington, 1997; 

Griswold et al., 1998, 1999, 2005). Some molecular analyses, however, have not recovered a monophyletic Orbiculariae (e.g., Wu et al. (2002), but see Garb et al. (2006) and 

Blackledge et al. (2009) who find support for the group).

This paper revises the genus Menneus worldwide and 

establishes the generic classification of Deinopidae. It also 

provides a morphologically and ethologically based estimate of the species level phylogeny of the former and 

cladistically based definitions of relevant higher taxa. We 

expect to follow this paper with a worldwide revision of 

Deinopis, but, given the length of time this study has been 

underway, it seems worthwhile to publish these results 

now in order to facilitate Menneus species identification.

Classically, Menneus is a genus of southern Africa 

and Australia. Earlier authors recognized 13 species (Platnick, 2010). Six species were from Africa: camelus from 

KwaZulu-Natal, dromedarius and capensis (as Avellopsis) 

from the Cape Province, South Africa, affinis from East 

Africa, and tetragnathoides from the Congo. Seven species 

were from Australia, New Guinea, and adjacent islands 

(mostly treated as Avella): angulatus and superciliosus from 

Queensland, despiciens from Queensland and New South 

Wales, unifasciatus from New South Wales, neocaledonicus from New Caledonia, reticulatus from New Guinea, 

and trinodosus from Lord Howe Island. One fossil species, 

M. pietrzeniukae, is known from Baltic Amber but seems 

to be a juvenile female (Wunderlich, 2004; Dunlop et al., 

2009). Owing mainly to the tendency of nineteenth century 

authors to describe as new species juveniles of obviously 

new lineages, such as Menneus, many older names are nomina dubia. Deinopid species are surprisingly difficult to 

diagnose, even with genitalic characters. Species concepts 

based only on somatic features are best suppressed.

METHODS and Materials

A total of 135 adult Menneus specimens were examined, representing 113 species records. Localities are listed 

as written on museum labels (usually direct quotes) without interpretation of abbreviations. In square brackets 

we provide the collection acronym, the specimen code (if 

any), count, and sex of specimens, as well as decimal latitude and longitude.

Abbreviations

The following museum abbreviations are used subsequently throughout the text, and we thank the curators, 

collection managers, and associated staff for their help in 

providing specimens and loans: 

Museums

AMNH American Museum of Natural 

History, New York, USA

AMS Australian Museum, Sydney, Australia

BMNH The Natural History Museum, 

London, UK
4 • smithsonian contributions to zoology

CAS California Academy of Sciences, San 

Francisco, USA

EMUC Essig Museum University of 

California, Berkeley, USA

Filmer Martin Filmer Private Collection, 

Johannesburg, South Africa

MCSNG Museo Civico di Storia Naturale, 

“Giacomo Doria,” Genoa, Italy

MCZ Museum of Comparative Zoology, 

Harvard University, Cambridge, 

Massachusetts, USA

MNHN Muséum National d’Histoire 

Naturelle, Paris, France

Murphy Coll. John Murphy Private Collection, 

Hampton, UK

NMP Natal Museum, Pietermaritzburg, 

South Africa

QM Queensland Museum, Brisbane, 

Australia

RMCA Musée Royal de l’Afrique Centrale, 

Tervuren, Belgium

SAM South African Museum, Cape Town, 

South Africa

SANC South African National Collection, 

Pretoria, South Africa

SMNH Swedish Museum of Natural History, 

Stockholm, Sweden

TM Transvaal Museum Pretoria, South 

Africa

USNM National Museum of Natural History, 

Smithsonian Institution, Washington, 

D.C., USA 

WAM Western Australian Museum, Perth, 

Australia

ZMUC Zoological Museum, University of 

Copenhagen, Denmark

Other Abbreviations

AC aciniform spigot

AL alveolus

ALE anterior lateral eye

ALS anterior lateral spinneret

AME anterior median eye

BH basal hematodocha

CB cymbium

CY cylindrical spigot

E embolus

FL frame line

MAP major ampullate spigot

mAP minor ampullate spigot

MH median hematodocha

ML midline

MS modified spigot

NSS nonsticky spiral

OC o’clock

PC paracribellar spigot

PLS posterior lateral spinneret

PME posterior median eye

PMS posterior median spinneret

Rl lower radius

Rm middle radius

Ru upper radius

SEM scanning electron microscope

ST subtegulum

T tegulum

Acknowledgments

We thank Karolyn Darrow, Gustavo Hormiga, and 

Elaine Hodges for illustrations; Brian Kahn, Jeremy 

Miller, Dana DeRoche, and Simona Kralj-Fišer for SEM 

and other help; Magda Naparus for distribution maps; 

and Val Davies, Charles Griswold, Mark Harvey, Norman 

Larsen, and Robert Raven for correspondence. Mark Harvey and an anonymous reviewer made helpful suggestions 

to improve the manuscript.

This research was supported in part by the Slovenian 

Research Agency (grants Z1-7082-0618 and J1-2063 to MK 

and USA–Slovenia bilateral grants to MK and JC) and the 

U.S. National Science Foundation (EAR-0228699 to JC).

We thank the following curators, collection managers, 

and associated staff for their help in providing specimens 

and loans: Norman I. Platnick and Lou Sorkin (AMNH); 

Mike Gray and Graham Milledge (AMS); Janet Beccaloni 

(BMNH); Charles Griswold and Darrel Ubick (CAS); Rosemary Gillespie (EMUC); Maria Tavano (MCSNG); Gonzalo Giribet and Laura Leibensperger (MCZ); Christine 

Rollard (MNHN); Guy Redman (NMP); Robert Raven 

(QM); Rudy Jocqué (RMCA); Margie Cochrane, Dawn 

Larsen, and Norman Larsen (SAM); Ansie DippenaarSchoeman (SANC); Torbjörn Kronestedt (SMNH); Martin 

Krüger (TM); Dana M. DeRoche (USNM); Mark Harvey and Julianne Waldock (WAM); and Nikolaj Scharff 

(ZMUC).

Taxonomy

All measurements, including those on figures, are reported in millimeters, made using a microscope micrometer eyepiece. Prosoma and opisthosoma length and height 
number 636 • 5

were measured in lateral view, and their widths were measured in dorsal view, all at widest points (prosoma height 

measured at head region). Eye widths are maximum diameters of the eye lens; eye separations were measured 

between lenses. See “MORPHOLOGY” for detailed descriptions of family apomorphic and diagnostic features. 

All other taxonomic methods are standard in spider 

systematics.

Illustrations and Imaging

Older illustrations were rendered on Vellum® film 

or fine-grained coquille board and further rendered with 

black, soft Prismacolour® pencils. These artworks were 

scanned and further manipulated with Photoshop CS3. 

The newer illustrations were made with a Visionary Digital imaging system, equipped with a Canon 40d digital 

camera body and an Infinity K2 microscope using Olympus metallurgical objectives. Image “stacks” were combined with Helicon Focus 4.0 and further manipulated in 

Photoshop CS3 extended. Scanning electron microscope 

images were captured using an Amray 1810 SEM at the 

Smithsonian’s National Museum of Natural History Scanning Electron Microscope Facility.

Phylogenetic Methods

For deinopid exemplars, we used all Menneus and 

three Deinopis species, and for outgroups, we used Uloborus diversus and Dictyna arundinacea. Specimens examined for phylogeny were as follows.

Outgroups

Dictyna arundinacea: ♀ ♂ - USNM, N Sakhalin Okha 

District Pil’tun Bay, A. M. Basarukin, 6–7.1991.

Uloborus diversus: ♀ - USNM, Fresno Co. Chase Orchard Navel. 9.iv.74.

Ingroups

Deinopis spinosa: ♀ ♂ - USNM, FL, Alachua Co., Devil’s Millhopper St. Pk., mixed hardwoods. J. Coddington, 2.viii.1985.

D. lamia: ♀ - Coll. Alayon, El Veral. Guanahacabibes, 

ix.1981; M-La Chispa, Ballesteros, T. De Collantes. ix.1981, Col J. Santos.

D. cylindrica: ♀ - Natal Mus, South Africa, KwaZuluNatal, Lake Sibayi, Rhodes Univ. Exped., vi.1966; 

M-PAN, Zambesi (no further data).

Menneus camelus: ♀, South Africa, Johannesburg, 

dried branches in garden, M Filmer, 20.iii.1988; 

♂ - Natal Museum, South Africa, KwaZulu-Natal, 

Amanzimtoti, Inanda Nature Res, 3030BB?, 

P Reavell, 20.iv.1981.

M. capensis: ♀ - SAM, South Africa, Cape Prov., 

Cape Town, Newlands Forest, Paradijs’ Stream, 

N Larsen, 9.v.1991; ♀ ♂ - USNM, South Africa, 

Western Cape, Cape Town, Table Mtn. NP, Newlands Forest. 33°58.934′S 18°26.686′E. Alt. 159 m, 

Leg. Kuntner & al., 4.v.2006.

M. darwini: ♂ - All material examined (see “TAXONOMY”).

M. dromedarius: ♂ - NMP (NM 5894): South Africa, 

KwaZulu-Natal, Ngoyi Forest, Lawrence & al., 

ii.1953. ♀ - CAS (CASENT9012406): Madagascar, 

Toliara, Réserve Privé Berenty, Forêt de Bealoka, 

Mandraré River, 14.6 km 329° NNW Amboasary, 

ER30 beating low vegetation, 35 m, 24°57′25′′S 

46°16′17′′E, B. L. Fisher et al. BLF5318, 3-8.ii.2002.

M. samperi: ♀ ♂ - All material examined (see 

“TAXONOMY”). 

M. tetragnathoides: ♂ - lectotype - MNHN, “Congo” 

(Angola, Cabinda, Chinchoxo); RMCA, Malawi, 

Chintheche, R Jocqué, 2.v.1978.

M. aussie: ♀ ♂ - QM, Australia, QLD, Rundle Range, 

24.iii.1975.

M. bituberculatus: ♀ - QM, Australia, QLD, Upper 

Brookfield, pittraps, rainforest, Raven & Green, 

6.x.1981.

M. nemesio: ♀ - AM, Australia, NSW, Wamberal, MR 

Gray, 7.xii.1983; ♂ - CAS, Australia, NSW, 5 mi 

N Hawkesbury r. br., 150 m, Ross & Cavagnaro, 

3.xii.1962.

M. neocaledonicus: ♀ ♂ - QM, New Caledonia, 

Aoupinie, 20 km NE Poya, 650 m, Monteith & 

Cook, 18.v.1984.

M. quasimodo: ♀ holotype (see “TAXONOMY”).

M. superciliosus: ♀ ♂ - AM, Australia, QLD, Edmonton, NC Coleman, 29.ix.1969.

M. trinodosus: ♀ - AM, Australia, QLD, Eurimbula, 

SE Gladstone, rainforest site 4, 24°11′S, 151°50′E, 

C Horseman, iii.1975; ♂ - QM, Australia, QLD, 

Kroombit Tops (Three Moon Ck.), 45 km SSW 

Calliope rainforest, Davies & Gallon, 9.xii.1983.

M. wa: ♀ ♂ - All material examined (see “TAXONOMY”).

Menneus species, as noted in the Introduction, are 

rare spiders. Four Menneus species are only known from 

a single sex: M. tetragnathoides and M. darwini only from 

males and M. bituberculatus and M. quasimodo only from 
6 • smithsonian contributions to zoology

females. Because so few adults were available, we avoided 

as much as possible destructive sampling of specimens, 

such as dissections or SEM preparations. The use of light 

microscopy alone meant that phylogenetically informative features were necessarily macroscopic and scarce. We 

found only 56 potentially informative characters among 

the 19 taxa (see Phylogeny). We used Mesquite (Maddison 

and Maddison, 2009) to maintain the matrix and NONA 

2.0 (Goloboff, 1993) in combination with Winclada 

1.00.08 (Nixon, 2002) for tree searches using unweighted 

parsimony.

MORPHOLOGY

Millot (1933) mentioned several deinopid apomorphies. The cephalothoracic intestinal diverticulae of Uloboridae are solely supra-oesophageal and simple. Four pairs 

of diverticulae lead to the walking legs and one anterior 

pair enter the mouth area. Both Deinopis baubo and Menneus camelus have one system homologous to the uloborid 

condition and an extra system of sub-oesophageal diverticula, which Millot regarded as unique in spiders. In addition, deinopids have a sixth pair of diverticulae beneath 

the homologs of the uloborid anterior diverticulae: these 

serve the bases of the pedipalps and the labrum. Deinopis, 

uniquely, has the right anterior diverticula (dorsal view) 

enlarged and expanded to form the “frontal diverticle” in 

a semilunate shape immediately behind the lateral eyes.

Deinopidae have several sorts of body setae: blunt, 

plumose, and feathery setae (Figure 5a–d). Chelicerae always bear anterior and posterior teeth, but the number 

varies within and between species (Figure 5c).

The fourth metatarsus, as in all cribellate spiders, 

bears a calamistrum (Figure 5e). The fourth tarsus has a 

ventral comb of distinctive setae, and the retrolateral claw 

is reduced (Figure 5f). Male legs are relatively much longer than female legs (Figure 1a–c), and male legs I and II 

are additionally modified in various ways (see characters 

31–37, below, and Figure 3c–e).

Deinopids present a complete complement of spinnerets and spigots (Figure 7a). The cribellum is entire and 

broad, with the annulate cribellate spigots apparently 

clustered more densely on the anterior cribellum surface 

(Figure 7b,c). The ALS have a fairly extensive piriform 

field, and the major ampullate field, unusually among spiders, contains multiple functional major ampullate spigots 

(Figure 7d). The PMS bear a single anterior minor ampullate spigot, multiple apical aciniform spigots, an extensive field of anterior paracribellar spigots (with annulate 

shafts), and an extensive posterior field of cylindrical spigots (Figure 7e). The PLS have a distal “modified” spigot 

(putative homolog of the uloborid pseudoflagelliform and 

araneoid flagelliform spigot), multiple aciniform spigots, 

and an extensive basal brush of cylindrical spigots (Figure 7f). Male epiandrous spigots are grouped (Figure 6a), 

smooth, and set in deep pockets (Figure 6b).

The ventral surface of the deinopid anal tubercle has a 

modified field of setae, symmetric and oriented toward the 

midline (Figure 6c).

Deinopid palpal bulbs are built on a simplified plan 

(Figure 4a,b), which probably reflects exacting functional 

requirements. The embolus is always a thin, relatively flat, 

spiral sclerite, and the number of coils correlates, to some 

extent, with the number of coils of the female copulatory 

ducts. (For the 12 deinopid taxa of Table 1 known from 

both sexes, the correlation coefficient, r2, is 0.51 and p = 

0.08.) The embolus has a complex cross section (Figure 

6f) with an inner ridge and an adjacent groove. The ridges 

and grooves of a given loop probably engage adjacent 

loops. This reinforcing mechanism may aid the male to 

insert the tip of the embolus all the way into the spermathecae of the female during copulation. The tip of the 

embolus may have subdistal structures whose function is 

enigmatic (Figure 6f).

The tegulum has only a single apophysis other than the 

embolus, which is probably the homologue of the uloborid 

and araneoid median apophysis (Figure 6d–f), not the conductor (contra Kuntner et al., 2008: fig. 20). The deinopid 

median apophysis is compact, and the apical part is always 

more or less hook-shaped (Figure 15g), with the concavity 

facing the median or apical margin of the palp. The basal 

part often has a lobe (Figures 6e, 10j), more or less visible in ventral view. The apical hook probably engages the 

epigastric rim adjacent to the median plate, or septum, of 

the female epigynum (Figure 22e–g) during copulation, thus 

locking the bulb to the epigynum. The median apophysis is 

flexibly attached to the tegulum, via a coiled, tough, tendonlike membrane. The tegulum is also highly modified. The 

ejaculatory duct is a simple spiral. The walls of the tegulum 

are smooth, and it is shaped like a squat sphere. There are 

no lateral projections. It is firmly attached to the subtegulum. The subtegulum is small and contains the fundus of 

the reservoir. It is attached to the cymbium by the extremely 

coiled, tough, basal hematodocha (Figure 4b).

We hypothesize that these features of the palp cooperate in a special copulatory mechanism. In order to introduce the embolus tip into the female spermathecae, the 

tegulum apparently rotates a number of times as the basal 

hematodocha expands, thus screwing the embolus into the 
number 636 • 7

copulatory ducts. All spider palps rotate as they expand, 

but the morphology of the deinopid palp seems specialized 

to rotate to an exaggerated extent. If the median apophysis is fixed to the epigynum during copulation, it cannot 

rotate. Likewise, the cymbium cannot rotate. However, 

via the median apophysis membrane and similarly coiled 

basal hematodocha, the tegulum can “unwind” through 

as many revolutions as there are coils, thus imparting a 

spiral motion to the embolus.

The female genitalia (Figure 23e–g) are entelegyne, 

unlike some genera of uloborids. Externally, the epigynum 

is simple, consisting of a median plate bordered laterally 

by the thickened rims of the epigastric furrow. Between 

the median plate and the thickened rims lie paired copulatory slits (Figure 27e,f). The thickened margins of the epigastric furrow adjacent to the median plate are excavated 

to a greater or lesser extent on their dorsal surface, thus 

forming a small hollow that probably engages the male 

median apophysis during copulation.

Internally, the copulatory ducts spiral around the 

fertilization ducts (e.g., Figure 27g). In posterior and 

dorsal view, the left copulatory duct, for example, spirals 

counterclockwise from the epigynum toward the spermatheca. The entrance to the copulatory duct is at the basal, 

wider portion of the median plate of the epigynum, more 

or less at the bottom of the copulatory slits. The fertilization ducts are heavily sclerotized and long (probably additional synapomorphies for Deinopidae). These thickened 

ducts might serve as a stiffening support or anchor for the 

coil of copulatory ducts. The lumen of the duct is quite 

straight. Sometimes the wall of the fertilization ducts are 

reinforced with auger-like spiral thickenings, to which the 

inner margins of the copulatory duct may be fused. The 

initial curve of the fertilization duct passes along the margin of the lateral epigynal margins and generally is quite 

visible in posterior view as a darkly sclerotized area adjacent to the rim of the median epigynal plate (Figure 23g).

Although the distal sclerotized portions of the female 

genitalia are often massive, actual limits of spermathecae are often difficult to discern. Glandular pore fields in 

Deinopis seem to be limited to the apical portion of the 

spermathecae.

TABLE 1. Phylogenetic character matrix. See “Character Descriptions” for explanation of the 56 characters and their states; see 

“Phylogenetic Methods” for list of exemplar taxa and sources. A dash (-) indicates inapplicable data; a question mark (?) indicates 

unknown data.

Character number

Terminal taxa 10 20 30 40 50 56

Dictyna arundinacea 00-0000000 00-0000--0 002100--00 00000000-0 100----30- 00-000

Uloborus diversus 00-0000000 00-0001000 000-00--00 0000000100 000-0--20- 00-010

D. cylindrica 1101000100 10-0011001 000-011001 11111110-1 0014010111 100???

D. spinosa 1101001111 1101011001 000-011010 0000000101 0010000011 110121

D. lamia 1101001110 10-0011001 000-011010 000000?101 0010000011 110???

Menneus bituberculatus 0100011211 00-1011100 000-01002? ?????????? ?????????? ?????? 

M. trinodosus 0100011211 0101011100 100-010210 0000000101 1111011111 100?21

M. capensis 0100011211 00-0011000 0011110111 0000000101 1110011112 100121

M. superciliosus 0100011201 00-0011000 0110110121 0000001101 1112100313 000?21

M. neocaledonicus 0100010111 00-0000--0 100-010200 11100110-1 1114100010 000???

M. quasimodo 0110010101 00-0011000 100-01003? ?????????? ?????????? ?????? 

M. wa 0110011211 00-0011000 0111110121 111111?101 1113000013 000???

M. aussie 0110011201 00-0011000 1011110151 1111111101 1112100013 000???

M. nemesio 0110011111 00-0011000 1111110131 11111110-1 1112100213 100???

M. camelus 0110111211 0111111110 100-010120 1111111111 1114100012 101121

M. tetragnathoides ?????????? ?????????? ?????????0 111111?111 1111100113 001???

M. dromedarius 0110111211 0101011110 100-010140 1111111111 1113100113 000???

M. darwini ?????????? ?????????? ?????????0 1111111111 1112100113 001???

M. samperi 0110111211 0111111110 110-010130 1111111111 1113100113 1001??
8 • smithsonian contributions to zoology

PHYLOGENY

Table 1 gives the matrix for 19 terminals and 56 phylogenetic characters and character states, which are described below.

Character Descriptions

Female Somatic Morphology

1. Female PME setal fringe: (0) absent (Figure 18a); 

(1) present (Figure 3a). The posterior median eyes 

of female Deinopis have an encircling fringe of 

short, broad, modified setae.

2. Female ALE tubercle: (0) absent (Opell, 1979: fig. 

23); (1) present (Figure 3a). The ALE of all known 

deinopids occur on prominent tubercles such that 

the eyes point ventrolaterally. The animals hunt 

ventral side up and this feature evidently enables 

the lateral eyes to see above and to the sides.

3. Female ALE tubercle: (0) small (Figure 3a); (1) 

large (Figure 25d). In some Menneus species the 

ALE tubercle is notably longer.

4. Female AME/PME ratio: (0) ≥0.50 (Figure 25d); 

(1) ≤0.30 (Figure 3a). All Deinopis species have 

greatly enlarged posterior median eyes.

5. Female clypeal setae: (0) absent (Figure 25d); (1)

present (Figure 18a). Some Menneus species have 

noticeably thicker and larger setae medially on 

the margin of the female clypeus.

6. Female cheliceral setal conformation. (0) multiple, irregular (Figure 10a); (1) distinct medial and 

frontal macrosetae (Figure 28b) Menneus species 

have two pairs of distinct cheliceral macrosetae.

7. Female endite trichobothria: (0) absent; (1) present (Figure 5b). The endite bears, usually, two 

basal shorter and two distal longer trichobothria 

that point in notably different directions than the 

surrounding setal fields.

8. Female sternum: (0) broad; (1) setose knob (Figure 

5a); (2) pointed (Figure 31k). The sternum posterior apex may be relatively broadly truncated, 

a blunter knob bearing setae, or sharply pointed.

9. Female sternum pattern: (0) uniform or mottled; 

(1) light median band (Figure 20d). The sternal 

color pattern may be uniform to mottled or with 

a median light band.

10. Female posterior carapace white mark: (0) absent; 

(1) present (Figure 32a). The posterior carapace 

margin may have a white V-shaped guanine mark.

11. Female dorsal femur I: (0) normal; (1) bulging 

(Figure 3b). The basal, dorsal femur is sometimes 

expanded into a bulge or knob.

12. Female femur III ventral hair brush: (0) absent; (1) 

present. The retrolateral surface of femora III and 

IV sometimes have a distinct brush of long setae.

13. Female femur III hair brush length: (0) short; (1) 

long. The third femoral hair brush may be short 

or long.

14. Female femur IV hair brush: (0) absent; (1) present.

15. Female coxae III–IV pleural bar: (0) absent; (1) 

present. Some deinopids have sclerotized bars 

between the third and fourth coxae, which join 

carapace to sternum.

16. Female tarsus IV retrolateral claw teeth: (0) normal; (1) reduced (Figure 5f, arrow). The fourth 

tarsal retrolateral claw may be smaller and may 

bear fewer, smaller teeth than the prolateral claw.

17. Female abdominal form: (0) smooth (Figure 28c); 

(1) tuberculate (Figure 10d). Deinopoid females 

generally have paired dorsal abdominal tubercles.

18. Female abdominal tubercle position: (0) mediananterior (Figure 25b,c); (1) posterior (Figure 

31c,d). Abdominal tubercles may be either median or posterior.

19. Female abdominal tubercle symmetry: (0) symmetric (Figure 20c); (1) asymmetric (Figures 1d, 

18c). Almost all deinopids have abdominal tubercles, but in some Menneus species the tubercles 

are greatly enlarged and single, either left or right, 

and both states may occur in the same species.

20. Female abdomen proportion: (0) width/length 

>0.4; (1) width/length <0.4 (Figure 11d). The 

Deinopis abdomen is relatively longer and thinner than other taxa in this study.

21. Female ventral abdominal dark band: (0) mottled/solid (Figure 30d); (1) paired (Figure 27d). 

The abdomen bears a median ventral dark area, 

which in some species is mottled or nearly solid 

and in others it is paired.

22. Female abdominal ventral guanine pattern: (0) 

mottled (Figure 11c); (1) paired conspicuous 

guanine spots (Figure 20d). Some species have 

paired, relatively anterior, ventral abdominal guanine spots or patches.

23. Female dorsal posterior abdominal pattern: (0) 

mottled (Figure 11d); (1) paired longitudinal narrow bands (Figure 27c); (2) median dark band.

24. Female dorsal posterior abdominal pattern 

shape: (0) straight to undulating (Figure 30c); (1) 
number 636 • 9

marked, regular undulations, chevron-like (Figure 27c).

25. Female abdominal mid-dorsal color pattern: (0)

longitudinal mottled or light band (Figure 28c); 

(1) transverse light bar (Figure 27c).

Female Genital Morphology

26. Female epigynal median area: (0) chamber or thin 

septum (Opell, 1979: fig. 43); (1) broad, platelike septum. The median epigynal area of deinopids always bears a well-developed median septum 

that appears as a broad plate (Figure 10e).

27. Female epigynal median plate: (0) flat (Figure 

18f); (1) involute (Figure 10e). In ventral view the 

anterior lateral margins of the median septum of 

Deinopis is excavated so that the septum appears 

to have a pronounced neck.

28. Female epigynal septum lateral margins: (0) subrectangular (Figure 9b); (1) triangular (Figure 

25f); (2) subparallel (Figure 31e). The overall outline of the epigynal median septum may be wider 

than long (subrectangular), triangular, or longer 

than wide (subparallel).

29. Female number of copulatory duct turns: (0) 1; 

(1) 2; (2) 3; (3) 4; (4) 5; (5) 6. The number of 

copulatory duct turns in deinopids can be quantified in various ways, depending on where the duct 

turns are considered to start and end. The least 

ambiguous seems to be to count them from figures representing a particular orientation. Consequently, we quantify duct turns in dorsal view, as 

the number of apparent turns on the lateral margin of the copulatory/fertilization duct complex 

(e.g., Figure 28g, 1 turn; Figure 27g, 4 turns). The 

number of turns in M. quasimodo is ambiguous; 

we code it as state three.

Male Somatic Morphology

30. Male carapace shape: (0) pear-shaped, sides of 

thoracic region parallel (Figure 22h); (1) diamondshaped, sides of thoracic region narrowing posteriorly (Figure 25h). As in females, male carapace 

shape varies informatively, either the plesiomorphic pear shape or a more angular diamond shape.

31. Male tibia II distal end: (0) normal; (1) with

apophysis (Figure 3d). Tibia II of some males 

bears a prolateral strong apophysis at the metatarsal joint. This and the next two characters 

pertain to distal tibial modifications and are distributed identically for this taxon sample, but we 

code all three conditions because the morphology 

of each is distinct.

32. Male tibia–metatarsus condyle: (0) normal; (1) 

stout (Figure 3d). The ectal condyle of the male 

tibia–metatarsal joint is sometimes stronger and 

stouter than normal.

33. Male tibia II tip: (0) normal; (1) swollen (Figure 

3d). In addition to the previous modifications, 

the male tibia II distal tip may be significantly 

broader or swollen.

34. Male distal Mt II setae: (0) normal; (1) curved 

and/or stout (Figure 3e). The distal end of metatarsus II bears several setae, of which the dorsal 

and/or prolateral may be distinctly longer, curved, 

or stouter in some species.

35. Male Mt–Ta II joint: (0) normal; (1) ball and 

socket (Figure 3e). The metatarsus–tarsal joint 

in some species is modified, with the metatarsus 

bearing a distal, dorsal protruding knob that fits 

into a corresponding socket on the tarsus.

36. Male Ta II dorsal setae: (0) normal; (1) spatulate 

(Figure 3e). The dorsal surface of male tarsus II may 

bear a longitudinal field of short, spatulate setae.

37. Male Ta I form: (0) entire; (1) divided (Figure 3c).

Like the tibia, the male tarsus I may be divided.

38. Male abdominal form: (0) strictly cylindrical (Figure 10i); (1) more oval and tuberculate (Figure 

20i). As in the females, some species have prominent dorsal abdominal tubercles. Tubercles in 

males can vary greatly in size.

39. Male abdominal tubercle symmetry: (0) paired 

(Figure 20i); (1) unpaired (Figure 24d). As in females, male abdominal tubercles may be paired 

or single and either left or right.

Male Genital Morphology

40. Male cymbial form: (0) oval, elongate (Opell, 

1979: fig. 47); (1) round, hemispherical (Figure 

4e). In ventral view, deinopid cymbia are remarkably circular in outline, and in lateral view nearly 

hemispherical.

41. Male apical cymbial margin: (0) smoothly 

rounded (Figure 4b); (1) pointed (Figure 4e). In 

some males the distal margin of the cymbial alveolus is rounded, and in others it is pointed.

42. Male apical cymbial setae: (0) absent (Figure 

4a,b); (1) present (Figure 4e). Some males have 
1 0 • smithsonian contributions to zoology

noticeable tufts of curved, often white setae on 

the apical margin of the alveolus.

43. Male tegulum form: (0) elongate, oval (Opell, 

1979: fig. 46); (1) round, radially symmetric (Figure 14b). In ventral view, all deinopid tegula are 

remarkably circular in outline.

44. Median hematodocha position: (0) 12–(4–6); 

(1) 2–(4–5); (2) 4–(7–8); (3) 6(8)–9(11); (4) (10–

11)–(1–2). The span of the median hematodocha 

insertion on the tegulum in ventral view varies 

both in position and extent and is often diagnostic for species. It can be quantified as occupying 

various sectors of an imaginary “clock” (Figure 

14b “10–2,” e “11–4,” h “6–9”). In an attempt 

to constrain the proliferation of singleton character states, we occasionally subsume variation between species, thus (10–11)–(1–2) means a span 

of 10–11 OC to 1–2 OC.

45. Median apophysis base: (0) smoothly rounded 

(Figure 14e); (1) with knob (Figure 15b,e,h). In 

ventral view, the base of the median apophysis may be rounded or produced into a blunt or 

somewhat acutely pointed knob.

46. Median apophysis distal lobe: (0) relatively 

broad, subrectangular (Figure 22l); (1) relatively 

narrow, elongate (Figure 10j). In ventral view, the 

distal tip of the median apophysis may be fairly 

narrow, such that the basal portion of the median 

apophysis is easily visible or broad such that no 

part of the basal portion shows.

47. Median apophysis distal lobe curvature: (0) 

abruptly angled (Figures 4e, 15g); (1) gently 

curved (Figure 17f). In retrolateral or prolateral 

view, the median apophysis may either bend 

sharply, at almost a right angle, or its curve may 

be gradual and gentle.

48. Embolic coil width: tegulum width: (0) << 1; (1) 

< 1; (2) subequal; (3) > 1. In ventral view, the 

diameter of the outermost embolic coil may be 

much less (Figure 16h), slightly less (Figure 14h), 

subequal (Figure 16e), or greater than (Figure 

17b) the diameter of the tegulum.

49. Embolus form: (0) curved (Opell, 1979: fig. 47); 

(1) spiraled (Figure 4c). Outgroups such as Uloboridae may have distinctly curved emboli, but 

all deinopid emboli are strongly and often extremely spiraled.

50. Embolic coil number: (0) 2; (1) 2.5; (2) 3; (3) 

3.5 or more. The same issue that arises when 

quantifying female copulatory duct turns or 

twists (character 29) also pertains to quantification of male embolic turns. In Menneus the tip 

of the embolus of a normal palp terminates close 

to the cymbium apex. Coils are counted to the 

nearest half rotation, from illustrations of ventral 

views, beginning from the tip of the embolus until 

it disappears beneath the median apophysis (e.g., 

Figure 14b [3.5 turns], e [2.5 turns]).

51. Embolus tip form: (0) simple, attenuate (Figure 

24e,f); (1) distally widened (Figure 10j,k). In 

ventral view, the embolus tip may be simply attenuate or needle-like, or it can be expanded or 

widened subapically.

52. Embolus tip symmetry: (0) entire (Figure 20k, l); 

(1) excavated (Figure 11m). The embolus tip may 

be symmetric (whether widened or not) or asymmetric, with one side of the embolus prolonged 

into a narrow, delicate tip.

53. Embolus tip: (0) bent smoothly away from embolus coil (Figure 14f); (1) relatively straight 

(Figure 14i). In retrolateral view, the tip of the 

embolus may lie in the plane of the embolic coils 

or may abruptly bend away from the tegulum toward the median apophysis.

Behavior

54. Female eggsac form: (0) oval to flattened (Opell, 

1979: fig. 254); (1) spherical (Figure 2e). All 

known deinopid eggsacs are markedly spherical, whereas most spider eggsacs, and in this case 

those of the outgroups, are more or less flattened 

and oval. Deinopid egg sac walls are also relatively dense and hard.

55. Web architecture: (0) sheet; (1) orb with many radii 

(Opell, 1979: pl. 8); (2) orb with few radii (Figure 

2d). All known deinopid webs are bilaterally symmetric, modified orbs (Coddington, 1986a) with 

three radii on each side. Although the web architecture of Menneus is known only for a few species, all show the expected deinopid architecture.

56. Sticky silk area size: (0) much larger than spider 

(Opell, 1979: pl. 8); (1) subequal to spider size 

(Figure 2c). Most web-spinning spiders spin webs 

much larger than their body size. Deinopids make 

webs whose sticky silk area is roughly 1–2 cm at 

rest and thus comparable to the body size and leg 

span of the spider.
number 636 • 1 1

RESULTS

The matrix of Table 1 yields a single tree of length 131 

under unweighted parsimony, with consistency index 0.54 

and retention index 0.69 (Figure 8a). Although Figure 8a 

is the single most parsimonious explanation for the data of 

Table 1 and a testable hypothesis, the matrix is so sparse 

that applying measures of nodal support would be pointless (most nodes within Menneus would show low levels 

of support, at best). All 131 steps are mapped on Figure 

8a. Ambiguous mappings were usually resolved with fast 

optimization, thus allocating homoplasy to secondary loss, 

but for several features, slow optimization seemed more 

appropriate. Deinopidae, Deinopis, and Menneus are 

monophyletic, and their synapomorphies are discussed in 

“TAXONOMY.” Of note, secondary sexual characters affecting male first and second legs evolve in a block at the 

node subtending Menneus wa. Large posterior median eyes 

are a Deinopis synapomorphy. Deinopid web architecture 

(Figures 1a,d, 2b–d) is a family synapomorphy. Given that 

Deinopis is circumtropical and Menneus is limited (mostly) 

to the Old World tropics, the family must predate the 

breakup of Gondwana. The biogeography as mapped on 

the cladogram (Figure 8a) suggests that the origin of Menneus may be Australasian. The cladistic position and African distribution of M. capensis among Australasian species 

is therefore anomalous. The earliest known fossil, Paleomicromenneus lebanensis (Penney, 2003), is Cretaceous (M. 

pietrzeniukae and other juveniles are known from Baltic 

amber; Wunderlich 2004), but by that time, Africa and 

Australasia had already rifted apart. Given the uncertainty 

as to the true ages of lineages, and, most importantly, the 

paucity of known specimens and severe undersampling bias, 

the biogeographic pattern cannot be ascribed at this point 

either to vicariance or dispersal. A distal clade of species is 

unique to Africa (Figure 8a) and defined by an unusual synapomorphy: the ability to develop asymmetric (i.e., left or 

right) abdominal tubercles. This feature, in addition to the 

family-level apomorphies of femoral setal fringes, mottled 

coloration, and daytime resting postures, no doubt form a 

complex of adaptations promoting crypsis.

TAXONOMY

Deinopidae C. L. Koch, 1850

Diagnosis. Entelegyne cribellate spiders; females with median epigynal plate and copulatory ducts 

coiling around long, heavily sclerotized fertilization ducts 

(Figure 23g), males with unique, flat, round tegulum held 

in a round, hemispherical cymbium, embolus long, coiled 

around a single tegular apophysis, hypothesized to be the 

median apophysis homologue (Figure 4a–e). Both sexes 

with anterior lateral eyes on distinct, ventrally oriented 

tubercles (Figure 10a,b,f,g). Tapetum absent from all secondary eyes. Fourth tarsal retrolateral claw teeth reduced 

(Figure 5f). Web architecture and attack behavior unique, 

a highly reduced and stylized orb web, about the same size 

as the body length of the spider, held in the anterior legs, 

and actively used to envelop aerial or pedestrian prey (Figures 1a,d, 2b–d). Egg sac probably unique: a hard, brown 

sphere attached to leaves or buried in leaf litter (Figure 2e).

Composition. Three lineages, Menneus, 

Deinopis, and Paleomicromenneus, treated as genera. 

Paleomicromenneus lebanensis Penney 2003 is known 

only from Cretaceous Lebanese amber (Penney, 2003). 

The male specimen is small (2.86 mm body length) but 

otherwise fits comfortably into the family, resembling M. 

capensis in genitalic form.

Species Diagnostic Characters. While 

external epigynum shape varies to some extent, the internal morphology (especially size, proportion, and number 

of coils of copulatory duct) is diagnostic. In males, the relative origin of the median hematodocha, the orientation 

of the median apophysis, and the position of the embolus 

tip, and its morphology, are diagnostic (Figures 14–17). In 

local faunas, abdominal form and coloration can be valuable identification clues (e.g., Figure 1b).

Distribution. Tropics and subtropics worldwide, extending to temperate SW Australia, from sea level 

to as much as 2300 m.

Deinopis MacLeay, 1839

Deinopis MacLeay, 1839:9.—Platnick, 2010. Type species D. lamia MacLeay, 1939, by monotypy.

Dinopis: Bonnet, 1956.—Lehtinen, 1967:230, 306.

Notes. Original genus spelling is used (MacLeay, 1839; Platnick, 2010), not the unjustified emendation Dinopis (as in Bonnet, 1956). The name is feminine 

(Bonnet, 1956).

Diagnosis. Synapomorphies in this analysis 

that define Deinopis are: the PME are less than a diameter apart and extremely enlarged (at least 3× the AME 

diameter) with a fringe of setae (characters 1 and 4, Figure 

3a). Female first femora are distally enlarged (character 
1 2 • smithsonian contributions to zoology

11, Figure 3b). Deinopis abdomens, although not unique 

among spiders in their proportions, are usually much longer than wide (character 20, Figure 9a).

Description. Large spiders, female total 

length from 15 to 28 mm, males from 12 to 23 mm. Body 

and legs usually gray to brown. Cephalothorax elongate, 

with a narrow cephalic region (Figure 9a,e), and a long, 

narrow sternum, prolonged posteriorly (Figure 5a). Eyes 

as diagnosed, with ALE on tubercles (Figures 10a,b,f,g; 

11a,b,i,j). Chelicerae with conspicuous pro- and retrolateral teeth, sometimes also with small denticles close to 

posterior rim (Figure 5c). Very long legs with leg formula 

1, 2, 4, 3. Abdomen cylindrical, usually many times longer than wide, with or without prominent paired tubercles (Figures 9a,e, 10c,d,h,i, 11c,d,k,l). Female epigyna 

inconspicuous, median septum triangular to rectangular, 

with small round spermathecae and long copulatory ducts 

coiled around straight sclerotized fertilization ducts (Figures 9b–d, 11e–h). Male palpal tegulum partly covered 

with conspicuous embolus coiled around median apophysis. Embolus makes 1.5 to many turns; embolic tip tapered, flattened, or sinuate (Figures 9f, 10j, 11m).

Species. The genus currently contains 46 species (Platnick, 2010). However, our generic revision in 

preparation will recognize approximately 30 valid species.

Distribution. Tropics, subtropics worldwide.

Paleomicromenneus Penney, 2003

Palaeomicromenneus Penney 2003: 571. Type species Palaeomicromenneus 

lebanensis Penney 2003, by monotypy.

Notes. Penney (2003) described this species 

based on a single male from Lebanese amber. On the 

basis of the size (2.9 mm) of the type species, the genus is 

probably valid. Features used to diagnose modern species 

(embolus position, median apophysis form, median hematodocha extent) were not visible in the fossil specimen.

Menneus Simon, 1876

Menneus Simon, 1876:218.—Lehtinen, 1967:217, 306.—Platnick, 2010. 

Type species Menneus tetragnathoides Simon, 1876, by monotypy.

Avella Pickard-Cambridge, 1877:574 [see Cambridge, O. P., in References].—

Lehtinen, 1967:217, 306.—Platnick, 2010. Type species Avella despiciens Pickard-Cambridge, 1877, from Rockhampton, Queensland, 

Australia, by monotypy. NEW SYNONYMY.

Avellopsis Purcell, 1904: 128.—Lehtinen, 1967:217, 306.—Platnick, 2010. 

Type species Avellopsis capensis Purcell, 1904, from Kalk Bay Mountain, South Africa, by monotypy. NEW SYNONYMY.

Note. The name is masculine (Bonnet, 1957).

Diagnosis. Menneus are easily recognized as 

deinopid spiders in which the PME are always at least a 

diameter apart and of relatively normal size (AME/PME 

≥ 0.40, Figure 18a,b,g,h). Synapomorphies for the genus 

at this point are subtle. Menneus have distinct frontal 

and medial cheliceral setae (character 6, Figures 20b, 

25d), the carapace usually has a distinct posterior guanine patch (character 10, Figure 32a), and the cymbium 

bears a distal group of long, curving setae (character 42, 

Figure 4e).

Description. Medium sized spiders, female total length from 7.2 to 19.5 mm, males from 5.0 

to 13.6 mm. Body and legs usually gray to brown, with 

patches of white and black. Cephalothorax elongate, 

pear or diamond shaped (Figures 18b,h, 20a,g). Eyes as 

diagnosed, with ALE on pronounced elongate tubercle, 

AME usually on smaller tubercle (Figure 19a,b). Sternum longer than wide, relatively acute posteriorly (Figure 20d). Long legs with leg formula 1, 2, 4, 3 (Figures 

1a–d, 2a–c,e). Abdomen cylindrical, longer than wide, 

usually with a single (all African species except capensis) 

(Figure 18c,d,i,j) or paired dorsal tubercles (capensis and 

all Australasian species except neocaledonicus) (Figure 

20c,i). Female epigyna relatively small, triangular, with 

inconspicuous posterior copulatory openings (Figure 

18e,f), spermathecae round, unlobed, long thick copulatory ducts coiled around long straight fertilization ducts 

(Figures 22g, 23g). Male palpal cymbium round, apically 

pointed, ventral palp view exposes median hematodocha 

at edge of flat tegulum; tegulum partly covered with conspicuous embolus coiled around median apophysis. Embolus makes 2–4 turns; embolic tip tapered, flattened, or 

sinuate (Figures 14–17).

Species. We currently recognize 14 Menneus

species, six from Africa, and eight from Australasia. Several old Australasian names were based on juvenile specimens, and because species are often sympatric and can 

only be identified confidently from adults, we regard these 

names as nomina dubia (see below).

Distribution. Central and southern Africa, 

Madagascar, Australia, New Guinea and New Caledonia 

(Figures 12, 13).
number 636 • 1 3

KEY TO MENNEUS SPECIEs

1. Africa or Madagascar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Australasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

2. Males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

3. Embolic coils in ventral view less than tegular width [Figure 14c,f] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Embolic coils subequal to or greater than tegular width [Figures 14i, 15i] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4. Embolic coils much less than tegular width, median apophysis distal lobe broad and flat [Figure 14a–c] . . . . M. camelus

Embolic coils slightly less than tegular width, median apophysis distal lobe narrow, extended [Figure 14d–f] . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. capensis

5. Embolus tip bent away from tegulum in prolateral view [Figure 15c,f], median apophysis distal–basal lobe axis points to 

2 OC [Figure 15b,e] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Embolus tip relatively straight and parallel to tegulum in prolateral view [Figures 14i, 15i], median apophysis distal–basal 

lobe axis points to 1 OC [Figures 14h, 15h] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6. Median hematodocha in ventral view begins at roughly 6 OC, median apophysis in pro- or retrolateral view relatively flat 

[Figure 15a–c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. dromedarius

Median hematodocha in ventral view begins at roughly 8 OC, median apophysis in pro- or retrolateral view relatively 

rounded [Figure 15d–f] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. samperi

7. Median apophysis in pro- or retrolateral view rounded [Figure 14g–i], embolus tip at 10:30 OC . . . . . . . . . M. darwini

Median apophysis in pro or retrolateral view flat and angled [Figure 15g–i], embolus tip at 11:30 OC . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tetragnathoides

8. Copulatory duct turns three or fewer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Copulatory duct turns four or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

9. Copulatory ducts with one full turn (or two if counting lateral margins in dorsal view) [Figure 20f] . . . . . . M. capensis

Copulatory ducts with two full turns (or three if counting lateral margins in dorsal view) [Figure 18k] . . . . M. camelus

10. Copulatory ducts with three full turns (or four if counting lateral margins in dorsal view) [Figure 23g] . . . . M. samperi

Copulatory ducts with four full turns (or five if counting lateral margins in dorsal view) [Figure 22g] . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. dromedarius

11. Males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

12. Embolic coils less than tegular width [Figures 16i, 17f,i] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Embolic coils subequal to or greater than tegular width [e.g., Figures 16e, 17a,b] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

13. Embolic coils only about half the tegular width [Figure 16i] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. neocaledonicus

Embolic coils more than half the tegular width [Figure 17f,i] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Median apophysis distal lobe narrow, only slightly curved [Figure 17d–f] . . . . . . . . . . . . . . . . . . . . . . . . . M. trinodosus

Median apophysis distal lobe broad, abruptly curved [Figure 17g–i] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. wa

15. Embolus tip abruptly curved away from tegulum [Figure 17a,c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. superciliosus

Embolus tip gradually curved away from tegulum [Figure 16c,f] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16. Median apophysis in ventral view subrectangular [Figure 16b] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. aussie

Median apophysis in ventral view curved, lunate [Figure 16e] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nemesio

17. Abdominal tubercles absent [Figure 28c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. neocaledonicus

Abdominal tubercles present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Abdominal tubercles posterior [Figures 26c, 31c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Abdominal tubercles anterior or median [Figures 25b, 30c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19. Abdominal tubercles single, not bifid [Figure 31c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. trinodosus

Abdominal tubercles bifid [Figure 26c] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. bituberculatus

20. Abdomen extends caudally beyond the spinnerets [Figure 27c,d] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nemesio

No caudal abdominal extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

21. Copulatory ducts with only two full spirals (or three counting lateral margins in dorsal view) [Figures 30g, 32g] . . . . 22

Copulatory ducts with three or more full spirals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

22. Abdomen relatively narrow, lateral margins mottled [Figure 30c,d] . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. superciliosus

Abdomen wide, with conspicuous lateral brown spots [Figure 32c,d] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. wa

23. Copulatory ducts with three or four full spirals (or four or five counting lateral margins in dorsal view) [Figure 29g is 

ambiguous] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. quasimodo

Copulatory ducts with five full spirals (or six counting lateral margins in dorsal view) [Figure 25g] . . . . . . . . . M. aussie
1 4 • smithsonian contributions to zoology

African Species

Menneus camelus Pocock, 1902

Figures 2a–c, 2e, 4e, 12, 14a–c 18, 19

Menneus camelus Pocock, 1902:325 (♀).—Lehtinen, 1967:247, fig. 40 (see 

M. dromedarius for incorrect synonymy).—Penney, 2003:572, figs.1–3 

(♂).—Platnick, 2010.

Types. Holotype female from Durban, Natal, 

South Africa, in BMNH, examined.

Diagnosis. Female camelus differ from all 

Menneus except M. dromedarius and M. samperi by the 

usual presence of an unpaired abdominal tubercle (Figure 

18c,d,i,j, but see variation). They differ from M. dromedarius and M. samperi by the epigynum with only two full 

(or three, counted as visible lateral margins in dorsal view) 

wide copulatory duct coils (Figure 18k). The median hematodocha in ventral view (left palp) spans 11–2 OC, the 

embolus tip lies at about 11 OC, and the median apophysis–basal lobe axis lies at about 2 OC (Figure 14b). The embolus makes 3.25 coils and is relatively straight and parallel 

to the tegulum in prolateral view (Figure 14c), and, unlike 

all other African Menneus, the embolic coils are narrower 

than the tegular width in ventral view (Figure 14a–c).

Description. Female from Umkomaas River, 

KwaZulu-Natal. Total length 14 mm. Carapace 5.3 mm 

long; 2.3 mm wide. Abdomen 8.8 mm long; 5.0 mm wide. 

Cephalic width 0.4× carapace width. PME diameter 1 mm; 

PME separation 1.9 diameters. AME/PME 0.7; AME separation 2.5 AME diameters; closer to PME than to clypeal 

margin. ALE/PME 0.8; ALE tubercle large, eye’s lower edge 

2/3 or more diameters from clypeus margin (Figure 18a,g); 

ALE 1.5 diameters below clypeal margin. PLE/PME 0.8; 

PLE separation 7.2 PLE diameters; PLE 1.3 diameters behind PME. Clypeus 3.1 AME diameters. Chelicerae with 5 

to 7 promarginal teeth; 4 retromarginal teeth; frontal macrosetae in transverse line; laterals stout. Sternum 0.5× as 

wide as long; maximum width at coxa I. First femur length 

11.7 mm; with 5 prolateral macrosetae; patella + tibia I, 

14.2 mm; metatarsus I, 12.5 mm; tarsus I, 3 mm. Lengths 

of patella + tibia II, 11.4 mm; III, 5.2 mm; IV, 6.5 mm. 

Calamistrum 0.4 metatarsus IV length.

Male from Amanzimtoti, KwaZulu-Natal. Total 

length 8.5 mm. Carapace 3.8 mm long, 2.2 mm wide. Abdomen 5.0 mm long, 2.0 mm wide. Cephalic width 0.6× 

carapace width. PME diameter 0.25 mm; PME separation 2 diameters. AME/PME 0.7; AME separation 2.4 

AME diameters; equally spaced between PME and clypeal 

margin. ALE/PME 0.6; ALE tubercle large; ALE 2 diameters below clypeal margin. PLE/PME 0.7; PLE separation 

6 PLE diameters; PLE 1.7 diameters behind PME. Clypeus 1.2 AME diameters. Chelicerae with 4 promarginal 

teeth; 4 retromarginal teeth; additional 2 or 4 small denticles close to retromargin. Sternum 0.5× as wide as long. 

First femur length 10.5 mm; with 3 prolateral macrosetae; 

patella + tibia I 13.6 mm; metatarsus I 15.0 mm; tarsus 

I 4.3 mm. Lengths of patella + tibia II, 9.9 mm; III, 4.6 

mm; IV, 6.0 mm. Tarsus I divided, with proximal portion 

distinctly swollen. Terminal portion of metatarsus II with 

curve bordered posteriorly by stout seta. Dorsal surface 

of tarsus II with stout setae extending down most of its 

length. Palpal femur length 2.6× cymbium length.

Variation. In both sexes the usually unpaired 

abdominal tubercle varies in size and shape. A female collected at Gwala-Gwala had a paired tubercle (Figure 2a) 

unlike her conspecific from the same locality. Embolus 

relative length varies. Female body length ranges from 7.7 

to 19.5 mm (n = 31), and males from 7.2 to 9.5 (n = 9).

Natural History. Menneus camelus employs the usual deinopid hunting strategy including a 

backward strike (M. Kuntner, pers. obs.; Figures1d, 2a–c).

Distribution. South Africa (Figure 12).

Specimens Examined. SOUTH AFRICA:

Gauteng Prov., Johannesburg, dried branches in garden, 

M Filmer, 20.iii.1988, [Filmer Coll: 1 ♀; −26.2: 28.03]. 

Gauteng Prov., Northcliff, Johannesburg, in bushes 

in garden, L Prendini, 4.iv.1988, [Murphy Coll.: 1 ♂; 

−26.14: 27.97]. Gauteng Prov., outside Pretoria, in low 

bush near buildings, M Filmer, [Filmer Coll.: 1 ♂;−25.74: 

28.19]. Gauteng Prov., Pretoria, HK Monro, iii.1952, 

[NMP: NM 5886, 1 ♀; −25.74: 28.19]. Gauteng Prov., 

Pretoria, A Viljoen, 8.iv.1981, [SANC: AcAT/82/342/ 

1 ♀; −25.74: 28.19]. Gauteng Prov., Pretoria, in house, 

C Cilliers, iv.1986, [SANC: AcAT/86/40/, 1 ♀; −25.74: 

28.19]. Gauteng Prov., Pretoria, in garden, C. Kok, 

4.vi.1979, [SANC: AcAT/79/295/, 1 ♀; −25.74: 28.19]. 

Gauteng Prov., Pretoria, V/d. Mark, 10.x.1973, [SANC: 

AcAT/76/1714, 1 ♀; −25.74: 28.19]. Gauteng Prov., 

Pretoria North, M Greyling, [TM: 1 ♀; −25.74: 28.19]. 

Gauteng Prov., Pretoria, Agric. Building, in toilet, A vdBerg, [SANC: AcAT/83/322/, 1 ♂; −25.74: 28.19]. 

Gauteng Prov., Weavind Park, Pretoria, on wall, S vdBerg, 

30.iii.1985, [SANC: AcAT/85/133/, 1 ♀; −25.74: 28.19]. 

KwaZulu-Natal, Amanzimtoti, Inanda Nature Res., 

3030BB?, P Reavell, 20.iv.1981, [NMP: NM 14065, 1 

♂; −30.09: 30.85]. KwaZulu-Natal, Charters Creek, 

St. Lucia, forest, 28°12′S 32°26′E, Doyen & Griswold, 

19–21.xi.1985, [NMP: 1 ♀, 2 juveniles; −28.2: 32.43]. 
number 636 • 1 5

KwaZulu-Natal, Dukuduku, forest in deep shade, on 

herbs, 28°23′S 32°90E, P Reavell, 18.i.1980, [NMP: NM 

12952, 1 ♀; 28.38: 30.59]. KwaZulu-Natal, Greater St. 

Lucia Wetlands Park, Gwala Gwala forest, 28°23.042′S 

32°24.436E, Leg. Kuntner & al., 28.iv.2006, [USNM: 

1 ♀; −28.384: 32.4073]. KwaZulu-Natal, Greater St. 

Lucia Wetlands Park, Gwala Gwala forest, 28°23.042′S 

32°24.436′E, Leg. Kuntner & al., 28.iv.2006, [USNM: 

1 ♀; −28.384: 32.4073]. KwaZulu-Natal, Greytown, 

[RMCA: 146.144, 1 ♀, 4 juveniles; −29.06: 30.59]. 

KwaZulu-Natal, Kosi Bay, Toppin, ii.1906, [NMP: 

NM 2048, 1 ♂, 4 juveniles; −26.89: 32.88]. KwaZuluNatal, Mtunzini, Twin Streams Farm, coastal dune forest, 28°57′S 31°46′E, Griswold & al, 19.i.1984, [NMP: 

2 ♀; −28.95: 31.77]. KwaZulu-Natal, Natal [no further 

data], Coll CM, [MNHN: 1 ♂, 1 ♀, 5 juveniles; −28.52: 

30.9]. KwaZulu-Natal, Natal, Zululand [no further 

data], [MNHN: AR 166, 1 ♀, 8 juveniles; −28.52: 30.9]. 

KwaZulu-Natal, Ngotshe, Toggekry, 27°46′S 31°36′E, 

TW Schofield, 12.iv.1968, [NMP: NM 12406, 2 ♀; 

−27.77: 31.6]. KwaZulu-Natal, Pietermaritzburg, RF 

Lawrence, v.1957, [NMP: NM 6836, 1 ♀; −29.61: 30.37]. 

KwaZulu-Natal, Pietermaritzburg, RF Lawrence, iv.1957, 

[NMP: NM 5302, 1 ♀; −29.61: 30.37]. KwaZulu-Natal, 

Pietermaritzburg, Holliday, x.1944, [NMP: NM 4269, 

1 ♂; −29.61: 30.37]. KwaZulu-Natal, Pietermaritzburg, 

RF Lawrence, vii.1944, [NMP: NM 4261, 1 ♂; −29.61: 

30.37]. KwaZulu-Natal, Pietermaritzburg, E Pratt Yule, 

iv.1941, [NMP: NM 3350, 1 ♀; −29.61: 30.37]. KwaZuluNatal, Pietermaritzburg, Townbush forest, R Oberprieler, 

27.ii.1984, [SANC: AcAT/84/442/, 1 ♀; −29.61: 30.37]. 

KwaZulu-Natal, St, Lucia Game Reserve, Fanies Island, 

25m, M Ramirez, 31.iii–4.iv.2001, [AMNH: 1 ♂; −28.11: 

32.43]. KwaZulu-Natal, Stella Bush, HW Bell Marley, 

ii.1916, [SAM: B 2173, 1 ♀; −33.93: 18.85]. KwaZuluNatal, Tembe Elephant Park, 115m, M Ramirez, 9–12.

iv.2001, [AMNH: 1 ♀; −27.04: 32.42]. KwaZulu-Natal, 

Umkomaas River, [BMNH: 1 ♀, 4 juveniles; −30.2: 30.79]. 

KwaZulu-Natal, Winkel Spruit, 30°06′S 30°51′E, C Akerman, xii.1916, [NMP: NM 1913, 1 ♀; −30.1: 30.85]. 

Limpopo Prov., Meetse-a-Bophelo mission, on roof of 

bus, M Zwiegelaar, 11.iv.1977, [SANC: AcAT/77/891/, 1 

♀; −24.25: 30.45]. Mpumalanga, Badplaas Embuleni Nature Reserve, grassvelt savanna, in wooded area, 1100m, 

25°5712′′S 30°3315′′E, D & S Ubick, 28.iii.2001, [CAS: 

1 ♀; −25.953333: 30.554167]. [no data; TM: 8962, 1 

♀]. W Transvaal, Rustenberg, AL Capener, ii.1965, 

[NMP: NM 9431, 1 ♀; −33.5: 24.5]. Weenen, TD Walsh, 

25.vii.1934, [NMP: 1511 Weenen Walsh, 1 ♀; −28.85: 

30.08].

Menneus capensis (Purcell, 1904), 

new combination

FIGURES 1a–c, 12, 14d–f, 20

Avellopsis capensis Purcell 1904:128 (♂♀).—Lehtinen, 1967:217, figs. 33, 

37 (transfer from Uloboridae).—Platnick, 2010.

Types. Lectotype female in SAM hereby designated, examined. Vial with lectotype contains three labels: 

one in ink stating “Dinopidae TYPE Avellopsis capensis 

Purcell ACC. NO. 12330 SAM/ARAN 3176. Side of Kalk 

Bay Mountain. N.F. Purcell. 3/1902”; one photocopied stating “MUS CAPE TOWN lectotype/PTL\Avellopsis capensis 

Purcell, 1904.”; and one in pencil stating “12330 Side of 

Kalk Bay Mountain 3/02 by W.F.P. of Avellopsis capensis 

Purc.” Paralectotype female from Camps Bay is immature.

Diagnosis. Female Menneus capensis differ 

from all other African species by the diamond-shaped 

abdomen with paired abdominal tubercle (Figure 20c,d). 

Copulatory ducts with one and a half spirals (or two counting lateral margins in dorsal view) (Figure 20f). The median hematodocha in ventral view (left palp) spans 12–4 

OC, the embolus tip lies at about 11 OC, and the median 

apophysis–basal lobe axis lies at about 8:30 OC (Figure 

14e), is narrow, and is gently curved in prolateral view 

(Figure 14e,f). The embolus makes 2.5 coils, and is bent 

away from the tegulum in prolateral view (Figure 14f).

Description. Female from Newlands Forest, 

Cape Town, South Africa. Total length 10 mm. Carapace 

3.4 mm long; 2.5 mm wide. Abdomen 6.6 mm long; 4.8 

mm wide. Cephalic width 0.7× carapace width. PME diameter 0.3 mm; PME separation 1.2 diameters. AME/

PME 0.5; AME separation 1.9 AME diameters; closer to 

PME than to clypeal margin. ALE/PME 0.5; ALE tubercle 

large; ALE 1 diameter below clypeal margin. PLE/PME 

0.8; PLE separation 5.2 PLE diameters; PLE 1.4 diameters 

behind PME. Clypeus 2 AME diameters. Chelicerae with 

4 promarginal teeth; 4 retromarginal teeth; additional 4 

small denticles close to retromargin. Sternum 0.8× as wide 

as long; maximum width at coxa I. First femur length 6.6 

mm; with 5 prolateral macrosetae; patella + tibia I, 7.2 

mm; metatarsus I, 6.4 mm; tarsus I, 1.4 mm. Lengths of 

patella + tibia II, 6.5 mm; III, 4.1 mm; IV, 4.2 mm. Calamistrum 0.3 metatarsus IV length.

Male from Cape Province, Goudini. Total length 6.5 

mm. Carapace 3.2 mm long; 2 mm wide. Abdomen 3 mm 

long; 1.4 mm wide. Cephalic width 0.5× carapace width. 

PME diameter 0.2 mm; PME separation 1.2 diameters. 

AME/PME 0.5; AME separation 1.8 AME diameters; 
1 6 • smithsonian contributions to zoology

closer to clypeal margin than to PME. ALE/PME 0.6; 

ALE tubercle large; eye’s ventral edge 2/3 or more diameters from clypeus margin; ALE 0.6 diameters below 

clypeal margin. PLE/PME 0.5; PLE separation 4 PLE 

diameters; PLE 2.2 diameters behind PME. Clypeus 0.8 

AME diameters. Chelicerae with 2 promarginal teeth; 3 

retromarginal teeth; median frontal macrosetae distal to 

laterals; laterals attenuate. Sternum 0.9× as wide as long. 

First femur length 9.4 mm; with 4 prolateral macrosetae; 

patella + tibia I 10.5 mm; metatarsus I 12.1 mm; tarsus I 

1.9 mm. Lengths of patella + tibia II, 8.6 mm; III, 4.8 

mm; IV, 5.2 mm. Tarsus I undivided. Terminal portion 

of tibia II unmodified. Terminal portion of metatarsus II 

without distal curve in dorsal view. Dorsal surface of tarsus II without stout setae. Palpal femur length 2.1× cymbium length.

Variation. Female body length ranges from 

8.5 to 11.4 mm (n = 7), and males from 5.0 to 7.0 (n = 3).

Natural History. The species inhabits forests around Table Mountain, Cape Town (N. Larsen, pers. 

comm.). We have recently collected fresh material from 

Newlands Forest, Cape Town. The web and predatory 

habits are typical of deinopids (Figure 1a–c). The spiders 

were found to build their webs close to the ground, typically under boulders adjacent to water.

Distribution. Western Cape, South Africa 

(Figure 12).

Specimens Examined. SOUTH AFRICA:

Cape Prov., Camps Bay, WF Purcell, [SAM: 748 SAM/

ARAN 3177, 1 juvenile; −33.95: 18.37]. Cape Prov., 

Cape Town, Newlands Forest, Paradijs Stream, N Larsen, 

9.v.1991, [SAM: C 2453, 2 ♀; −33.98: 18.44]. Cape Prov., 

Cape Town, Newlands Forest, Paradijs Stream, N Larsen, 

v.1991, [SAM: C 1864, 1 juvenile; −33.98: 18.44]. Cape 

Prov., Cape Town, Newlands Forest, Paradijs Stream 

(NL153), N Larsen, 9.v.1991, [SAM: C 1858, 1 ♀; 

−33.98: 18.44]. Cape Prov., Cape Town, Newlands Forest, Paradijs Stream (NL154), N Larsen, 9.v.1991, [SAM: 

C 1859, 1 ♂; −33.98: 18.44]. Cape Prov., Kalk Bay Mountains, WF Purcell, iii.1902, [SAM: Acc. No. 12330 SAM/

ARAN 3043, 14 juveniles; −33.93: 18.46]. Cape Prov., 

side of Kalk Bay Mountain, WF Purcell, iii.1902, [SAM: 

Acc. no. 12330 SAM/ARAN 3176, 1 ♂; −33.93: 18.46]. 

Western Cape, Cape Town, Table Mtn. NP, Newlands 

Forest, 159 m, 33°58.934′S 18°26.686′E, M Kuntner & 

al, 4.v.2006, [USNM: 1 ♀; −33.9822: 18.4448]. Western Cape, Cape Town, Table Mtn. NP, Newlands Forest, 159 m, 33°58.934′S 18°26.686′E, M Kuntner & al, 

4.v.2006, [USNM: 1 ♂; −33.9822: 18.4448]. Western 

Cape, Cape Town, Table Mtn. NP, Newlands Forest, 159 

m, 33°58.934′S 18°26.686′E, M Kuntner & al, 4.v.2006, 

[USNM: 1 ♀; −33.9822: 18.4448]. Western Cape, Capetown, Newland Forest Preserve, SE Table Mtn., indigenous 

forest, 33°58′S 18°28′E, N Larsen, K Muller, S Prinsloo, D 

& S Ubick, 4.iv.2001, [CAS: 1 ♂, 1 ♀; −35.08: 18.47]. 

Western Cape, Table Mountain, Capetown, Fernwood 

Gully, indigenous forest, 150 m, 33°58′S 18°27′E, CE 

Griswold, 18.xii.1996, [CAS: 1 ♀; −33.97: 18.53].

Menneus darwini, new species

FIGURES 12, 14g–i, 21

Types. Holotype male: TANZANIA: Iringa, 

Mufindi Dist., Uzungwa Scarp For. Res., canopy fog, 1515 

m, 08°30′S 35°52′E, McKamey & al, 4.iii.1996, [ZMUC: 

1 ♂; −8.5: 35.87].

Etymology. Named for Charles Darwin in 

honor of the 150th anniversary of the publication of On

the Origin of Species and the 200th anniversary of his 

birth. Gender is masculine.

Diagnosis. Male Menneus darwini is most 

similar to M. dromedarius, M. tetragnathoides, M. camelus, and M. samperi among African species. The median 

hematodocha in ventral view (left palp) spans 4:30–8 OC, 

unlike any other African species, the embolus tip lies at 

about 10:30 OC, and the median apophysis–basal lobe 

axis lies at about 1 OC (Figure 14h). The embolus makes 

3.5 coils, and the tip is straight rather than bent away 

from the tegulum in prolateral view (Figure 14i).

Description. Female unknown.

Male (holotype). Total length 8.7 mm. Carapace 3.5 

mm long. Carapace 2.4 mm wide. Abdomen 5.0 mm 

long. Abdomen 2.5 mm wide. Cephalic width 0.4× carapace width. PME diameter 0.3 mm; PME separation 1.7 

diameters. AME/PME 0.7; AME separation 1.8 AME diameters; equidistant between PME and clypeal margin. 

ALE/PME 0.7; ALE tubercle large; eye’s ventral edge 2/3 

or more diameters from clypeus margin; ALE 1 diameter 

below clypeal margin. PLE/PME 0.8; PLE separation 5.2 

PLE diameters; PLE 1.3 diameters behind PME. Clypeus 

0.8 AME diameters. Chelicerae with 3 promarginal teeth; 

7 retromarginal teeth; median frontal macrosetae absent 

(or inconspicuous); laterals attenuate. Sternum 0.5× as 

wide as long. First femur length 10.7 mm; with 4 prolateral macrosetae; patella + tibia 12.4 mm; metatarsus I 

12.7 mm; tarsus I 3.5 mm. Lengths of patella + tibia II, 

9.1 mm; III, 3.7 mm; IV, 4.9 mm. Tarsus I divided. Terminal portion of tibia II with stout condyles and prolateral 

apophysis. Terminal portion of metatarsus II with curve 
number 636 • 1 7

bordered posteriorly by stout seta. Dorsal surface of tarsus II with stout setae extending down most of its length. 

Palpal femur length 2.3× cymbium length.

Variation. Known only from holotype male.

Natural History. Unknown.

Distribution. Only known from the type locality, Tanzania.

Menneus dromedarius Purcell, 1904

FIGURES 12, 15a–c, 22

Menneus dromedarius Purcell, 1904:127 (♀).—Lehtinen, 1967:247, fig. 40. 

Here removed from synonymy of M. camelus (contra Lehtinen, 1967).

Types. Holotype female from Pirie Bush, King 

Williamstown, Natal, South Africa, in SAM, spec. no. 

11536, examined. Type label reads: “ . . . F. A. Pym, 

March, 1902.”

Note. Four females recently collected in Madagascar fit well into the known variation of South African 

M. dromedarius.

Diagnosis. Female M. dromedarius differ 

from other known African females by having copulatory 

ducts with four full spirals (or five counting lateral margins in dorsal view; Figure 22g). The median hematodocha 

in ventral view (left palp) spans 6–10 OC, unlike any other 

African species, the embolus tip lies at about 11:30 OC, 

and the median apophysis–basal lobe axis lies at about 2 

OC (Figure 15b). The embolus makes 3.5 coils and is bent 

away from the tegulum in prolateral view (Figure 15c).

Description. Female from Kleinmond, Cape 

Province, South Africa. Total length 8.8 mm. Carapace 

3.2 mm long; 1.9 mm wide. Abdomen 5.4 mm long; 3.0 

mm wide. Cephalic width 0.8× carapace width. PME diameter 0.23 mm; PME separation 2.6 diameters. AME/

PME 0.6; AME separation 2.1 AME diameters; closer to 

PME than to clypeal margin. ALE/PME 0.7; ALE tubercle 

large; ALE 2 diameters below clypeal margin. PLE/PME 

0.8; PLE separation 7.4 PLE diameters; PLE 2 diameters 

behind PME. Clypeus 3 AME diameters. Chelicerae with 

4 promarginal teeth; 4 or 5 retromarginal teeth. Sternum 

0.5× as wide as long; maximum width at coxa I. First 

femur length 6.5 mm; with 3 prolateral macrosetae; patella + tibia I, 7.5 mm; metatarsus I, 6.2 mm; tarsus I, 1.6 

mm. Lengths of patella + tibia II, 5.4 mm; III, 2.6 mm; IV, 

3.5 mm. Calamistrum 0.5 metatarsus IV length.

Male from Grahamstown. Total length 10 mm. Carapace 4.7 mm long. Carapace 3 mm wide. Abdomen 5.1 

mm long. Abdomen 2.2 mm wide. Cephalic width 0.6× 

carapace width. PME diameter 0.3 mm; PME separation 

1.8 diameters. AME/PME 0.6; AME separation 1.7 AME 

diameters; closer to PME than to clypeal margin. ALE/

PME 0.5; ALE tubercle large; eye’s ventral edge 2/3 or 

more diameters from clypeus margin; ALE 1.9 diameters 

below clypeal margin. PLE/PME 0.7; PLE separation 7.3 

PLE diameters; PLE 1.8 diameters behind PME. Clypeus 

0.9 AME diameters. Chelicerae with 3 promarginal teeth; 

1 retromarginal teeth; median frontal macrosetae distal to 

laterals; laterals attenuate. Sternum 0.5× as wide as long. 

First femur length 11.9 mm; with 4 prolateral macrosetae; 

patella + tibia I 15.7 mm; metatarsus I 16 mm; tarsus I 5.2 

mm. Lengths of patella + tibia II, 11.4 mm; III, 5.5 mm; IV, 

6.9 mm. Tarsus I divided, with proximal portion distinctly 

swollen. Terminal portion of tibia II with stout condyles 

and prolateral apophysis. Terminal portion of metatarsus 

II with curve bordered posteriorly by stout seta. Dorsal 

surface of tarsus II with stout setae extending down most 

of its length. Palpal femur length 2.8× cymbium length.

Variation. In both sexes the unpaired abdominal tubercle varies in size and shape. Female body length 

ranges from 10.0 to 17.5 mm (n = 5), and males from 10.0 

to 13.3 (n = 5).

Natural History. Unknown.

Distribution. South Africa, Madagascar (Figure 12). Copenhagen (ZMUC) has a specimen of M. dromedarius with a label saying only “Cameroon” and no other 

data. At this time we regard this as a locality error.

Specimens Examined. MADAGASCAR:

Toliara, Réserve Privé Berenty, Forêt de Bealoka, Mandraré 

River, 14.6 km 329° NNW Ambooasary, gallery forest, general collecting night, 35 m, 24°57′25′′S 46°16′17′′E, BL Fisher 

et al. BLF5319, 3–8.ii.2002, [CAS: CASENT9012443, 1 ♀; 

−24.956944: 46.271389]. Toliara, Réserve Privé Berenty, 

Forêt de Bealoka, Mandraré River, 14.6 km 329° NNW 

Amboasary, ER30 beating low vegetation, 35 m, 24°57′25′′S 

46°16′17′′E, BL Fisher et al. BLF5318, 3–8.ii.2002, [CAS: 

CASENT9012406, 1 ♀; −24.956944: 46.271389]. Toliara, Réserve Privé Spéciale de Cap Sainte Marie 12.3 km 

262°W Marovato, spiny forest/thicket EH21 beating low 

vegetation, 200 m, 25°34′54′′S 45°10′6′′E, BL Fisher et 

al. BLF5502, [CAS: CASENT9012629, 1 ♀, 1 juvenile; 

−25.581667: 45.168333]. Toliara, Réserve Privé Spéciale de Cap Sainte Marie, 12.3 km 262°W Marovato, 

spiny forest/thicket EH21 beating low vegetation, 200 m, 

25°34′54′′S 45°10′6′′E, BL Fisher et al. BLF5505, [CAS: 

CASENT9012682, 1 ♀; −25.581667: 45.168333]. SOUTH 

AFRICA: Cape Prov., Diepwalle Forest Sta., 22 km NE 

Knysna, indigenous forest, 1800 ft, 33°57′S 23°10′E, Griswold & Griswold, 10.i.1985, [NMP: 1 ♂; −33.95: 23.17]. 
1 8 • smithsonian contributions to zoology

Cape Prov., Kleinmond, on ground in fynbos vegetation, 

S Neser, 16.xii.1985, [SANC: AcAT/86/13/, 1 ♀; −34.34: 

19.02]. Eastern Cape, Grahamstown, behind bookcase, 

33°18′S 26°32′E, PMC Croeser, 26.i.1979, [NMP: 1 ♂; 

−33.3: 26.53]. KwaZulu-Natal, Ngoyi Forest, Lawrence 

& al., ii.1953, [NMP: NM 5894, 1 ♂, 1 juvenile; −28.83: 

31.66]. Transvaal, Barberton, 25°47′S 31°03′E, RF Lawrence, iii.1960, [NMP: NM 7966, 1 ♂; −25.78: 31.05]. 

Fairy Glen Farm, Bathurst, forest floor, RF Lawrence, 

25.i.1980, [SANC: AcAT/82/127/, 1 ♂; −33.5: 26.82].

Menneus samperi, new species

FIGURES 12, 15d–f, 23

Type. Holotype male in NMP from Kenya, Kaibos; see locality data below.

Etymology. Named for Cristián Samper K., 

Director of the National Museum of Natural History, 

Smithsonian Institution, in recognition of his leadership and 

contributions to biodiversity science. Gender is masculine.

Diagnosis. Female Menneus samperi differ 

from other known African females by having copulatory 

ducts with three full spirals (or four counting lateral margins in dorsal view) (Figure 23g). The median hematodocha 

in ventral view (left palp) spans 8–11 OC, the embolus tip 

lies at about 12 OC, and the median apophysis–basal lobe 

axis lies at about 2 OC (Figure 15e). The embolus makes 

3.5 coils and is bent away from the tegulum in prolateral 

view (Figure 15f). Both males and females have noticeably 

long third femoral setal fringes.

Description. Female from Tanzania. Total 

length 10.5 mm. Carapace 4.4 mm long; 2.9 mm wide. 

Abdomen 6.1 mm long; 4.3 mm wide. Cephalic width 

0.7× carapace width. PME diameter 0.23 mm; PME separation 3.3 diameters. AME/PME 0.8; AME separation 2.2 

AME diameters; equidistant between PME and clypeal 

margin. ALE/PME 0.7; ALE tubercle large; ALE 2.7 diameters below clypeal margin. PLE/PME 1.0; PLE separation 7.8 PLE diameters; PLE 1.7 diameters behind PME. 

Clypeus 1 AME diameters. Chelicerae with 4 promarginal 

teeth; 7 retromarginal teeth. Sternum 0.5× as wide as long; 

maximum width at coxa I. First femur length 10.6 mm; 

with 3 prolateral macrosetae; patella + tibia I, 12.1 mm; 

metatarsus I, 11.3 mm; tarsus I missing. Lengths of patella 

+ tibia II, 10.0 mm; III, 4.6 mm; IV, 5.4 mm. Calamistrum 

0.3 metatarsus IV length.

Male (holotype). Total length 7.6 mm. Carapace 

3.3 mm long. Carapace 2.2 mm wide. Abdomen 4.1 

mm long. Abdomen 2.2 mm wide. Cephalic width 0.6× 

carapace width. PME diameter 0.19 mm; PME separation 2.6 diameters. AME/PME 0.97; AME separation 1.9 

AME diameters; closer to clypeal margin than PME. ALE/

PME 0.8; ALE tubercle large; eye’s ventral edge 2/3 or 

more diameters from clypeus margin; ALE 1.8 diameter 

below clypeal margin. PLE/PME 1.1; PLE separation 5.8 

PLE diameters; PLE 1.0 diameters behind PME. Clypeus 

0.8 AME diameters. Chelicerae with 4 promarginal teeth; 

1 retromarginal tooth; median frontal macrosetae absent 

(or inconspicuous); laterals attenuate. Sternum 0.6× as 

wide as long. First femur length 8.5 mm; prolateral macrosetae absent; patella + tibia 11.0 mm; metatarsus I 11.4 

mm; tarsus I 3.2 mm. Lengths of patella + tibia II, 8.2 mm; 

III, 3.4 mm; IV, 4.3 mm. Tarsus I divided. Terminal portion of tibia II with stout condyles and prolateral apophysis. Terminal portion of metatarsus II with curve bordered 

posteriorly by stout seta. Dorsal surface of tarsus II with 

stout setae extending down half of its length. Palpal femur 

length 2.0× cymbium length.

Variation. Only one male known. Female 

body length ranges from 13.0 to 15.0 mm (n = 2).The 

Uganda female had 4–5 first femoral macrosetae.

Natural History. The Uganda female was 

collected with a typical deinopid spherical eggsac, 9 mm 

in diameter.

Distribution. Eastern Africa (Kenya, Tanzania, Uganda; Figure 12).

Specimens Examined. KENYA: Kaibos, indigenous forest, 01°12′N 35°08′E, B Lamoral, 23.v.1980, 

[NMP: 1 ♂; 1.2: 35.13]. TANZANIA: Bukoba Dist., 

Minzero Forest Reserve, 01°05′S 31°30′E, M Andersen, 

x.1998, [ZMUC: 1 ♀; −1.08: 31.5]. UGANDA: Kabale 

Dist., Ruhizha Bwindi Impenetrable NP, “Nature Trail,” 

forest, 2300 m, 01°03′S 29°47′E, CE Griswold, 13–16.

ix.1996, [CAS: 1 ♀; −1.05: 29.78].

Menneus tetragnathoides Simon, 1876

FIGURES 12, 15g–i, 24

Menneus tetragnathoides Simon 1876:219 (♂).—Simon, 1892:210, figs. 

155, 156.—Platnick, 2010.

Menneus affinis Tullgren, 1910:91, pl. 1, fig. 2 (♂).—Platnick, 2010. NEW 

SYNONYMY.

Notes. Platnick (2010) notes that Berland 

(1914:53) considered M. affinis as a junior synonym 

of M. tetragnathoides, which was not accepted subsequently. Our examination of the holotype confirms such 

synonymy.
number 636 • 1 9

Types. Menneus tetragnathoides syntype series 

of one male and two immatures in MNHN, examined, lectotype male designated here. Vial with lectotype contains 

two labels, one stating “AR 163” and the other “Museum 

Paris AE 163Menneustetragnathoides Sim. Congo TYPE.” 

According to Simon (1876) the type material was collected 

by Louis Petit ca. 1876 at “Chinchonxo in Congo” (today 

in Angola, Cabinda, Chinchoxo, –5.166667: 12.116667). 

The remaining specimens have been designated as paralectotypes.

Holotype male M. affinis in SMNH, examined, label 

reads “Kilimandjaro, Kibonoto, Colleg. Y. Sjot.”

Diagnosis. The median hematodocha in ventral view (left palp) spans 3–6 OC, the embolus tip lies 

at about 11:30 OC, and the median apophysis–basal lobe 

axis lies at about 1 OC (Figure 15h). The embolus makes 

3.5 coils and is relatively straight and parallel to the tegulum in prolateral view (Figure 15i).

Description. Female unknown.

Male from Malawi, Chintheche. Total length 9.8 

mm. Carapace 4.2 mm long; 2.3 mm wide. Abdomen 5.4 

mm long; 2.6 mm wide. Cephalic width 0.3× carapace 

width. PME diameter 0.3 mm; PME separation 1.8 diameters. AME/PME 0.7; AME separation 1.4 AME diameters; closer to clypeal margin than to PME. ALE/PME 

0.6; ALE tubercle large; eye’s ventral edge 2/3 or more 

diameters from clypeus margin; ALE 1.4 diameters below 

clypeal margin. PLE/PME 0.8; PLE separation 7.2 PLE 

diameters; PLE 1.3 diameters behind PME. Clypeus 0.7 

AME diameters. Chelicerae with 4 promarginal teeth; 1 

retromarginal teeth; frontal macrosetae absent. Sternum 

0.5× as wide as long. First femur length 10.5 mm; with 3 

prolateral macrosetae; patella + tibia I 15.3 mm. Lengths 

of patella + tibia II, 12.3 mm; III, 4.6 mm; IV, 5 mm. 

Terminal portion of tibia II with stout condyles and prolateral apophysis. Terminal portion of metatarsus II with 

distal curve in dorsal view, bordered by prolateral stout 

seta. Dorsal surface of tarsus II with stout setae extending 

down most of its length. Palpal femur length 3× cymbium 

length.

Variation. Male body length ranges from 9.5 

to 11.0 mm (n = 2).

Natural History. Unknown.

Distribution. West and East Africa (Cabinda, Malawi, Kilimanjaro) (Figure 12).

Specimens Examined. ANGOLA: Cabinda, “Chinchonxo,” 05°10′S 12°07′E, Louis Petit, ca. 

1876, [MNHN: AE 163, 1 ♂, 2 juveniles; −5.17: 12.12]. 

MALAWI: Chintheche, R Jocque, 2.v.1978, [RMCA: no. 

153.633, 1 ♂; −11.83: 34.16]. TANZANIA: Kilimanjaro, 

Kibonoto [Kibongoto], Y. Sjot, xii.1905 [SMNH: 1 ♂; 

−3.18: 37.09].

Australasian Species

Menneus aussie, new species

FIGURES 13, 16a–c, 25

Etymology. “Aussie” is a fifth declension 

feminine Latin noun, meaning “from Australia.”

Types. Holotype male designated here, in QM, 

from Rundle Range; see below.

Diagnosis. Female M. aussie have paired median dorsal abdominal tubercles, like M. superciliosus, M. 

nemesio, M. quasimodo, and M. wa, but, uniquely, have 

copulatory ducts with five full spirals (or six counting 

lateral margins in dorsal view; Figure 25g). The median 

hematodocha in ventral view (left palp) spans 6–9 OC, 

the embolus tip lies at about 11:30 OC, and the median 

apophysis–basal lobe axis lies at about 2 OC (Figure 16b). 

The embolus makes 3.25 coils and is bent away from the 

tegulum in prolateral view (Figure 16c).

Description. Female from Australia, NSW, 

“Avalon Bch.” Total length 10.6 mm. Carapace 4 mm 

long; 3 mm wide. Abdomen 6.8 mm long; 4.2 mm wide. 

Cephalic width 0.5× carapace width. PME diameter 0.3 

mm; PME separation 1.5 diameters. AME/PME 0.7; AME 

separation 1.4 AME diameters; equidistant between PME 

and clypeal margin. ALE/PME 0.6; ALE tubercle large, 

eye’s lower edge 2/3 or more diameters from clypeus 

margin; ALE 0.7 diameters below clypeal margin. PLE/

PME 0.9; PLE separation 4.7 PLE diameters; PLE 1.6 diameters behind PME. Clypeus 1 AME diameters; lacking 

large marginal macrosetae. Chelicerae with 4 promarginal 

teeth; 5 retromarginal teeth; median frontal macrosetae 

proximal to laterals. Sternum 0.7× as wide as long; width 

at coxae I and II about equal. First femur length 8.9 mm; 

with 6 to 7 prolateral macrosetae; patella + tibia I, 9.7 

mm; metatarsus I, 10 mm; tarsus I, 1.8 mm. Lengths of 

patella + tibia II, 9.1 mm; III, 5.1 mm; IV, 5.2 mm. Calamistrum 0.4 metatarsus IV length.

Male (holotype) from Australia, ME. Queensland, 

Rundle Range. Total length 10.5 mm. Carapace 4 mm 

long. Carapace 2.9 mm wide. Abdomen 6.5 mm long. Abdomen 3 mm wide. Cephalic width 0.4× carapace width. 

PME diameter 0.3 mm; PME separation 1.1 diameters. 

AME/PME 0.6; AME separation 1.5 AME diameters; 

closer to clypeal margin than to PME. ALE/PME 0.7; ALE 

tubercle large; eye’s ventral edge 2/3 or more diameters 
2 0 • smithsonian contributions to zoology

from clypeus margin; ALE 1.3 diameters below clypeal 

margin. PLE/PME 0.9; PLE separation 3.4 PLE diameters; 

PLE 1.8 diameters behind PME. Clypeus 1 AME diameters. Chelicerae with 2 promarginal teeth; 1 retromarginal 

teeth; median frontal macrosetae proximal to laterals; laterals attenuate. Sternum 0.7× as wide as long. First femur 

length 13.4 mm; with 6 prolateral macrosetae; patella + 

tibia I 14.9 mm; metatarsus I 16.2 mm; tarsus I 4.3 mm. 

Lengths of patella + tibia II, 12.2 mm; III, 7 mm; IV, 7.2 

mm. Tarsus I divided and of uniform width. Terminal portion of tibia II with stout condyles and prolateral apophysis. Terminal portion of metatarsus II with curve bordered 

posteriorly by stout seta. Dorsal surface of tarsus II with 

stout setae extending down most of its length. Palpal 

femur length 2.9× cymbium length.

Variation. The female from New Caledonia 

lacks the typical abdominal tubercles, which are consistently present in Australian specimens. Female body length 

ranges from 9.0 to 12.0 mm (n = 6), and males from 9.3 

to 9.8 (n = 2).

Natural History. Unknown.

Distribution. Australia (Queensland, New 

South Wales), New Caledonia (Figure 13).

Specimens Examined. AUSTRALIA: NSW, 

Avalon Bch., B Collins, 11.xi.1972, [AMS: KS 16841, 1 

♀; −33.64: 151.33]. NSW, Manly, NO Fletcher, 6.v.1945, 

[AMS: KS 16840, 1 ♀; −33.8: 151.29]. QLD, Dandabah, Bunya NP, VED, 6.iii.1976, [QM: 2 ♀; −27.36: 

152.93]. QLD, Lamington NP, OReillys, night collecting, 

RR&VED, 16.xi.1977, [QM: 1 juvenile; −28.22: 153.13]. 

QLD, Rundle Range, 24.iii.1975, [QM: 4 ♂, 3 ♀; −23.65: 

150.97]. QLD, Rundle Range, R Kohout, 24.iii.1975, 

[QM: 2 ♂, 2 ♀; −23.65: 150.97]. NEW CALEDONIA: Col 

dAmieu Forest Stn., 400 m, Monteith & Cook, 8.v.1984, 

[QM: 1 ♀; −21.58: 165.76].

Menneus bituberculatus, new species

FIGURES 13, 26

Types. Holotype female designated here, in 

QM, from Upper Brookfield; see below.

Etymology. Named for the bifid posterior 

abdominal tubercles, an adjective.

Diagnosis. Uniquely in Menneus, symmetric abdominal tubercles in M. bituberculatus are doubly 

paired (Figure 26c). Female M. bituberculatus have copulatory ducts with two full (or three if counting lateral margins) spirals (Figure 26g).

Description. Female (holotype). Total length 

11.5 mm. Carapace 3.8 mm long; 2.6 mm wide. Abdomen 7.5 mm long; 3.3 mm wide. Cephalic width 1.5× 

carapace width. PME diameter 0.2 mm; PME separation 

3.7 diameters. AME/PME 0.8; AME separation 1.8 AME 

diameters; equidistant between PME and clypeal margin. 

ALE/PME 0.7; ALE tubercle small, eye’s lower edge contiguous with clypeus margin; ALE 0.8 diameters below 

clypeal margin. PLE/PME 1; PLE separation 6.1 PLE 

diameters; PLE 0.6 diameters behind PME. Clypeus 0.2 

AME diameters. Chelicerae with 4 promarginal teeth; 4 

retromarginal teeth; frontal macrosetae in transverse line; 

laterals attenuate. Sternum 0.6× as wide as long; maximum width at coxa II. First femur length 7.8 mm; with 

6 prolateral macrosetae; patella + tibia I, 9.1 mm; metatarsus I, 8.9 mm; tarsus I, 1.8 mm. Lengths of patella + 

tibia II, 7.8 mm; III, 3.3 mm; IV, 3.7 mm. Calamistrum 

0.4 metatarsus IV length.

Male. Unknown.

Variation. Unknown.

Natural History. Unknown.

Distribution. Australia (Queensland), 

probably New Guinea, based on immature male record 

(Figure 13).

Specimens Examined. AUSTRALIA: QLD, 

Upper Brookfield, pit traps, rainforest, Raven & Green, 

6.x.1981, [QM: 1 ♀, 1 juvenile; −27.49: 152.86]. INDONESIA: West Papua, Hollandia [Jayapura], rain forest, 250ft, H Hoogstraal, xii.1944, [AMNH: 1 juvenile; 

−2.53: 140.71], identification uncertain.

Menneus superciliosus (Thorell, 1881)

FIGURES 13, 17a–c, 30

Avella superciliosa Thorell, 1881:200 (♀).—Lehtinen, 1967: 217 (transfer 

from Menneus). —Platnick, 2010.

Menneus superciliosus Rainbow, 1911:124.

Types. Adult female from Somerset, Cape York, 

in MCSNG, examined. Vial with holotype contains three 

labels, one in pencil stating “Avella superciliosa Thor. 

Typus Ragni Austr. Males. p. 200. sp. 122 Somerset (Cape 

York) 1875. rac. D’Alb.,” one in ink “Avella superciliosa 

Typus! Thorell (Ann. Mus. XVII, p. 200) Somerset (Capo 

York) 1875 – L. M. D’Albertis,” and one in ink stating, stating “Museo Civico di Storia Naturale di Genova 

Avella superciliosa Typus! Thorell Somerset (Capo York) 

leg. L. M. D’Albertis, 1875.”
number 636 • 2 1

Diagnosis. Among Australian Menneus with 

paired median dorsal abdominal tubercles female M. superciliosus closely resemble M. wa in having copulatory 

ducts with two full spirals (or three counting lateral margins in dorsal view) (Figure 30g). Female M. superciliosus

and M. wa, apart from their apparently disjunct distributions, are difficult to distinguish. Menneus superciliosus

seem to have narrower, more elongate abdomens and lack 

the brown lateral abdominal spot (Figure 30c,d). The median hematodocha in ventral view (left palp) spans 4–6 

OC, the embolus tip lies at about 1 OC, and the median 

apophysis–basal lobe axis lies at about 2:30 OC (Figure 

17b). The embolus makes 3.5 coils and is bent away from 

the tegulum in prolateral view (Figure 17c).

Description. Female (paratype) from Mount 

Cook. Total length 10.7 mm. Carapace 4.2 mm long; 2.7 

mm wide. Abdomen 5.1 mm long; 4.3 mm wide. Cephalic 

width 0.7× carapace width. PME diameter 0.3 mm; PME 

separation 1.7 diameters. AME/PME 0.8; AME separation 

1.5 AME diameters; closer to PME than to clypeal margin. 

ALE/PME 0.9; ALE tubercle large, eye’s lower edge 2/3 or 

more diameters from clypeus margin; ALE 1.6 diameters 

below clypeal margin. PLE/PME 0.8; PLE separation 3.8 

PLE diameters; PLE 1.8 diameters behind PME. Clypeus 

1.8 AME diameters. Chelicerae with 5 promarginal teeth; 

5 retromarginal teeth; median frontal macrosetae proximal to laterals; laterals stout. Sternum 0.7× as wide as 

long; maximum width at coxa I. First femur length 9.4 

mm; with 4 prolateral macrosetae; patella + tibia I, 12.6 

mm; metatarsus I, 10.8 mm; tarsus I, 1.9 mm. Lengths of 

patella + tibia II, 9.8 mm; III, 5.3 mm; IV, 5.5 mm. Calamistrum 0.3 metatarsus IV length.

Male (holotype) from Mt. Cook. Total length 8 mm. 

Carapace 3.4 mm long. Carapace 2.4 mm wide. Abdomen 

4.5 mm long. Abdomen 1.7 mm wide. Cephalic width 0.5× 

carapace width. PME diameter 1.1 mm; PME separation 

1.4 diameters. AME/PME 0.8; AME separation 1.3 AME 

diameters; equidistant between PME and clypeal margin. 

ALE/PME 0.6; ALE tubercle large; eye’s ventral edge 2/3 

or more diameters from clypeus margin; ALE 0.6 diameters below clypeal margin. PLE/PME 0.8; PLE separation 

7 PLE diameters; PLE 1.6 diameters behind PME. Clypeus 

1 AME diameters. Chelicerae with 6 promarginal teeth; 6 

retromarginal teeth; median frontal macrosetae distal to 

laterals; laterals attenuate. Sternum 0.5× as wide as long. 

First femur length 9.8 mm; with 7 prolateral macrosetae; 

patella + tibia I 11.1 mm; metatarsus I 13 mm; tarsus I 2.5 

mm. Lengths of patella + tibia II, 9.2 mm; III, 4.9 mm; IV, 

5 mm. Tarsus I divided and of uniform width. Terminal 

portion of tibia II with stout condyles and prolateral 

apophysis. Terminal portion of metatarsus II without distal curve. Dorsal surface of tarsus II without stout setae. 

Palpal femur length 2.6× cymbium length.

Variation. Female body length ranges from 

7.2 to 13.5mm (n = 21), and males from 7.0 to 9.5 

(n = 9).

Natural History. Label data suggest the 

standard deinopid web.

Distribution. Australia (Queensland, New 

South Wales; Figure 13).

Specimens Examined. AUSTRALIA:

NSW, Royal Nat Park, R Mascord, 9.xii.1967, [AMS: 

KS 16861, 1♂; −34.12: 151.06]. NSW, Royal Nat Park, 

R Mascord, 9.xii.1967, [AMS: KS 16836 2♀; −34.12: 

151.06]. NSW, Royal Nat Park, A Speechley, 14.iii.1969, 

[AMS: KS 16847, 1♂; −34.12: 151.06]. NSW, Tarre, MR 

Gray FN1839, [AMS: KS 9371, 1♂; −31.91: 152.46]. 

QLD, 2.5 km SW Mt Hartley, 35 km S Cooktown, Monteith & al., 23.iv.1982, [QM: 2♀; −15.79: 145.21]. QLD, 

Beating 12 m Scrub Site 20 NE Q, 15°50′S 145°19′E, 

RM VED, 22.xi.1975, [QM: 1♂; −15.83: 145.32]. QLD, 

Bellenden Ker Range, Cableway base Stn, 100 m, Earthwatch, 17.x.1981, [QM: 1♂, 1♀, 3 juveniles; −17.27: 

145.92]. QLD, Cairns, RG Wind, viii.–ix.1838, [MCZ: 

1♀; −16.92: 145.79]. QLD, Cape Tribulation NE QLD, 

Monteith, Yates & Thompson, 22ix–7x.1982, [QM: 

1♀; −16.09: 145.45]. QLD, Davies Creek, NC Coleman, 

11.i.1972, [QM: 1♀; −17: 145.57]. QLD, Edmonton, RE 

Mascord, 27.vii.1971, [AMS: KS 16834, 1♀; −17.02: 

145.74]. QLD, Edmonton, NC Coleman, 29.ix.1969, 

[AMS: KS 16868, 2♂, 1♀; −17.02: 145.74]. QLD, Home 

Rule NEQ, night collecting; in web holding net, TT&VED, 

31.x.1974, [QM: 1♀; −15.73: 145.29]. QLD, Homevale 

Riverine Rainforest, 1.iv.1975, [QM: 1♀; −21.72: 148.7]. 

QLD, Kellep Hut, Windsor Tb, ANZSES Exp, xii.1980, 

[QM: 1♂; −27.43: 153.04]. QLD, Kenilworth State Forest, R Raven, 14.ii.1972, [QM: 1♀; −26.59: 152.72]. 

QLD, Koonawarra Cmpgrd. Kianagarow Track Bunya 

Mts. NP, EI Schlinger, 29.i.1978, [EMUC: 1♀; −26.79: 

153.08]. QLD, Michaelsens Farm Mt Cotton, vine thicket, 

V Gallon, 20.xi.1983, [QM: 1♀; −27.62: 153.24]. QLD, 

Mt Cook, site 34, RR&VED, 12.xi.1975, [QM: 1♂, 1♀; 

−15.49: 145.26]. QLD, Mt Finlay, site 29, RM JC VED, 

2.xii.1973, [QM: 1 juvenile; −15.82: 145.35]. QLD, 

OF Kroombit Tops SEQ, R Raven, 25.ii.1982, [QM: 

2♀; −24.42: 150.97]. QLD, Palm Cove, NC Coleman, 

10.xi.1971, [QM: 1♀; −16.75: 145.63]. QLD, The Hummock Mt Hedley, on shrub on path, VE Davies, [QM: 1♀; 
2 2 • smithsonian contributions to zoology

−15.75: 145.3]. QLD, Somerset, LM D’Albertis, 1875, 

[MCSNG: 1♀; −10.76: 142.56].

Menneus nemesio, new species

FIGURES 13, 16d–f, 27

Lehtinen, 1967:217, fig. 41

Types. Male holotype (CAS) and two female 

paratypes (AMS); see below.

Etymology. Noun in apposition, named for 

Alexander Nemesio Coddington.

Diagnosis. Menneus nemesio is unique in 

having a prolonged abdominal caudal extension beyond 

the spinnerets (Figure 27c–d,j–k). Females have copulatory ducts with four full spirals (Figure 27g). The median 

hematodocha in ventral view (left palp) spans 5–7 OC, the 

embolus tip lies at about 11 OC, and the median apophysis–basal lobe axis lies at about 1 OC (Figure 16e). The 

embolus makes 3.5 coils and is bent away from the tegulum in prolateral view (Figure 16f).

Description. Female from Wamberal. Total 

length 10.9 mm. Carapace 3.2 mm long; 2.2 mm wide. 

Abdomen 7.4 mm long; 2.8 mm wide. Cephalic width 0.6× 

carapace width. PME diameter 0.2 mm; PME separation 

1.3 diameters. AME/PME 0.7; AME separation 1.6 AME 

diameters; equidistant between PME and clypeal margin. 

ALE/PME 0.9; ALE tubercle large, eye’s lower edge 2/3 

or more diameters from clypeus margin; ALE 0.3 diameters below clypeal margin. PLE/PME 0.9; PLE separation 

4.1 PLE diameters; PLE 1.9 diameters behind PME. Clypeus 1.5 AME diameters. Chelicerae with 4 promarginal 

teeth; 4 retromarginal teeth; median frontal macrosetae 

proximal to laterals; laterals stout. Sternum 0.7× as wide 

as long; width at coxae I and II about equal. First femur 

length 7.6 mm; with 4 to 6 prolateral macrosetae; patella 

+ tibia I, 8.6 mm; metatarsus I, 8.6 mm; tarsus I, 2 mm. 

Lengths of patella + tibia II, 8 mm; III, 5.7 mm; IV, 5 mm. 

Calamistrum 0.3 metatarsus IV length.

Male (holotype). Total length 8.2 mm. Carapace 2.8 

mm long. Carapace 2 mm wide. Abdomen 5.5 mm long. 

Abdomen 1.2 mm wide. Cephalic width 0.5× carapace 

width. PME diameter 0.1 mm; PME separation 1.8 diameters. AME/PME 0.8; AME separation 0.6 AME diameters; closer to clypeal margin than to PME. ALE/PME 1; 

ALE tubercle large; eye’s ventral edge 2/3 or more diameters from clypeus margin; ALE 1 diameters below clypeal 

margin. PLE/PME 0.8; PLE separation 3.8 PLE diameters; 

PLE 2.4 diameters behind PME. Clypeus 1.1 AME diameters. Chelicerae with 3 promarginal teeth; 1 retromarginal 

teeth; median frontal macrosetae present, but laterals absent. Sternum 0.6× as wide as long. First femur length 8.8 

mm; with 3 prolateral macrosetae; patella + tibia I 10.3 

mm; metatarsus I 12.4 mm; tarsus I 2.9 mm. Lengths of 

patella + tibia II, 8.4 mm; III, 4.9 mm; IV, 5.2 mm. Tarsus 

I divided and of uniform width. Terminal portion of tibia 

II with stout condyles and prolateral apophysis. Terminal 

portion of metatarsus II with curve bordered posteriorly 

by stout seta. Dorsal surface of tarsus II with proximal 

stout setae. Palpal femur length 2.3× cymbium length.

Variation. Both females vary in the degree 

of pigmentation; only one male is known. Female body 

length ranges from 10.5 to 11.5 mm (n = 2).

Natural History. Unknown.

Distribution. New South Wales (Figure 13).

Notes. Lehtinen (1967:217, fig. 41) illustrated 

the epigynum of a female specimen in the British Museum 

(BMNH: 19.9.18.5733), which currently bears a label 

“Avella despiciens.” However, the holotype of M.despiciens, at Oxford, is juvenile and therefore a nomen 

dubium (see below). The BMNH female, illustrated by 

Lehtinen, is M. nemesio.

Specimens Examined. AUSTRALIA: NSW, 

5 mi N Hawkesbury R. br., 150 m, Ross & Cavagnaro, 

3.xii.1962, [CAS: 1♂; −33.53: 151.19]. NSW, Epping, 

Sydney, N Allan, 30.viii.1969, [AMS: KS 16866, 1♀; 

−33.77: 151.08]. NSW, Sydney, no collector or date, 

BMNH: 19.9.18.5733, 1♀; −33.86: 151.2]. NSW, Wamberal, MR Gray, 7.xii.1983, [AMS: KS 13622, 1♀; 

−33.42: 151.45].

Menneus neocaledonicus (Simon, 1889)

FIGURES 13, 16g–i, 28

Avella neo-caledonica Simon, 1889:242 (juvenile).

Menneus neocaledonicus: Simon, 1892:210.—Berland, 1924:174, figs.11–

15 (♂).

Avella neocaledonica: Platnick, 2010.

Types. Presumed syntypes (2 juveniles) from 

Noumea in MNHN, examined.

Diagnosis. Unlike other Menneus species, 

female M. neocaledonicus abdomen is more or less fusiform, lacks any prominent tubercles (Figure 28c,d), the 

copulatory ducts make a single full spiral (Figure 28g), 

and the male lacks short stout setae on metatarsus II. The 
number 636 • 2 3

median hematodocha in ventral view (left palp) spans 

9–12:30 OC, the embolus tip lies at about 11 OC, and 

the median apophysis–basal lobe axis lies at about 3 

OC (Figure 16h). The embolus makes 2 coils and is bent 

away from the tegulum in prolateral view (Figure 16i). 

The embolus takes only half the size of the tegulum in 

ventral view.

Description. Female from Aoupini. Total 

length 16 mm. Carapace 5.1 mm long; 2.9 mm wide. Abdomen 5.5 mm long; 2.9 mm wide. Cephalic width 1.6× 

carapace width. PME diameter 0.2 mm; PME separation 

2.4 diameters. AME/PME 0.9; AME separation 1.6 AME 

diameters; closer to PME than to clypeal margin. ALE/

PME 0.8; ALE tubercle small, eye’s lower edge contiguous with clypeus margin; ALE 0.6 diameters below clypeal 

margin. PLE/PME 0.8; PLE separation 6.3 PLE diameters; 

PLE 2.2 diameters behind PME. Clypeus 0.8 AME diameters. Chelicerae with 4 promarginal teeth; 6 retromarginal teeth; frontal macrosetae in transverse line; laterals 

attenuate. Sternum 0.5× as wide as long; maximum width 

at coxa II. First femur length 14.4 mm; with 4 prolateral 

macrosetae; patella + tibia I, 16 mm; metatarsus I, 15.7 

mm; tarsus I, 3.2 mm. Lengths of patella + tibia II, 12.4 

mm; III, 8.4 mm; IV, 9.2 mm. Calamistrum 0.3 metatarsus 

IV length.

Male from Aoupini. Total length 14 mm. Carapace 

4.5 mm long. Carapace 2.2 mm wide. Abdomen 9.1 mm 

long. Abdomen 1.7 mm wide. Cephalic width 1.2× carapace width. PME diameter 0.2 mm; PME separation 2.2 

diameters. AME/PME 1; AME separation 1.3 AME diameters; equidistant between PME and clypeal margin. 

ALE/PME 0.8; ALE tubercle small; eye’s ventral edge contiguous with clypeus margin; ALE 0.8 diameters below 

clypeal margin. PLE/PME 0.9; PLE separation 5.2 PLE 

diameters; PLE 1.8 diameters behind PME. Clypeus 0.1 

AME diameters. Chelicerae with 4 promarginal teeth; 5 

to 6 retromarginal teeth; frontal macrosetae in transverse 

line; laterals attenuate. Sternum 0.5× as wide as long. First 

femur length 16.4 mm; with 4 prolateral macrosetae; patella + tibia I 17.7 mm; metatarsus I 10.9 mm; tarsus I 

4.5 mm. Lengths of patella + tibia II, 12.9 mm; III, 8.4 

mm; IV, 9.4 mm. Tarsus I divided and of uniform width. 

Terminal portion of tibia II with stout condyles and prolateral apophysis. Terminal portion of metatarsus II with 

curve bordered posteriorly by stout seta. Dorsal surface of 

tarsus II without stout setae. Palpal femur length 3× cymbium length.

Variation. Female body length ranges from 

15.5 to 17.0 mm (n = 2); only one male is known.

Natural History. Unknown.

Distribution. New Caledonia (Figure 13).

Specimens Examined. NEW CALEDONIA: Aoupinie, 20 km NE Poya, 650 m, Monteith & Cook, 

18.v.1984, [QM: 1♂, 1♀; −21.15: 165.31]. Mandjelia, 

above Pouebo, 6–750, Monteith & Cook, 11.v.1984, 

[QM: 1♀; −20.4: 164.57]. Noumea, Saves, [MNHN: AR 

165, 2 juveniles; −22.23: 166.47].

Menneus quasimodo, new species

FIGURES 13, 29

Types. Holotype female, designated here, in 

poor condition in WAM from Norseman; details below.

Etymology. Named after Victor Hugo’s famous literary character, Quasimodo, the hunchback of 

Notre Dame. The holotype spider resembles Quasimodo 

in having a “wart” and a “hump” (Figure 29c). Gender 

is masculine.

Diagnosis. The single female has multiple median and frontal macrosetae. The epigynum of the only female is slightly damaged, so that the number of copulatory 

duct spirals is ambiguous—either three or four full spirals 

(or four or five counting lateral margins in dorsal view; 

Figure 29g). The paired, relatively anterior, abdominal tubercles are markedly large and truncate, and the female 

lacks the abdominal extension beyond the spinnerets that 

characterizes M. nemesio.

Description. Female (holotype). Total length 

8.8 mm. Carapace 3.3 mm long; 2.2 mm wide. Abdomen 5.3 mm long; 4.6 mm wide. Cephalic width 0.42× 

carapace width. PME diameter 0.21 mm; PME separation 0.27 diameters. AME/PME 0.67; AME separation 

1.64 AME diameters; closer to clypeal margin than PME. 

ALE/PME 0.67; ALE tubercle large, eye’s lower edge 1/3 

diameter from clypeus margin; ALE only slightly below 

clypeal margin. PLE/PME 0.62; PLE separation 5.69 PLE 

diameters; PLE 2.46 diameters behind PME. Clypeus 1.8 

AME diameters. Chelicerae with 2 promarginal teeth; 3 

retromarginal teeth; multiple macrosetae on median and 

frontal surfaces. Sternum 0.8× as wide as long; widest at 

coxae I. First femur length 7.1 mm; with 4 prolateral macrosetae; patella + tibia I, 7.5 mm; metatarsus I, 7.4 mm; 

tarsus I, 1.4 mm. Lengths of patella + tibia II, 7.4 mm; III, 

4.4 mm; Leg IV missing.

Variation. Unknown.

Natural History. Unknown.

Distribution. Western Australia (Figure 13).
2 4 • smithsonian contributions to zoology

Specimens Examined. AUSTRALIA: WA, 

Norseman, 32°12′S 121°46′E, “senex,” 1926, [WAM: 

26/465, 1♀; −32.2: 121.77].

Menneus trinodosus Rainbow, 1920

FIGURES 13, 17d–f, 31

Menneus trinodosus Rainbow, 1920:230, pl. 28, figs. 3, 4 (♀).

Deinopis insularis Rainbow, 1920:229, pl. 28, figs. 1, 2 (♂). NEW SYNONYMY.

Avella trinodosa: Lehtinen, 1967:217 (transfer from Menneus).—Platnick, 

2010.

Avella insularis: Lehtinen, 1967:217, fig. 38 (transfer from Deinopis).—Coddington, 1990a:11, fig. 44 (♂).—Platnick, 2010.

Types. Holotype female of M. trinodosus in 

South Australian Museum, Adelaide, examined (Australia, NSW, Lord Howe Island: see Entomology Insecta Register I11509, no latitude given, no longitude given, A. M. 

Lea, 12/1915–1/1916, SAM: N1981233 and ARA 5325). 

Holotype male of D. insularis in South Australian Museum, examined (Australia, NSW, Lord Howe Island: see 

Entomology Insecta Register I11508, A. M. Lea, 12/1915–

1/1916, SAM: N1981232 and ARA 5325). 

Diagnosis. Uniquely in Menneus, female 

M. trinodosus have posterior paired abdominal tubercles, 

the anterior portion of the epigynum is blunt; in ventral 

view the spermathecae are large, and the copulatory ducts 

make one full spiral (or two if counting lateral margins; 

Figure 31c,d,g). The median hematodocha in ventral view 

(left palp) spans 1–6 OC, the embolus tip lies at about 

11 OC, and the median apophysis–basal lobe axis lies 

at about 2:30 OC (Figure 17e). The embolus makes 2.5 

coils and is bent away from the tegulum in prolateral view 

(Figure 17f).

Description. Female from Gladstone. Total 

length 11.7 mm. Carapace 4.1 mm long; 2.7 mm wide. 

Abdomen 7 mm long; 3.1 mm wide. Cephalic width 1.5× 

carapace width. PME diameter 0.2 mm; PME separation 

3 diameters. AME/PME 0.9; AME separation 1.9 AME 

diameters; equidistant between PME and clypeal margin. 

ALE/PME 0.7; ALE tubercle small, eye’s lower edge contiguous with clypeus margin; ALE 0.6 diameters below 

clypeal margin. PLE/PME 1; PLE separation 6.5 PLE 

diameters; PLE 1.7 diameters behind PME. Clypeus 0.5 

AME diameters. Chelicerae with 4 promarginal teeth; 5 

to 6 retromarginal teeth; frontal macrosetae in transverse 

line; laterals attenuate. Sternum 0.6× as wide as long; 

maximum width at coxa II. First femur length 8.1 mm; 

with 8 to 9 prolateral macrosetae; patella + tibia I, 8.1 

mm; metatarsus I, 8.7 mm; tarsus I, 2.1 mm. Lengths of 

patella + tibia II, 8.1 mm; III, 3.6 mm; IV, 4.2 mm. Calamistrum 0.3 metatarsus IV length.

Male from Burdaberg. Total length 9.1 mm. Carapace 

3.3 mm long. Carapace 2 mm wide. Abdomen 5.6 mm 

long. Abdomen 1.8 mm wide. Cephalic width 0.6× carapace width. PME diameter 0.2 mm; PME separation 2.5 

diameters. AME/PME 1.1; AME separation 1.2 AME diameters; closer to PME than to clypeal margin. ALE/PME 

0.7; ALE tubercle small; eye’s ventral edge contiguous with 

clypeus margin; ALE 0.6 diameters below clypeal margin. PLE/PME 1; PLE separation 4.9 PLE diameters; PLE 

1.5 diameters behind PME. Clypeus 0.1 AME diameters. 

Chelicerae with 4 promarginal teeth; 3 to 4 retromarginal 

teeth; frontal macrosetae in transverse line; laterals attenuate. Sternum 0.6× as wide as long. First femur length 9.6 

mm; with 6 to 7 prolateral macrosetae; patella + tibia I 

8.6 mm; metatarsus I 8.7 mm; tarsus I 2.1 mm. Lengths 

of patella + tibia II, 11.4 mm; III, 3.2 mm; IV, 3.7 mm. 

Tarsus I undivided. Terminal portion of tibia II with stout 

condyles. Terminal portion of metatarsus II without distal 

curve. Dorsal surface of tarsus II without stout setae. Palpal femur length 2.2× cymbium length.

Variation. While in most females the abdominal tubercles are posterior as illustrated, in some this is 

less pronounced. Female body length ranges from 9.1 to 

18.0 mm (n = 9), and males from 7.0 to 9.1 (n = 10).

Natural History. Unknown.

Distribution. Australia (Queensland, New 

South Wales, including Lord Howe Island) (Figure 13).

Specimens Examined. AUSTRALIA: NSW, 

L. H. I. 25 [Lord Howe I.], in grass-litter, [AMS: KS 16843, 

2♀; −31.55: 159.08]. NSW, L. H. I. 39 [Lord Howe I.], 

[AMS: KS 16846, 1♀; −31.55: 159.08]. NSW, Lord Howe 

Island, Intermediate Hill, Big Creek, malaise-trough, tall 

forest #5, 50 ft, S & J Peck, 18.v.1980, [AMNH: 1♂; 

−31.55: 159.08]. NSW, Minnamurra Falls Res., C Horseman, 20.iv.1982, [AMS: KS 9204, 1♀, 1 juvenile; −34.62: 

150.85]. QLD, Bulburin (Forestry nursery), NW Bundaberg, rainforest site 3A, 580 m, 24°31′S 151°29′E, Gray & 

Horseman, iii.1975, [AMS: KS 0092, 4♂; −24.52: 151.48]. 

QLD, Bulburin SF, 24°31′S 151°29′E, [QM: 2♂, 2♀; 

−24.52: 151.48]. QLD, Bulburin SF, 24°31′S 151°29′E, 

[QM: 1♂; −24.52: 151.48]. QLD, Eurimbula, SE Gladstone, rainforest site 4, 0 m, 24°11′S 151°50′E, C Horseman, iii.1975, [AMS: KS 0256, 1♀; −24.18: 151.83]. 

QLD, Finch Hatton, RK&VED, 7.iv.1975, [QM: 1♂, 1♀, 

2 juveniles; −21.14: 148.63]. QLD, Kroombit Tops (Three 

Moon Ck.), 45 km SSW Calliope, rainforest, Davies & 
number 636 • 2 5

Gallon, 9.xii.1983, [QM: 1♂, 1 juvenile; −24.49: 151.2]. 

QLD, Kuranda (12 mi N of), Coleman & Mascord, 

30.i.1972, [AMS: KS 16867 1♀; −16.82: 145.63]. QLD, 

Malaan S.F., RR&VED, 20.iv.1978, [QM: 1♂, 1 juvenile; 

−17.58: 145.58]. QLD, Mt. Goonaneman, nr Childers, 

670 m, Davies & Raven, 3.xi.1980, [QM: 1♀; −25.12: 

152.18]. QLD, Spear Ck nr Mt Mallay, CNVF Site 34, 

RR&VED, 5.xi.1975, [QM: CNVF Site 34?, 1♂, 1♀, 7 

juveniles; −16.68: 145.33].

Menneus wa, new species

FIGURES 13, 17g–i, 32

Types. Holotype male from Denmark and three 

paratype females from Quaalup Homestead and Quaranup Road, all in WAM; details below.

Etymology. Named for Western Australia, 

construed as a noun in apposition.

Diagnosis. Among Australian Menneus with 

paired median dorsal abdominal tubercles female M. wa

closely resemble M. superciliosus in having copulatory ducts 

with two full spirals (or three counting lateral margins in 

dorsal view; Figure 32g). Menneus wa seem to have broader, 

shorter, abdomens with a lateral brown abdominal spot 

(Figure 32c,d). The median hematodocha in ventral view 

(left palp) spans 6–9 OC, the embolus tip lies at about 11 

OC, and the median apophysis–basal lobe axis lies at about 

1:30 OC (Figure 17h). The embolus makes 3.5 coils and is 

bent away from the tegulum in prolateral view (Figure 17i).

Description. Female from Quaranup Rd. 

Total length 10.8 mm. Carapace 3.3 mm long; 2.2 mm 

wide. Abdomen 7.6 mm long; 5.6 mm wide. Cephalic 

width 0.3× carapace width. PME diameter 0.23 mm; PME 

separation 0.28 diameters. AME/PME 0.7; AME separation 1.6 AME diameters; closer to PME than clypeal 

margin. ALE/PME 0.7; ALE tubercle large, eye’s lower 

edge 1/3 diameter from clypeus margin; ALE only slightly 

below clypeal margin. PLE/PME 1.0; PLE separation 3.7 

PLE diameters; PLE 1.6 diameters behind PME. Clypeus 

1.8 AME diameters. Chelicerae with 4 promarginal teeth; 

4 retromarginal teeth; median frontal macrosetae proximal to laterals; laterals stout. Sternum 0.6× as wide as 

long; widest at coxae II. First femur length 7.1 mm; with 

5 prolateral macrosetae; patella + tibia I, 7.8 mm; metatarsus I, 7.6 mm; tarsus I, 1.5 mm. Lengths of patella + 

tibia II, 7.2 mm; III, 4.4 mm; IV, 4.6 mm. Calamistrum 

0.35 metatarsus IV length.

Male (holotype). Total length 8.5 mm. Carapace 2.8 

mm long. Carapace 2 mm wide. Abdomen 5.8 mm long. 

Abdomen 1.9 mm wide. Cephalic width 0.4× carapace 

width. PME diameter 0.23 mm; PME separation 1.1 diameters. AME/PME 0.57; AME separation 1.6 AME diameters; closer to clypeal margin than to PME. ALE/PME 

0.83; ALE tubercle large; eye’s ventral edge 1.2 or more 

diameters from clypeus margin; ALE 0.1 diameters below 

clypeal margin. PLE/PME 1.0; PLE separation 3.7 PLE diameters; PLE 1.15 diameters behind PME. Clypeus 1.54 

AME diameters. Chelicerae with 4 promarginal teeth; 5 

retromarginal teeth; median frontal and lateral macrosetae missing (or absent?). Sternum 0.6× as wide as long. 

First femur length 9.8 mm; with at least 3 prolateral macrosetae; patella + tibia I 11.3 mm; metatarsus I 12.7 mm; 

tarsus I missing. Lengths of patella + tibia II, 8.8 mm; III, 

5.4 mm; IV, 5.7 mm. Terminal portion of tibia II with 

stout condyles and prolateral apophysis. Terminal portion 

of metatarsus II with curve bordered posteriorly by stout 

seta. Dorsal surface of tarsus II with proximal stout setae. 

Palpal femur length 2.3× cymbium length.

Variation. Three females range in length from 

11.0 to 12.5 mm. Two of three females had conspicuous 

brown spots on the lateral abdominal surface. Only one 

male is known.

Natural History. Unknown.

Distribution. Western Australia (Fig. 13)

Specimens Examined. AUSTRALIA: WA, 

Denmark, inside house, 34°58′S 117°21′E, R. P. McMillan, 2iii1994, [WAM: 97/2350, 1♂; −34.97: 117.35]. WA, 

Quaalup Homestead, night collecting, 34°17′S 119°26′E, 

MS Harvey & JM Waldock, 25.v.1994, [WAM: 97/2351, 

1♀; −35.08: 117.92]. WA, Torndirrup Nat. Pk., Quaranup Rd., 35°05′S 117°55′E, MS Harvey & JM Waldock, 

25.iv.1990, [WAM: 97/2352, 1♀; −34.28: 119.43]. WA, 

Torndirrup Nat. Pk., Quaranup Rd., 35°05′S 117°55′E, 

MS Harvey & JM Waldock, 25.iv.1990, [WAM: 97/2353, 

1♀; −33.97: 18.45].

Menneus pietrzeniukae Wunderlich, 2004

?Menneus pietrzeniukae Wunderlich 2004:892 (juv.), figs. 1–6.

Types. Juvenile female from Tertiary Baltic amber 

forest, Paleontological Museum of the Humboldt University 

Berlin, MB.A114, collected by Simon. Not examined.

Notes. Judging from Wunderlich (2004, fig. 

6), the specimen is juvenile, which otherwise would make 

the name a nomen dubium. However, given that it is a 

Baltic Amber fossil and that other fully diagnosable specimens may be discovered in the future, we leave it to future 

authors to decide formally the status of this name.
2 6 • smithsonian contributions to zoology

Nomina dubia

Avella angulata L. Koch, 1878

Avella angulata L. Koch, 1878:1037, pl. 91, fig. 1 (♀).—Platnick, 2010.

Menneus angulatus: Rainbow, 1911:123.—Davies, 1988:282, fig. 7 (♂).

Types. Female holotype in Hamburg, examined.

Notes. The specimen consists only of the cephalothorax and legs. The abdomen is missing. It is unclear 

if the animal was adult. Without the epigynum, or even 

knowledge of the form of abdominal tubercles, no known 

Menneus species can be unambiguously linked to this 

name.

Avella despiciens O. P.-Cambridge, 1877

Avella despiciens O. P.-Cambridge, 1877:574, pl. 57, fig. 10 (♀) [see Cambridge, O. P., in References].—Lehtinen, 1967:217, fig. 41 (misidentification).—Platnick, 2010.

Menneus despiciens: Rainbow, 1911:124.

Types. Juvenile female (?) holotype in Oxford, 

examined (Australia, QLD, Rockhampton, E. W. Janson, 

1877, OUMNH: Bottle no. 169, 1 juvenile). See Note 

under M. nemesio.

Avella unifasciata L. Koch, 1878

Avella unifasciata L. Koch, 1878:1035, pl. 90, fig. 6 (♂).—Platnick, 2010. 

Menneus unifasciatus: Rainbow, 1911:124.

Types. The holotype male cannot be located 

and is presumed lost. 
number 636 • 2 7

FIGURES

FIGURE 1. Menneus natural history: a–c are Menneus capensis, Cape Town; (a) female holding web; (b) subadult female dorsal; (c) male; 

(d) Menneus camelus, female from Sodwana Bay, KwaZulu-Natal, holding web (note unpaired abdominal tubercle).
2 8 • smithsonian contributions to zoology
number 636 • 2 9

FIGURE 2. (facing page) Deinopid natural history: (a) Menneus camelus female from Gwala-Gwala, KwaZulu-Natal, in cryptic posture on 

twig, head marked with black arrow, note unusual morphology with paired abdominal tubercles (white arrows); (b, c) presumed female Menneus camelus in its web, voucher from Fanies Island, KwaZulu-Natal, not available for examination; (d) web of Deinopis spinosa, redrawn from 

Coddington and Sobrevila (1987); (e) presumed female Menneus camelus spinning her eggsac, voucher from Sodwana Bay, KwaZulu-Natal, 

not available for examination.

FIGURE 3. Deinopid somatic morphology: a, b are Deinopis spinosa, female from Florida, (a) frontal view 

of prosoma and (b) dorsal view of first left femur; c, e are Menneus aussie, male from Queensland: (c) first 

left tarsus, (d) distal second left tibia, and (e) second left metatarsus-tarsus joint. Numbers = characters 

(states); see text.
3 0 • smithsonian contributions to zoology

FIGURE 4. Deinopid male palpal morphology: a–d are Deinopis spinosa, expanded left palp in (a) prolateral, (b) retrolateral, 

(c) ventral, and (d) apical view; (e) Menneus camelus from South Africa, apical view.
number 636 • 3 1

FIGURE 5. Deinopid female morphology (D. spinosa): (a) sternum, ventral view, showing setose knob (arrow); (b) labium, endites, trichobothria 

(arrows); (c) detail of chelicera, ventral view, showing teeth (arrows) and denticles; (d) detail of carapace showing feathery, blunt, and plumose 

setae; (e) metatarsus IV showing calamistrum; (f) tarsus IV claws showing reduced retrolateral teeth (arrow) and ventral tarsal deinopoid setal 

comb (arrows). All scales = 0.1 mm, except a = 1 mm.
3 2 • smithsonian contributions to zoology

FIGURE 6. Deinopid male morphology (D. spinosa): (a) epiandrous spigots, ventral view; (b) detail of epiandrous spigots, ventral view; (c) anal 

tubercle, ventral view; (d) left palp, prolateral view; (e) median apophysis, ventral view; (f) detail of embolus tip. All scales = 0.1 mm, except b 

and f = 0.01 mm.
number 636 • 3 3

FIGURE 7. Deinopis spinosa female spinneret morphology, all ventral views: (a) spinneret overview; (b) cribellum; (c) cribellar spigot detail; 

(d) ALS with MAP and PI spigots; (e) PMS with mAP, AC, PC, and CY spigots; (f) PLS with CY, AC, and MS spigots. All scales = 0.l mm, 

except c = 0.01 mm.
3 4 

• smithsonian contributions to zoology

FIGURE 8. (a) Cladogram for outgroups, Deinopis exemplars, and all Menneus species. Length 131, ci 0.54, ri 0.69. All 131 steps are mapped on the tree, open ticks are characters with homoplasy, solid without. Ambiguous optimizations (5, 7–9, 12, 14, 18–19, 28–29, 34–35, 38–39, 41, 44–45, 47–48, 50, 55), were resolved to maximize homology of states and minimize parallel origin. (b) Growth of deinopid museum collections from 1800 to the present, by decade.
number 636 • 3 5

FIGURE 9. Deinopis lamia, from Pico Turquino, Cuba: a–d are female; (a) habitus, dorsal; (b) epigynum, ventral; (c) epigynum, caudal; (d) epigynum, dorsal; e–g are male; (e) habitus, dorsal; (f) left palpal bulb, ventral; (g) embolus tip.
3 6 • smithsonian contributions to zoology

FIGURE 10. Deinopis cylindrica: a–e are female from Lake Sibayi, KwaZulu-Natal; (a) prosoma, frontal; (b) prosoma, dorsal; (c) abdomen, 

ventral; (d) abdomen, dorsal; (e) epigynum, ventral; f–j are male from Empangeni, KwaZulu-Natal; (f) prosoma, frontal; (g) prosoma, dorsal; 

(h) abdomen, ventral; (i) abdomen, dorsal; (j) left palpal bulb, ventral; (k) embolus tip.
number 636 • 3 7

FIGURE 11. Deinopis spinosa: a–e are female from Florida; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, ventral; (d) abdomen, dorsal; (e) epigynum, ventral; f–h are epigynum of female from Jamaica; (f) ventral; (g) caudal; (h) dorsal; i–m are male from Florida; (i) prosoma, 

frontal; (j) prosoma, dorsal; (k) abdomen, ventral; (l) abdomen, dorsal; (m) left palpal bulb, ventral.
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FIGURE 12. Distribution of African Menneus species.
number 636 • 3 9 FIGURE 13. Distribution of Australasian Menneus species.
4 0 • smithsonian contributions to zoology

FIGURE 14. Menneus male left palps in (left) prolateral, (middle) ventral, and (right) retrolateral view: (a–c) 

M. camelus, (d–f) M. capensis, and (g–i) M. darwini. Diagnostic features of species in b, e, h: short bold arrows 

indicate extent of median hematodocha; long slender arrow indicates embolus tip; dashed line indicates the axis 

of the median apophysis and basal lobe.
number 636 • 4 1

FIGURE 15. Menneus male left palps in (left) prolateral, (middle) ventral, and (right) retrolateral view: (a–c) 

M. dromedarius, (d–f) M. samperi, and (g–i) M. tetragnathoides. Diagnostic features of species in b, e, h: short 

bold arrows indicate extent of median hematodocha; long slender arrow indicates embolus tip; dashed line indicates the axis of the median apophysis and basal lobe.
4 2 • smithsonian contributions to zoology

FIGURE 16. Menneus male left palps in (left) prolateral, (middle) ventral, and (right) retrolateral view: (a–c) 

M. aussie, (d–f) M. nemesio, and (g–i) M. neocaledonicus. Diagnostic features of species in b, e, h: short bold arrows indicate extent of median hematodocha; long slender arrow indicates embolus tip; dashed line indicates the 

axis of the median apophysis and basal lobe.
number 636 • 4 3

FIGURE 17. Menneus male left palps in (left) prolateral, (middle) ventral, and (right) retrolateral view: (a–c)

M. superciliosus, (d–f) M. trinodosus, and (g–i) M. wa. Diagnostic features of species in b, e, h: short bold arrows 

indicate extent of median hematodocha; long slender arrow indicates embolus tip; dashed line indicates the axis 

of the median apophysis and basal lobe.
4 4 • smithsonian contributions to zoology

FIGURE 18. Menneus camelus: a–f are female from Dukuduku, KwaZulu-Natal; (a) prosoma, frontal; (b) prosoma, dorsal; (c) abdomen, 

ventral; (d) abdomen, dorsal; (e) epigynum, caudal; (f) epigynum, ventral; g–k are female from Pietermaritzburg, KwaZulu-Natal; (g) prosoma, 

frontal; (h) prosoma, dorsal; (i) abdomen, ventral; (j) abdomen, dorsal; (k) epigynum, dorsal.
number 636 • 4 5

FIGURE 19. Menneus camelus, male from Amanzimtoti, KwaZulu-Natal: (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, 

ventral; (d) abdomen, dorsal; (e) left palpal bulb, ventral; (f) embolus tip.
4 6 • smithsonian contributions to zoology

FIGURE 20. Menneus capensis: a–f are female from Newlands Forest, Cape Town; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, 

dorsal; (d) abdomen, ventral; (e) epigynum, ventral; (f) epigynum, dorsal; g–l are male from Goudini, Cape Province; (g) prosoma, dorsal; 

(h) prosoma, frontal; (i) abdomen, dorsal; (j) abdomen, ventral; (k) left palpal bulb, ventral; (l) embolus tip.
number 636 • 4 7

FIGURE 21. Menneus darwini, holotype male from Tanzania: (a) prosoma, 

dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, ventral.
4 8 • smithsonian contributions to zoology

FIGURE 22. Menneus dromedarius: a–d are female from Table Mountain, South Africa; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum of female from Kleinmond, South Africa; (e) ventral; (f) caudal; (g) dorsal; h–l are male 

from Grahamstown, South Africa; (h) prosoma, dorsal; (i) prosoma, frontal; (j) abdomen, dorsal; (k) abdomen, ventral; (l) left palpal bulb, 

ventral.
number 636 • 4 9

FIGURE 23. Menneus samperi: a–d are female from Tanzania; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, 

ventral; e–g are epigynum of female from Uganda; (e) ventral; (f) caudal; (g) dorsal; h–k are male from Kenya; (h) prosoma, dorsal; (i) prosoma, 

frontal; (j) abdomen, dorsal; (k) abdomen, ventral.
5 0 • smithsonian contributions to zoology

FIGURE 24. Menneus tetragnathoides, male from Chintheche, Malawi: (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, 

ventral; (d) abdomen, dorsal; (e) left palpal bulb, ventral; (f) embolus tip.
number 636 • 5 1

FIGURE 25. Menneus aussie: a–g are female from Manly, NSW; (a) prosoma, dorsal; (b) abdomen, dorsal; (c) abdomen, ventral; (d) prosoma, 

frontal; (e) epigynum, caudal; (f) epigynum, ventral; (g) epigynum, dorsal; h–l are male from Rundle Range, Queensland; (h) prosoma, dorsal; 

(i) prosoma, frontal; (j) abdomen, ventral; (k) abdomen, dorsal; (l) left palpal bulb, ventral.
5 2 • smithsonian contributions to zoology

FIGURE 26. Menneus bituberculatus, female from Upper Brookfield, Queensland: (a) prosoma, dorsal; (b) prosoma, frontal; 

(c) abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum; (e) ventral; (f) caudal; (g) dorsal.
number 636 • 5 3

FIGURE 27. Menneus nemesio: a–g are female from Wamberal, New South Wales (NSW); (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum; (e) ventral; (f) caudal; (g) dorsal; h–m are male from Hawksburg, NSW; (h) prosoma, 

frontal; (i) prosoma, dorsal; (j) abdomen, dorsal; (k) abdomen, ventral; (l) left palpal bulb, ventral; (m) embolus tip.
5 4 • smithsonian contributions to zoology

FIGURE 28. Menneus neocaledonicus from Aoupinie, New Caledonia: a–g are female; (a) prosoma, dorsal; (b) prosoma, frontal; (c) 

abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum; (e) ventral; (f) caudal; (g) dorsal; h–m are male; (h) prosoma, dorsal; (i) prosoma, frontal; (j) abdomen, ventral; (k) abdomen, dorsal; (l) left palpal bulb, ventral; (m) embolus tip.
number 636 • 5 5

FIGURE 29. Menneus quasimodo, female from Western Australia: (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum; (e) ventral; (f) caudal; (g) dorsal.
5 6 • smithsonian contributions to zoology

FIGURE 30. Menneus superciliosus from Edmonton, Queensland: a–g are female; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, 

dorsal; (d) abdomen, ventral; (e) epigynum, caudal; (f) epigynum, ventral; (g) epigynum, dorsal; h–l are male; (h) prosoma, dorsal; (i) prosoma, frontal; (j) abdomen, dorsal; (k) abdomen, ventral; (l) left palpal bulb, ventral.
number 636 • 5 7

FIGURE 31. Menneus trinodosus, a–g are female from Eurimbula, Queensland; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum; (e), ventral; (f) caudal; (g) dorsal; h–m are male from Bundaberg, Queensland; 

(h) prosoma, dorsal; (i) prosoma, frontal; (j) abdomen, dorsal; (k) abdomen, ventral; (l) left palpal bulb, ventral; (m) embolus tip.
5 8 • smithsonian contributions to zoology

FIGURE 32. Menneus wa, from Western Australia: a–g are female; (a) prosoma, dorsal; (b) prosoma, frontal; (c) abdomen, dorsal; (d) abdomen, ventral; e–g are epigynum; (e) ventral; (f) caudal; (g) dorsal; h–k are male; (h) prosoma, dorsal; (i) prosoma, frontal; (j) abdomen, dorsal; 

(k) abdomen, ventral.
References

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