the genera of the spider family theridiosomatidae · 2013. 12. 3. · abstract coddington, jonathan...

The Genera of the SpiderFamily Theridiosomatidae

JONATHAN A. CODDINGTON

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 422

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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 4 2 2

The Genera of the Spider FamilyTheridiosomatidae

Jonathan A. Coddington

SMITHSONIAN INSTITUTION PRESS

City of Washington

1986

A B S T R A C T

Coddington, Jonathan A. The Genera of the Spider Family Theridioso-matidae. Smithsonian Contributions to Zoology, number 422, 96 pages, 220figures, 8 maps, 1 table, 1986.—The cosmotropical spider family Theridio-somatidae is revised at the generic level to contain 9 genera: Theridiosoma O.Pickard-Cambridge, 1879, Ogulnius O. Pickard-Cambridge, 1882, Wendil-garda Keyserling, 1886, Epeirotypus O. Pickard-Cambridge, 1894, Baalzebub,new genus (type-species B. baubo, new species), Epilineutes, new genus (type-species Theridiosoma globosum O. Pickard-Cambridge), Plato, new genus (type-species P. troglodita, new species), Naatlo, new genus (type-species N. sutila,new species), and Chthonos, new name. Of the 22 genera historically associatedwith the family, 17 have been rejected, transferred, or synonymized. Theri-dilella Chamberlin and Ivie, 1936 (damaged specimen), and Allototua Bryant,1945 (unique specimen lost), are considered unrecognizable nomina dubia;Haliger Mello-Leitao lacks the defining features of theridiosomatids and isconsidered incertae sedis. Diotherisoma di Caporiacco, 1947, is transferred tothe Araneidae and Totua Keyserling, 1891, to the Linyphiidae. The previoustransfers of Colphepeira Archer, 1941, to the Araneidae, Billima Simon, 1908,Helvidia Thorell, 1890, and Spheropistha Yaginuma, 1957, to the Theridiidae,Cyatholipulus Petrunkevitch, 1930, to the Symphytognathidae, CyatholipusSimon, 1894, and Tekella Urquhart, 1894, to the Cyatholipinae (Tetragnath-idae), and Parogulnius Archer, 1953, and Phricotelus Simon, 1895, to theMysmenidae are not contested. The genus Andasta Simon, 1895, is synony-mized with Theridiosoma, and Enthorodera Simon, 1907, and Cyathidea Simon,1907, with Wendilgarda. Theridiosoma argentatum Keyserling, 1886, and T.radiosum (McCook, 1881) are synonymized with T. gemmosum (L. Koch,1878), and Wendilgarda panamica Archer, 1953, W. hassleri Archer, 1953,and W. theridionina Simon, 1895, with W. clara Keyserling, 1886. Tecmessatetrabuna Archer, 1958, and Epeirotypus gloriae Petrunkevitch, 1930, aretransferred to Ogulnius. Maymena bruneti Gertsch, 1960, and Wendilgardaguacharo Brignoli, 1972, W. miranda Brignoli, 1972, and W. bicolor Keyser-ling, 1886, are transferred to Plato. Theridiosoma fauna Simon, 1897, T.splendidum (Taczanowski, 1873), and T. sylvicola Hingston, 1932, are trans-ferred to Naatlo. Theridiosoma albinotatum Petrunkevitch, 1930, and T. brauniWunderlich, 1976, are transferred to Baalzebub. Theridiosoma nigrum (Key-serling, 1886) is returned to Wendilgarda. The genus Tecmessa O. Pickard-Cambridge, 1882, is valid but the name is preoccupied {Tecmessa Burmeister,1878: Lepidoptera); the new name Chthonos replaces it. The new generaBaalzebub, Epilineutes, Naatlo, and Plato, and the new species Baalzebub baubo,Plato troglodita, Naatlo sutila, and Epeirotypus chavarria are described. Thesister taxon of the Theridiosomatidae is the Mysmenidae-Symphytognathi-dae-Anapidae clade. A cladogram for theridiosomatid genera is presented.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and isrecorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coralMontastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication DataCoddington, Jonathan A.The genera of the spider family Theridiosomatidae.(Smithsonian contributions to zoology ; no. 422)Bibliography: p.Supt. of Docs, no.: SI 1.27:422I. Theridiosomatidae—Classification. I. Title. II SeriesQL1.S54 no. 422 [QL458.42.T55] 591s [595.4'4] 85-600150

Contents

Page

Introduction 1Acknowledgments 2Methods 3

Abbreviations 4Taxonomic History 4

Morphology and Phylogeny 8Monophyly of Theridiosomatidae 8Comparative Morphology of Theridiosomatid Genitalia 10Interfamilial Relationships 13Intergeneric Relationships 17

THERIDIOSOMATIDAE 22

Key to Genera 27PLATONINAE, new subfamily 28

Plato, new genus 28Plato troglodita, new species 29Plato bruneti (Gertsch), new combination 31Plato miranda (Brignoli), new combination 33Plato guacharo (Brignoli), new combination 33Plato bicolor (Keyserling), new combination 33

Chthonos, new name 33Chthonos pectorosa (O. Pickard-Cambridge), new combination . 35Chthonos peruana (Keyserling), new combination 37Chthonos tuberosa (Keyserling), new combination 37Chthonos quinquemucronata (Simon), new combination 37

EPEIROTYPINAE Archer 37Epeirotypus O. Pickard-Cambridge 37

Epeirotypus brevipes O. Pickard-Cambridge 39Epeirotypus chavarria, new species 43

Naatlo, new genus 44Naatlo sutila, new species 45Naatlo splendida (Taczanowski), new combination 47Naatlo fauna (Simon), new combination 50Naatlo sylvicola (Hingston), new combination 52

OGULNIINAE, new subfamily 52

Ogulnius O. Pickard-Cambridge 52Ogulnius obtectus O. Pickard-Cambridge 55Ogulnius gloriae (Petrunkevitch), new combination 57Ogulnius tetrabuna (Archer), new combination 61

in

IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

THERIDIOSOMATINAE Simon 61Theridiosoma O. Pickard-Cambridge 61

Theridiosoma gemmosum (L. Koch) 64Theridiosoma semiargenteum (Simon), new combination 71Theridiosoma genevensium (Brignoli), new combination 71

Baalzebub, new genus 71Baalzebub baubo, new species 72Baalzebub albinotatus (Petrunkevitch), new combination 74Baalzebub brauni (Wunderlich), new combination 74

Epilineutes, new genus 74Epilineutes globosus (O. Pickard-Cambridge), new combination . 79

Wendilgarda Keyserling 82Wendilgarda mexicana Keyserling 83Wendilgarda clara Keyserling 88Wendilgarda atricolor (Simon), new combination 89

References 92

The Genera of the Spider FamilyTheridiosomatidae

Jonathan A. Coddington

Introduction

The spider family Theridiosomatidae exem-plifies a common taxonomic problem: a vaguelydefined, little-known, poorly understood, sup-posedly small, and yet exotic cosmotropicalgroup of animals. The exasperatingly small sizeof the spiders (often less than 2 mm total length)invited superficial descriptive work by taxono-mists and ensured neglect of their natural his-tory. However, the recent series of papers byForsterand Platnick (1977), Platnickand Shadab(1978a,b, 1979), Eberhard (1981, 1982), andCoddington (in press) on related araneoid spidersdemonstrates that knowledge of these smallgroups will probably be critical for our under-standing of the superfamily Araneoidea.

Study of theridiosomatids is important for anumber of reasons. Refutation of their tradi-tional placement within or near Araneidae (sensulato) makes the latter group more homogeneous,thus facilitating the eventual recognition of mon-ophyletic groups within that ill-defined assem-blage. The behavior and morphology of theridio-somatids will also help to advance our under-standing of the superfamily Araneoidea. Under-standing of that superfamily has always beenbased primarily on character polarities inferredfrom a few very large taxa (Araneidae, Linyphi-

Jonathan A. Coddington, Department of Entomology, NationalMuseum of Natural History, Smithsonian Institution, Washing-ton, DC 20560.

idae, Theridiidae). As groups of tropical ara-neoids become better known, they become reli-able outgroups for the remaining Araneoidea.Delimitation of these taxa thus can only improveour understanding of character transformationsin the superfamily as a whole.

Theridiosomatidae promises to be a muchlarger family than catalogs suggest (e.g., Roewer,1942; Bonnet, 1955-1959; Brignoli, 1983). Atpresent roughly 120 species, described and un-described, are known world-wide, and certainlythat is only the beginning. Probably rather fewof the 60-odd available species names will turnout to be synonyms. Most species are known onlyfrom the type series.

This revision was originally envisaged as atreatment of the neotropical theridiosomatid spe-cies. Tropical Africa, Australia, Malaysia, andNew Guinea, however, are rich in theridioso-matid species. As it turned out, putative synapo-morphies inferred for the neotropical groupswere contradicted by those in the Old WorldTropics; distribution patterns of characters insome cases are neither simple nor obvious.Therefore, it was clearly unwise to diagnose anygenus in the Neotropics without simultaneoustreatment of the family on a world-wide basis.This result expanded the work to its presentscope and rendered the idea of an exhaustiverevision of theridiosomatids at the species levelimpractical. The results published herein are acompromise: the family is revised on a generic

1

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

basis, interfamilial and intergeneric relationshipsare reviewed, a key to genera is provided, buttreatment of each genus is synoptic rather thancomplete. For each genus I have described orredescribed one, two, or three species to illus-trate diversity, including the type-species, andhave discussed the placement of all species in thegenus. By no means were all species redescribed,however. That task will be taken up in a seriesof generic revisions.

Even so, several more new genera could havebeen described. In this initial work, however, arather conservative approach has been taken to-ward the description of new genera. For exam-ple, Theridiosoma gemmosum forms a monophy-letic group with T. epeiroides, and possibly alsowith T. goodnightorum and its relatives. Takentogether this group may be the sister taxon ofanother species group including at least T. savan-num, T. nechodomae, T. davisi, and T. benoiti, butmore research is necessary to confirm that con-clusion. The two groups together are monophy-letic, and the name Theridiosoma is applied tothat more inclusive taxon. A similar situationoccurs in Ogulnius, which ranks with Theridio-soma as one of the largest genera in the family(20 to 30 species each).

ACKNOWLEDGMENTS

I wish to thank Herbert W. Levi for unstintingguidance, myriad favors, and continual assistancethroughout this study. Opell (1979) said it suc-cinctly and elegantly: "His excellent advice wasalways available but never imposed." W.G. Eber-hard generously shared his knowledge of thebehavior and natural history of theridiosomatids,as well as teaching me a very great amount aboutethological field technique. Cecile Villars, JohnHunter, and Wayne Maddison solved many last-minute problems that seemed overwhelming atthe time. Fieldwork was financed by a Jesse SmithNoyes Predoctoral Fellowship, the Organizationfor Tropical Studies, and the Richmond, Bar-bour, Atkins, and Anderson Funds of HarvardUniversity. National Science Foundation Grant

DEB 80-20492 to H.W. Levi defrayed much ofthe cost of laboratory research. Mario Dary ofthe Universidad de San Carlos in Guatemalamade possible field research in Purulha. JoseOttenwalder helped immeasurably during field-work in the Dominican Republic, and James Wy-lie of the Endangered Species Office, U.S. Fishand Wildlife Service, provided accommodationsand aid during fieldwork in Puerto Rico. JoeFelsenstein provided free copies of his computerprograms ("PHYLIP"), which I used initially toanalyze phylogenetic data. As a member of theMaryland Center for Systematic Entomology, Iwas also able to use the PHYSYS package writtenbyJ.S. Farris and M.F. Mickevich, with gratefulthanks to the University of Maryland ComputerScience Center for computer time.

Specimens or locality data used during thisstudy were made available by the following peo-ple and institutions (abbreviations in parenthe-ses): G. Arbocco and L. Capocaccia, Museo Civ-ico di Storia Naturale (MCSN), Genoa; N.P. Ash-mole (specimens collected by the joint Ecuado-rean-British Los Tayos Expedition, deposited inthe Museum of Comparative Zoology, HarvardUniversity (MCZ), Cambridge); D. Azuma, Acad-emy of Natural Sciences of Philadelphia (ANSP);C.L. Craig; CD. Dondale, Canadian NationalCollection (CNC), Ottawa; W.G. Eberhard; W.J.Gertsch (deposited in the American Museum ofNatural History (AMNH), New York); J. Gruber,Naturhistorisches Museum (NMW), Vienna;P.D. Hillyard, British Museum (Natural History)(BMNH), London; J. Heiss; H. Homann; M.Hubert, Museum National d'Histoire Naturelle(MNHN), Paris; J.A. Kochalka (JAK); T. Kro-nestedt, Naturhistoriska Riksmuseet (NRS),Stockholm; A. La Touche; H.W. Levi, (MCZ);G.H. Locket; Y. Lubin; N.I. Platnick, (AMNH);S.E. Riechert; C.L. Remington and D. Furth,Peabody Museum of Natural History (PMNH),New Haven; M.J. Scoble and I. Lansbury, HopeDepartment of Entomology (HDEO), Oxford;W.C. Sedgwick; W. Starega, Polska AkademiaNauk Instytut Zoologiezny (PANIZ), Warsaw;M.K. Stowe; University of Vermont Collection

NUMBER 422 3

(UVM), Burlington; C.E. Valerio, Universidadde Costa Rica (UCR), San Jose; J. Wunderlich.

METHODS

This revision is based primarily on the largetheridiosomatid collections of the AMNH andthe MCZ. I made no thorough attempt to borrownon-type material from other institutions, partlybecause the goal was treatment at the genericlevel, and partly because most theridiosomatidmaterial is not sorted as such, but usually is mixedin with theridiids, araneids, or other small ara-neoid groups.

Specimens examined with the AMR 1000 scan-ning electron microscope (SEM) were firstcleaned by hand agitation or ultrasonics, dehy-drated in acetone, and critical point dried incarbon dioxide. Specimens were sputter-coatedwith carbon and gold palladium prior to obser-vation. Micrographs of right-hand structureswere flipped during printing to make the struc-ture appear left-handed in order to ease compar-ison between species.

Most drawings of genitalia were prepared withan Olympus drawing tube mounted on a LeitzSmith Interference Contrast compound micro-scope. Specimens on the stage were manipulatedand oriented as described in Coddington (1983).

Male palpi were expanded by quick (2-5 min)immersion in concentrated KOH (0.2-1.0 g/mlH_»O), followed by several rinses with, and thenprolonged soaking in, distilled water. Full expan-sion in many cases was only obtained after severalKOH-Hv>O cycles. Also, in the case of generawith extensive conductors covering the embolicdivision (Ogulnius, Theridiosoma, Baalzebub, Epi-lineutes, Wendilgarda) the embolic division hadto be levered out from under the conductor witha fine needle, an operation that often damagedthe conductor. Full expansion can be ascertainedby examination with interference microscopy; inan incompletely expanded palp, the hematodo-chal folds are still visible inside the bulb. Thecomplex routing of the reservoir was duplicatedin a wire model, then checked against the speci-

men in several orientations.Female genitalia of non-type material were

dissected out from the abdomen, macerated in awarm trypsin solution for 1 to 5 hours to removeall proteinaceous tissue, and then mounted forobservation with compound microscopy as inCoddington (1983). In the case of holotypes, theentire spider was cleared in clove oil, mountedas above, and examined with incident and trans-mitted light by compound microscopy.

Features consistent for the family or for generaare described in the family description or in thegeneral generic description and not repeatedunder each species description. Measurements ofsomatic morphology were taken with a grid re-ticle in a dissecting microscope. In the case of legarticle lengths, the legs were separated from thespecimen and mounted on a glass slide under acover slip (accuracy usually ±0.02-0.03 mm).Eye diameters are difficult to measure accuratelyon such small spiders, and, in any case, the eyesare rarely round. For the eyes themselves, thedimensions given are of the span of the lens, notincluding any raised tubercle or pigment. Meas-urements are of the maximum span with the eyefeature in question oriented perpendicular to theoptical axis, insofar as that is possible. Similarly,accurate measurements of carapace or abdomendimensions are difficult to obtain. Cephalothoraxheight measurements were made in lateral view,from the surface of the sternum to the top of thecarapace (or posterior median eyes, if higher).Carapace lengths were measured in side viewfrom the rearmost extension of the cephalo-thorax to the clypeal rim (or anterior medianeyes, if longer). Carapace width was measured indorsal view. I have not routinely reported dataon cheliceral teeth, because the great variationin tooth size and placement defies simple descrip-tion (e.g., Figure 2). Length and height of theabdomen were measured in side view, the formerparallel, and the latter perpendicular to the sag-ittal plane of the cephalothorax, not includingany extension of the spinnerets below the ventralsurface of the abdomen. Total length rangesreported in the taxonomic descriptions are for at


least 10 specimens, or of all specimens available,if less than 10.

I give detailed locality information only forspecies known from few specimens, otherwise tothe level of county (USA) or elsewhere to thelevel of state or similar political unit (e.g., Com-isaria, Departamento, Estado, Intendencia, Prov-incia, etc.). In the taxonomic treatments andfigure legends, these units are set in small capitalletters.

As a rule, most old theridiosomatid "type"material is a syntype series. Some authors favorroutine lectotype designation in such cases, but Ifeel that for species in which the syntype series iswholly of that taxon, such designation circum-scribes the action of future taxonomists, andought to be avoided. Also, in some cases (e.g.,Epeirotypus brevipes O. Pickard-Cambridge,1894, or Theridiosoma radiosum (McCook, 1881)(= T. gemmosum (L. Koch, 1878)), the only spec-imens located thus far are probably not of thetype series. Neotypes might be designated, butagain as long as the available specimens fit theoriginal description and no confusion over thename exists, I have avoided such action.

For phylogenetic analysis I initially used theWagner tree algorithm written by Joe Felsenstein(PHYLIP). Those results were corroborated withthe PHYSYS package written by J.S. Farris andM.F. Mickevich and maintained at the Universityof Maryland by the Maryland Center for System-atic Entomology. Characters were coded as pres-ence-absence states, with additive binary codingemployed where necessary to represent complexcharacters.

ABBREVIATIONS USED IN THE FIGURES AND

T E X T .

AC'sAG'sA IKALSAM

AMK(.

( I .

CY (CY's)F.

acimform spigotsaggregate gland spigotsanterior lateral eyesanterior lateral spinneretainpullate spigotanterior median evesconductor<vmhi.il lamellacylindrical gland spigot(s)embolus

EA

ED

FL

MA

()PP i s

PLEPLS

PMEPMS

S T

T

embolic apophysisembolic divisionflagelliform gland spigotmedian apophysisopening of ejaculatory ductparacymbiumpiriform gland spigotsposterior lateral eyesposterior lateral spinneretposterior median eyesposterior median spinneretsubtegulumtegulum

TAXONOMIC HISTORY

Neither the family Theridiosomatidae nor anyof the genera properly included in it have everbeen revised. Archer (1953) reviewed the family,but his work was not in any sense revisionary. Heaccepted all the genera then placed in the family(e.g., Roewer, 1942) and described a new genus(Parogulnius) and several new species. He alsotransferred Chthonos (= Tecmessa) to the theri-diids. Unfortunately he did not borrow any typespecimens and therefore based his nomenclaturaland taxonomic conclusions on published descrip-tions and figures. The opinions of earlier authorsare often accurate, but usually are so poorlydocumented that no sound inferences can bedrawn from their illustrations.

The literature on the family is meager (lessthan 100 papers, including original descriptionsof species), and rather few papers discuss thestatus of the group as a whole (cf. Archer, 1953;Brignoli, 1979; Wunderlich, 1980). Theridioso-matid taxonomy has been chaotic, mainly due tothe lack of a clear, objective diagnosis of thefamily. For example, all the genera originallydescribed in Theridiosomatidae (except, ob-viously, the type genus Theridiosoma) seem tobelong elsewhere (or are synonyms), and no ge-nus accepted herein as a valid theridiosomatidtaxon was ever originally described as belongingto the family. More or less complete turnover ingroup membership at the generic level has oc-curred (Table 1). Of the 21 genera historicallyassociated with the family, 17 clearly belong in

NUMBER 422

TABLE 1.—Allocation of genera historically associated with Theridiosomatidae (* indicatesmonotypy; citations in parentheses indicate authority for previous generic transfers).

Genus Family Comments

*Allototua Bryant, 1945Andasta Simon, 1893

*Billima Simon, 1908* Colphepeira Archer, 1941*Cyathidea Simon, 1907* Cyatholipulus Petrunkevitch,

1930Cyatholipus Simon, 1894

* Diotherisoma di Caporiacco, 1947*Enthorodera Simon, 1907

Epeirotypus Cambridge, 1894* Haliger Mello-Leitao* Helvidia ThoreW, 1890

Ogulnius Cambridge, 1882* Parogulnius Archer, 1953*Phricotelus Simon, 1895*Spheropistha Yaginuma, 1957

Tecmessa O. Pickard-Cambridge,1882

*Tekella Urquhart, 1894* Theridilella Chamberlin and Ivie,

1936Theridiosoma Cambridge, 1879

(in part)Theridiosoma Cambridge, 1879



(in part)* Totua Keyserling

Wendilgarda Keyserling, 1886 (inpart)

Wendilgarda Keyserling, 1886 (inpart)

TheridiosomatidaeTheridiidae (Levi, 1968)Araneidae (Levi, 1978)TheridiosomatidaeSymphytognathidae (Wunderlich, 1978)

TetragnathidaeAraneidaeTheridiosomatidaeTheridiosomatidae

Theridiidae (Levi, 1972)Theridiosomatidae

?Mysmenidae (Brignoli, 1982)Theridiidae (Brignoli, 1982)Theridiosomatidae

Tetragnathidae

Theridiosomatidae

Theridiosomatidae

Theridiosomatidae

Theridiosomatidae

LinyphiidaeTheridiosomatidae

Theridiosomatidae

nomen dubium, type lost= Theridiosoma, new synonymy

= Wendilgarda, new synonymy

Cyatholipinae (Wunderlich, 1978)

= Wendilgarda, new synonymy

incertae sedis

incertae sedis

= Chthonos, new name

Cyatholopinae (Wunderlich, 1978)nomen dubium

= Epilineutes, new genus

= Baalzebub, new genus

= Naatlo, new genus

= Plato, new genus (Platoninae, newsubfamily)

other families, are synonyms, or are nomina du-bia. Most of the genera whose placement hasbeen unstable are monotypic.

Taczanowski described the first theridiosoma-tid as Theridium splendidum in 1873, based onmaterial from Brazil; however, the affinities ofthat species were not recognized until later (Key-serling, 1884). The type genus of the family isinstead Theridiosoma O. Pickard-Cambridge,with T. gemmosum (L. Koch), one of the fewtemperate taxa, as type-species. Koch described

it in 1878 from Germany as Theridium gemmosum,Pickard-Cambridge in 1879 from Britain as Ther-idiosoma argenteolum, and McCook in 1881 fromNorth America as Epeira radiosa.

Keyserling, O. Pickard-Cambridge, and Simonfurther established the character of Theridioso-matidae by describing or transferring Andasta,Epeirotypus, Ogulnius, Wendilgarda, and Phricote-lus to the group. Petrunkevitch (1923:178,1928:144) broadened the definition by includingCyatholipus, Cyathidea, Helvidia, and Totua. Pe-

6 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

trunkevitch also described Cyatholipulus luteus in1930, Mello-Leitao Haliger corniferus in 1943,Bryant Allototua guttata in 1945(a), L. di Ca-priacco Diotherisoma strandi in 1947, and ArcherParogulnius hypsigaster in 1953. Archer (1953)also included Colphepeira catawba in the theridio-somatids, and Roewer (1942) transferred Billimaand synonymized Cyatholipulus with Cyatholipus.Levi and Levi (1962) transferred Enthorodera,Spheropistha, Theridilella, and Tekella. In this, itsmost engorged state, the family contained 21genera.

Since that time the group has shrunk. Levi(1968b, 1972, 1978) transferred Helvidia andBillima to the theridiids and Colphepeira to thearaneids. Gertsch (1960) synonymized Parogul-nius with Trogloneta (Mysmenidae). Wunderlich(1978) provisionally placed Cyatholipus and Tek-ella with the tetragnathids, and Brignoli (1981)returned Spheropistha to theridiids and provision-ally placed Phricotelus in the Mysmenidae.

To what extent these transfers are valid re-mains to be seen. For example, Brignoli (1981)gave no diagnosis of Mysmenidae that supportedthe transfer of Phricotelus. If the relimitation ofMysmenidae by Platnick and Shadab (1978a) isaccepted, neither Phricotelus nor Parogulniusshares the synapomorphies those authors listedfor Mysmenidae. Parogulnius has a structurereminiscent of a parmula in its epigynum—per-haps it is a linyphiid. Wunderlich (1978) basedthe monophyly of cyatholipines (Teemenaarus,Tekella, Tekellatus, and Cyatholipus) on a wide,posterior tracheal spiracle situated far in advanceof the spinnerets, a character otherwise knownin diverse taxa (not necessarily araneoids) whoseaffinities are not well established (Forster, 1959).The alliance of those genera with tetragnathids(Wunderlich, 1978) because of a low clypeus andthe absence of a cheliceral condyle is certainlygrouping on the basis of symplesiomorphies, asoutgroup comparison with Dinopidae or Ulobor-idae demonstrates (see Coddington (in press) forjustification of Dinopoidea as the sister taxon toAraneoidea). Cyatholipulus, Cyatholipus, Haliger,Parogulnius, Phricotelus, Spheropistha, Tekella,

Tekellatus, and Teemenaarus should probably beincertae sedis until a convincing argument basedon synapomorphies allies them with some othertaxon. By incertae sedis I mean "of uncertainaffinities"; if the genera can't be placed withassurance in any well-defined araneoid family, itcan only compound systematic and nomencla-tural confusion to shuffle them between poorlydefined groups. On the other hand, no positiveevidence is available that these transfers are in-correct, so they may as well stand. Certainly theabove taxa exhibit no known characters thatjustify their retention in Theridiosomatidae.Diotherisoma di Caporiacco, 1947, is a synonymof Bertrana (Levi, pers. comm.), and Totua Key-serling, 1891, and Parogulnius Archer, 1953,have epigyna very like linyphiids. At any rate, allof the genera excluded from Theridiosomatidaein Table 1 lack the defining synapomorphies ofthe family (see below).

Two names in Table 1 are nomina dubia. Themonotypic Theridilella Chamberlin and Ivie,1936 (type-species T. zygops, in AMNH, exam-ined), is a theridiosomatid. Levi and Levi (1962)state that the specimen was immature; in any casethe genitalic region has been dissected out andlost. Without the genitalia nothing definite canbe said, but somatically the animal does resembleTheridiosoma goodnightorum Archer, 1953. Thespecimen is undoubtedly a Theridiosoma and, inany case, is undiagnosable, hence the name is anomen dubium. By the published figures, themonotypic Allototua Bryant, 1945 (type-speciesA. guttata, lost), may be a synonym of Ogulnius;on the other hand, it may not be a theridioso-matid at all. Bryant (1945a) described the genusfrom a single adult female, but at present itcannot be found. She mentioned that the labiumwas fused to the sternum; in all theridiosomatids,however, the labial suture is distinct. Also, shegives the order of leg lengths as 1-2-4-3, whereasin Ogulnius the order is 4-1-2-3. Eye proportionsand sternum shape are similar to Ogulnius. With-out any extant specimens, Allototua is also a no-men dubium.

Since the inception of the family, theridioso-

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matids have been heterogeneous and difficult toplace. Their superficial appearance obviouslysuggested Theridiidae to many authors (Chthonos(= Tecmessa), Epeirotypus, Enthorodera (= Wendil-garda), Ogulnius, Theridiosoma, and Wendilgardawere originally described as theridiids). Web ar-chitecture, however, linked theridiosomatidswith the araneoid orb weavers (McCook, 1881,1889a). In synonymizing Theridium gemmosumand Theridiosoma argenteolum, Simon (1881)mentioned that a closely related species occurredin North America, but presumably he was una-ware or skeptical of its status as an orb weaver,because he placed the monotypic genus in its ownsection in the "Theridionidae" (= Theridiidae).He cited L. Koch in stating that its web is "formeede quelques fils irreguliers" (Simon, 1881:27);thus he was unaware that the species spun an orbweb. Theridiosomatids were also placed as ther-idiids by Keyserling (1884, 1886). McCook(1889a) reiterated his evidence that Theridiosomawas an araneoid orb weaver (his European col-leagues apparently thought it might be a cribel-late orb weaver), and Simon (1895) soon con-curred by making the group a subfamily of hisArgiopidae. Simon's concept of Argiopidae wasso broad, however, that by modern standardsthat placement is equivalent to giving it familystatus.

Most subsequent authors have treated theri-diosomatids as a subfamily of Araneidae (= Ar-giopidae), e.g., Comstock (1912), Simon (1926),Petrunkevitch (1928), Wiehle (1931), Gerhardtand Kaestner (1938), and L. di Caporiacco(1938). Berland (1932:95) apparently still consid-ered them theridiids, but in view of his trainingunder Simon it is difficult to believe that theassignment was not a mistake. Apparently, Vel-lard (1924) was the first author to propose fullfamily status for the taxon, but the suggestionwas ignored by his colleagues. Kaston (1948)again assigned the group family status, butArcher (1953) argued against family rank. Opin-ion since then has continued to be divided. Someauthors maintained it as a subfamily of araneids(Locket and Millidge, 1953; Lehtinen, 1967),

although others give it family status (Kaestner,Levi, and Levi, 1980; Yaginuma, 1968; Wunder-lich, 1976, 1980; Brignoli, 1983).

The controversy over theridiosomatids as asubfamily or family is not a puerile argumentabout taxonomic rank; valid issues involving sis-ter group relations are involved. Subfamilial sta-tus in Araneidae implies that theridiosomatidsare more closely related to araneids than to anyother araneoid higher taxon. Inclusion of thegroup in Araneidae has been due primarily tothe occurrence of orb webs in both groups. In-deed, the operational definition of Araneidae,for the most part, has been "a generalized ara-neoid spider producing an orb web," as demon-strated by the steady transfer of "theridiid" spe-cies into Theridiosomatidae upon discovery oftheir orb webs. For example, O. Pickard-Cam-bridge (1894:135) described his new genus Epei-rotypus as a theridiid, saying:

This spider, which is allied to both Theridiosoma, Cambr.,and Ogulnius, Cambr., is even nearer to the true epeirids[= araneid orb weavers] than the former of these two; italso comes near the genus Mesopneustes, Cambr. [= Ther-idula, Theridiidae] . . . .

Two years later O. Pickard-Cambridge(1896:161) transferred Epeirotypus to the ara-neids when he learned that it spun an orb web.

Mr. Smith has the following note on them [that the spidermade an orb web], which is of great interest as showingthat the spider belongs to the Epeiridae rather than theTheridiidae, in which family I had first placed it beforethe facts related by Mr. Smith were known to me . . . .

However, current evidence shows that the orbweb, per se, is plesiomorphic for araneoid spi-ders, and thus cannot be used as evidence ofclose relationship within the superfamily or evenin araneoid family diagnoses (Coddington, inpress; Levi and Coddington, 1983).

Although theridiid-nesticids have not beenconsidered closely related to theridiosomatids forover a century, that lineage may well be the sistergroup of the Theridiosomatidae and symphytog-nathoids together (see below). By "symphytog-nathoids" I mean the Mysmenidae, Anapidae,


and Symphytognathidae; I do not mean to implysuperfamily status for the group. Many of thecomponent groups of Araneoidea still lack ex-plicit, objective diagnoses (for example, the mon-ophyly of araneids, tetragnathids, metids, andlinyphiids is tenuous). At present it seems best tosplit off monophyletic groups from these larger"taxa" and to give them family rank, thus em-phasizing both the objectivity of specific groupsand also our ignorance of their interrelation-ships.

Morphology and Phylogeny

MONOPHYLY OF THERIDIOSOMATIDAE

Throughout the taxonomic history of Theri-diosomatidae, various characters have been usedto define or to identify the group. Here thesecharacters are considered as potentially diagnos-tic and will be discussed and evaluated in turn.

STERNUM BROADLY TRUNCATE BEHIND.—(Si-

mon, 1895, 1926; Wiehle, 1931; Kaston, 1948;Levi, 1982). The character is often used in keysand does, in general, separate T. gemmosum (thebest known north temperate theridiosomatid spe-cies) from other north temperate araneoids, butthe feature is not at all consistent within theridio-somatids (e.g., Figures 76, 85). In addition var-ious other small-sized taxa also have truncatesterna (Platnick and Shadab, 1978a,b; Forsterand Platnick, 1984). Finally, how truncate doesthe sternum have to be before it is "broadlytruncate"? Sternum shape is probably influencedby overall body proportions, and certainly by theobserver's angle of view, so while the featuremight be informative in the context of a morecircumscribed phylogenetic analysis, it is toopoorly defined and too widespread to use in afamily diagnosis.

HIGH CLYPEUS HEIGHT.—(Levi, 1982; Kas-

ton, 1948). Many theridiids, anapids, mysmenids,linyphiids, and even some araneids have a clypeusmore than 2 or 3 times the height of an anteriormedian eye. Also, in some theridiosomatids (Naa-tlo, Baalzebub, Figures 49, 87, 182) the clypeus isrelatively low.

LARGE MALE SEXUAL ORGANS.—(Simon,

1895; Archer, 1953). Some male theridiids (Ti-darren) have palpi that are similarly huge, butthe coincidence is surely homoplasy; however,Chthonos species uniformly have small palpi, asdoes Wendilgarda mexicana.

LEGS WITHOUT SPINES.—(Simon, 1895,

Wiehle, 1931; Kaston, 1948). At present, a"spine" is considered to be a simple extension ofthe cuticle—solid and immovable with respect tothe exoskeleton. In the past authors have notrecognized this difference between spines andhairs, setae, macrosetae, or bristles, or more tech-nical differences such as ennervation, so it isdifficult to know what "lack of spines" specificallymeans. At any rate, spiders in general lack spinesin this strict sense, whereas theridiosomatids dohave robust setae on their legs. The character isbound to cause confusion and does not diagnosetheridiosomatids from other araneoids in eitherits narrow or broad meaning.

FEMALE PALP WITHOUT A CLAW.—(Simon,

1895, 1926; Wiehle, 1931; Kaston, 1948; Levi,1982). Other minute araneoid female spidersthat are probably closely related to theridioso-matids lack palpal claws (Forster, 1959).

RUDIMENTARY PARACYMBIUM.—(Simon,

1926; Archer, 1953; Levi, 1982). The theridio-somatid paracymbium (e.g., Figures 12, 72, 153)is no more rudimentary than that in most ara-neids—a simple hook on the basal lateral marginof the cymbium. Presence of a paracymbiumappears to be plesiomorphic for Araneoidea,and, in any case, the various kinds of araneoidparacymbia (nesticid, araneid, metid, tetrag-nathid, theridiid, cyatholipid, linyphiid, etc.)have not been analyzed in enough detail to arguethat the particular form or placement of thetheridiosomatid paracymbium is unique.

HETEROGENEOUS EYES.—(Wiehle, 1931; Kas-ton, 1948). Homann (1971) showed that canoetapeta are plesiomorphic features of secondaryeyes for a very large group of spiders, as is theabsence of tapeta in the "main," or anterior me-dian eyes. The tapeta of the posterior medianeyes and sometimes the posterior lateral eyes

NUMBER 422

have been lost in numerous araneoid genera(Levi and Coddington, 1983). Although that lossmay be synapomorphic, presence, and thereforethe character "heterogeneous eyes," is certainlysymplesiomorphic in theridiosomatids.

MEDIAN CLAW ELONGATE.—(Simon, 1895,

1926; Wiehle, 1931; Kaston, 1948). This char-acter (Figure 3) may define a monophyleticgroup of spiders including theridiosomatids (seebelow), but it does not define that family alone.It also occurs in mysmenids (Brignoli, 1974; pers.obs.) and nesticids (Wiehle, 1963; pers. obs.).

CHELICERAE WITHOUT A PROXIMAL BOSS.—

(Wiehle, 1931; Kaston, 1948; Levi, 1982). Thecheliceral boss that appears in some araneoids isabsent in many others (linyphiids, theridiids,etc.). Its absence in theridiosomatids is not auseful character.

MEDIAN APOPHYSIS BROAD-BASED OR PRONE

AND PROJECTING.—(Archer, 1953). The medianapophysis of many araneoids is broad based, orprone and projecting, e.g., Araniella (Araneidae:Araneinae). The shape and insertion of the ther-idiosomatid median apophysis varies from a thinlamina (Theridiosoma, Figure 134) to a sclerotizedcurved spur {Plato, Figure 10), to a wide plate(Ogulnius, Figure 99). It is difficult to infer whatthe primitive condition may have been, andhence whether it could be synapomorphic for thefamily.

LARGE TEGULUM.—(Archer, 1953). Mysmen-

ids (Gertsch, 1960; Platnick and Shadab, 1978a)also have relatively large tegula.

None of the characters traditionally used,therefore, accurately diagnose the family Theri-diosomatidae. Undoubtedly this ambiguity hasencouraged the use of the taxon as a repositoryfor genera vaguely like Theridiosoma but sharingno clear-cut derived features. Wunderlich (1980)made a very valuable contribution to the diag-nosis of theridiosomatids when he pointed outthat the sternal pit organs, mentioned by Simon(1907) and Archer (1953), were unique to ther-idiosomatid genera and thus were a convincingsynapomorphy of the family. In the course ofthis revision, other synapomorphies of Theridio-

somatidae have been discovered, and so the mon-ophyly of the group can hardly be doubted.These synapomorphies are as follows.

STERNAL PIT ORGANS.—(Wunderlich, 1980).These structures are located on the promarginof the sternum, adjacent to the labium. In mu-seum material they appear as deep pits (Figures76, 85, 140), but in cleared preparations theyare revealed as glandular structures, with sac-likeinvaginations (Wunderlich, 1980, figs. 3, 4).Their purpose is unknown. Sternal pits are pres-ent in all genera of theridiosomatids exceptChthonos. The presence of several synapomor-phies linking that genus with Plato, which haspits, implies that pits are secondarily lost inChthonos.

PALP CONFORMATION.—The juxtaposition of

sclerites in the unexpanded palp and their ori-entation to each other is unique to the family(Figures 30, 42, 70, 130, 161, 188, 196; seebelow for further explanation).

RESERVOIR.—In all theridiosomatid genera,the route of the reservoir of the sperm duct inthe male palpus is consistent and similar (Figures27, 28, 40, 63, 98, 119, 146, 147, 176). Specifi-cally, the ejaculatory duct is limited to the bodyof the embolus proper. The reservoir (cf. Corn-stock, 1910, 1912; Opell, 1979, for the defini-tions of the regions of the sperm duct acceptedhere) begins at the point where the embolicdivision inserts on the tegulum, loops retrolater-ally to the rear of the tegulum where it makes asharp U-turn around the base of the conductor(Figure 176), and then circles forward and mes-ally beneath the median apophysis (Figure 118).It then bends sharply into the center of thetegulum and out again to the lateral surface,where it executes one or more sharp loops (Fig-ures 62, 176). It again curves retrolaterally tothe rear of the tegulum, again around the baseof the conductor, and mesally forward beneaththe median apophysis, thus paralleling the firstloop described above (Figure 176). It continuesto the ventral wall of the tegulum, and bendssharply into the subtegulum, where it opens intothe large, thin-walled fundus (Figures 28, 41,


118). Many features of the course of the reservoirare consistent among all theridiosomatid generaand thus serve as synapomorphies for the family.Outgroup comparison to other orb-weavinggroups such as uloborids (Opell, 1979), araneids(Levi, 1971), or theridiids (Levi, 1961; Levi andLevi, 1962) indicates that the primitive course ofthe reservoir in the bulb is a simple spiral startingat the insertion of the embolus, continuingaround the margin of the tegulum for one ormore turns, passing into the subtegulum, andending in the fundus. In a sense theridiosomatidreservoirs do the same, with two significant elab-orations. At the outset the reservoir executes asharp turn around the conductor, thus reversingfrom a right-handed to left-handed spiral in api-cal view (Figure 119), and second, in the vicinityof the completion of the first spiral turn (Figures146, 147, 186), the reservoir executes a numberof sharp switchbacks. Of course some theridiids,metids, nephilids, and tetragnathids also have"complex" reservoir trajectories, but no convinc-ing similarity beyond "complexity" itself has beenfound between the trajectory in these taxa andthat in theridiosomatids. (See below for com-ments on symphytognathoids.)

LONG TIBIAL TRICHOBOTHRIA.—Theridioso-

matids have on all tibiae, but especially on theirthird and fourth, numerous dorsal trichobothriawhose length is often 2-4 times the diameter ofthe tibia (Figures 48, 171, 181). The distal tri-chobothrium of the fourth tibia is exceptionallylong. Tibial trichobothria on the dorsum of thefourth tibia of araneids, anapids, mysmenids,symphytognathids, metids, theridiids, nesticids,and linyphiids are relatively much shorter.

Based on the above suite of characters, thespider family Theridiosomatidae is defined toinclude Baalzebub, Chthonos, Epeirotypus, Epili-neutes, Naatlo, Ogulnius, Plato, Theridiosoma, andWendilgarda.

COMPARATIVE MORPHOLOGY OF

THERIDIOSOMATID GENITALIA

Theridiosomatid genitalia have not been de-scribed or illustrated in detail. In view of the

importance of genitalic morphology to phyloge-netic studies, it is desirable to explain clearly andto homologize their morphology with that ofother monophyletic araneoid groups, where pos-sible.

Terms used to describe the parts of the palpfollow Comstock (1910), Merrett (1963), Mil-lidge(1977, 1980), Levi (1971, 1978), and Opell(1979) (see these authors for the basic morphol-ogy of the palp of other orb-weaving spidergroups). Orienting terms (mesal, lateral, distal,proximal, etc.) in the following description followthe usual morphological conventions. In generalthe left palp is figured. Terms such as clockwiseor counterclockwise are used from an observer'spoint of view, looking at a left palp in ventral orapical view. Because the sclerites change theirorientations to each other and to the cymbiumwhen the palp expands, strictly accurate descrip-tion would require different orienting terms forboth states. The change would be very confusing.In the following description, unless otherwisenoted, terms refer to the cymbium, bulb, andsclerites as they appear in the unexpanded bulb.

BASAL PALPAL ARTICLES.—The palpal en-

dites, femur, patella, and tibia are essentiallyunmodified, lacking the stridulatory structures,tubercles, thorns, or apophyses that sporadicallyappear on other orb weavers (e.g., uloborids,araneids, anapids, etc).

CYMBIUM.—The cymbium is a broad, cup-shaped segment with the usual basal alveolus onthe ventral surface. The mesal basal margin ofthe cymbium bears a paracymbium (Figures 12,32, 50, 72, 102, 153, 164, 191, 216), always amore or less simple hook. A spine may (Figure102) or may not (Figure 216) occur on the distalend of the paracymbium. The cymbium of Baal-zebub (Figure 164), Epilineutes (Figure 191),Ogulnius (Figure 102), Plato (not figured), andWendilgarda (Figure 216) has an additional la-mella situated just distally to the paracymbiumon the margin of the cymbium (lacking inChthonos, Epeirotypus, and Naatlo). The tip of thecymbium and the distal margin of the alveolus isdistinctly pointed in most species of Plato and

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Chthonos, blunt in the remaining genera. Themesal margin of the cymbium in most species ofPlato has several notches (Figure 10).

PALPAL BULB.—The theridiosomatid palpalbulb is tripartite, consisting of the subtegulum,tegulum, and embolic division. The bulb is at-tached to the cymbium by a basal hematodocha.The basal hematodocha attaches proximally tothe alveolus of the cymbium, and its distal endinserts on the cylindrical basal margin of thesubtegulum. The petiole is apparently absent,but I am not certain of this fact. During expan-sion, the basal hematodocha inflates considerablyand may rotate the bulb through as much as 200degrees with respect to the cymbium. The bulbalways rotates counterclockwise with respect tothe cymbium in the left palpus and so the origi-nally mesal median apophysis (e.g., Figure 10,palp unexpanded) approaches the paracymbiumon the lateral cymbial margin (Figure 28, palpexpanded). In my preparations the medianapophysis did not, however, touch or engage theparacymbium as Heimer (1982) would predict.In an unexpanded bulb, therefore, a diagramshows the mesal aspect of cymbium and bulb, butin an expanded bulb it will show, for example,the mesal aspect of the cymbium and the lateralaspect of the bulb or visa versa.

The subtegulum (ST, Figures 11, 30, 42, 70,135) is a cup-shaped or cylindrical sclerite,slightly longer or deeper on the ventral side thanthe dorsal (in an unexpanded palp). The distalventral margin appears to be weakly fused to thetegulum. The point of fusion provides a fulcrumabout which the tegulum and embolic divisionpivot slightly during expansion (Figures 62, 63).Movement of the distal portions is accomplishedby expansion of the median hematodocha, by farthe largest hematodocha of the palpus. The sub-tegulum also contains most of the fundus of thesperm duct, a large, thin-walled sac near theventral wall of the subtegulum, where the reser-voir of the sperm duct inserts. In many palps theform of the fundus is difficult to decipher, andso the appearance of the structure in the illustra-tions is only diagrammatic.

The tegulum is a ring-shaped sclerite whoselateral aspect is hugely expanded. The lateralsurface usually has a dark stripe (Figure 27). It isdeeply split on its mesal, dorsal side, and theconductor sits inside the cleft (Figure 21). Themedian apophysis sits at the end of the mesal armof the split, and the embolic division originateson the lateral margin of the split. The tegulumalso contains the reservoir of the sperm duct,whose trajectory, as described above, is ex-tremely complex. Even though the tegulum ismuch modified, it can be recognized as suchbecause the course of the reservoir is entirelycontained within it, and it bears the medianapophysis and conductor, as is the case in nearlyall spiders. The median apophysis (Figures 10,29, 42, 70, 99, 133, 163, 188, 196) is an articu-lated sclerite arising from the mesal arm of thesplit tegulum. The median apophysis is variouslyshaped and can have different lobes or apo-physes, but in mesal view it is always in the sameplace, at the distal end of mesal arm of thetegulum. The reservoir of the sperm duct neverpasses through the median apophysis itself.

The conductor lies within the split of the te-gulum. It is a large, over-arching structure thatto a greater (Baalzebub, Epilineutes, Ogulnius,Theridiosoma, Wendilgarda) or lesser extent(Chthonos, Epeirotypus, Naatlo, Plato) covers theembolus when the palp is contracted. The con-ductor may have various lobes (Figures 11, 30,133, 137, 189). The more distal apophyses areoften pointed, and during expansion the pointrocks forward and rubs against the surface of thetegulum (Epeirotypus, Theridiosoma). The conduc-tor also seems to have a hematodocha between itand the lateral portion of the tegulum. Whenexpanded, this hematodocha to some extentpushes the embolus away and out from under-neath the conductor.

The embolic division of the palp arises fromthe tegulum near the connection with the con-ductor. It curls in a counterclockwise fashion (leftpalp, ventral view). The embolus is a simplestrong sclerite (Chthonos, Plato, Epeirotypus, Naa-tlo) or it has a mesal apophysis emerging at the


base, here termed the "embolic apophysis" (Baal-zebub, Epilineutes, Ogulnius, Theridiosoma, Wen-dilgarda). No hematodocha occurs between theembolic apophysis and the embolus, and nonebetween the embolic division and the tegulum.The embolic apophysis may either be a bifurcatebristle {Ogulnius, Figure 101), fragmented (Ther-idiosoma, Figures 131, 133, 136), or tripartite(Baalzebub, Epilineutes, Wendilgarda, Figures162, 190, 198). The embolic apophysis is appar-ently an autapomorphy of these genera, becausethe immediate outgroups of theridiosomatidslack any embolic apophyses at all, and further-more the apophysis of theridiosomatids differsconsiderably from the "terminal apophysis" ofother araneoid taxa such as linyphiids or ara-neids. Plato species may also exhibit mesal em-bolic apophyses (Figures 10, 22), but the homol-ogy of these more distal structures with thosediscussed above is not certain.

The male palp therefore offers many taxo-nomically useful characters. No doubt the femalegenitalia are similarly useful, but because thestructures are soft, less sclerotized, and involvedissecting the animal, I have studied them moresuperficially. Of course, male and female geni-talia complement each other. One expectsthe oversized male genitalia to be equalled by thecapacious copulatory bursae of the female. Like-wise the simplest routing of the copulatory ductsis matched by those males having simple embolicdivisions. Functionally related parts are not, afterall, homologues, and thus can be considered in-dependently derived apomorphic features.

The theridiosomatid epigynum is always a con-tinuous, usually simple, plate covering the open-ing of the copulatory bursae (Figures 25, 37,112, 151, 206). Similar plates occur in symphy-tognathoids, but the feature seems too nondes-cript to use in phylogenetic studies. The poste-rior rim may (Chthonos, Epeirotypus, Naatlo, Ogul-nius, Plato) or may not (Baalzebub, Epilineutes,Theridiosoma, Wendilgarda) have a transversegroove, sometimes interrupted medially by aslight longitudinal ridge. Most genera also havea knob, pit, or cuticular thickening in the central

region of the epigynal plate (Figures 18, 50, 173,183, 219). The function of this structure is un-known, but in Epeirotypus a well-developed mus-cle extends from it towards the pedicel, probablyenabling the female to reflect the epigynal rimwhile mating (Figure 5). Thus it may be anapodeme.

The copulatory pores are large and widelyspaced, often merging into a common atrium.The copulatory bursae, by which I mean theouter ends of the copulatory ducts and theircommon atrium, are also capacious. It is some-times difficult to tell from published illustrations,but apparently such capacious bursae are rare inAraneoidea (cf. Theridiidae, Levi and Levi,1962; Araneidae, Levi, 1968b; metids, tetrag-nathids, Levi, 1980b; Symphytognathidae, Fors-ter and Platnick, 1977; Anapidae, Mysmenidae,Platnick and Shadab, 1978a,b). In Baalzebub,Epilineutes, and Wendilgarda the bursae have lat-eral wings or pockets (Figures 174, 184, 215).

In many cases, the juncture between copula-tory bursae and ducts is indistinct. In some prep-arations the bursae seem to extend as blind pock-ets beyond the beginning of the narrow sper-mathecal ducts (Figures 26, 60, 174); in themajority the two seem continuous, the bursaesimply narrowing to form the copulatory ducts.Sectioned, stained preparations might settle thequestion.

The course of the bursae or ducts varies froma simple hairpin turn (Epeirotypus, Naatlo, Fig-ures 60, 95) to more convoluted arrangements(e.g., Ogulnius, Figure 113; Wendilgarda, Figure215). Duct routings tend to be diagnostic forgenera, but no feature beyond their relativelylarge size seems to be synapomorphic for thefamily.

On the other hand, the fused condition oftheridiosomatid spermathecae (Figures 26, 195,220) is probably synapomorphic for the family(compare references cited above for other ara-neoid taxa). The spermathecae share their me-dian wall, although in some derived groups theyare connate only at their distal tips (e.g., Baalze-bub, Epilineutes, Figures 174, 184).

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The copulatory ducts usually insert on thelateral sides of the spermathecae, no doubt be-cause the medial sides are fused. However, theducts of derived genera nevertheless pass overthe dorsal surface of the spermathecae towardsthe posterior and enter the spermathecae moreor less on their mesal surfaces (Baalzebub, Epili-neutes, Theridiosoma, Wendilgarda, Figures 145,174, 184,207).

The fertilization ducts are simple, short spursthat pass from the spermathecae to the vagina,and I made no attempt to use them in the analy-sis. Implications of these characters and othersfor the phylogeny of the family and its relation-ship to other araneoid families are evaluated inthe following section.

INTERFAMILIAL RELATIONSHIPS

Our understanding of the phylogeny of theAraneoidea is so chaotic that it is impossible todiscuss succinctly the placement of Theridioso-matidae in the superfamily. Some attention hasto be paid to the phylogeny of the superfamily asa whole, if only to evaluate the data for anyparticular placement of Theridiosomatidae.

At present, Araneoidea is the largest spidersuperfamily, containing about 12,000 describedspecies (estimated from Levi, 1982), or about athird of all described spider species. Araneoidspiders have an easily recognized gestalt, and thesuperfamily is doubtless monophyletic (see Cod-dington (in press) for a review of the evidence).The only controversy about the composition ofAraneoidea concerns the archaeid, microphol-commatid, textricellid, and mimetid lineages. Onthe basis of two synapomorphies (peg-shapedcheliceral teeth, distinctive cheliceral glandmound), Forster and Platnick (1984) removethese families from the araneoids and place themwith the huttoniids, stenochilids, and palpiman-ids, in the superfamily Palpimanoidea. It may besimpler (in the sense of competing phylogenetichypotheses) instead to transfer those families intoAraneoidea because of the contingent necessityof supposing the similarities between the former

taxa and araneoids to be convergent. For exam-ple, a paracymbium-like structure occurs in Pal-pimanoidea, some have serrate hairs, and somealso have labia wider than long, all features oth-erwise typical of Araneoidea. However, the ho-mology of those rather vaguely defined charac-ters in Araneoidea and Palpimanoidea is onlyspeculative, and, in any case, their absence inother spider taxa that might serve as outgroupsto either superfamily has not been documented.On the other hand, the removal of archaeids andmimetids conveniently makes Araneoidea a com-pact group of web-spinning spiders capable ofproducing sticky silk (as far as is known thePalpimanoidea sensu Forster and Platnick (1984)lack aggregate silk glands and consequently theability to produce sticky silk). The problem is adifficult one requiring further research on palp,spinneret, and silk gland morphology, at least.

The controversy affects the placement ofTheridiosomatidae within the Araneoidea onlybecause Archaeidae and Mimetidae need nolonger be considered as potential sister groupsof theridiosomatids if their removal to the Pal-pimanoidea is correct. In any case, I know of noconvincing features that would ally theridioso-matids with any of these taxa, so the questionthen becomes, which araneoid group (theridiid-nesticid, symphytognathoid, linyphiid-araneid,metid-tetragnathid) is the sister group of theri-diosomatids? For justification of the monophylyof theridiids-nesticids and symphytognathoids,see Coddington (in press). The monophyly ofaraneids and linyphiids is simply a working hy-pothesis, based on the admittedly slim but appar-ently synapomorphic evidence of a radix in theembolic division of the male palp. On the otherhand, the group "metid-tetragnathid" is almostcertainly para- or polyphyletic. I link metids andtetragnathids here only to simplify the followingarguments. As is argued below, the monophylyof metid-tetragnathids is in any case irrelevant tothe question of the placement of theridiosoma-tids.

The traditional placement of theridiosomatidshas been next to the "araneids"—itself appar-


ently not a monophyletic group if broadly de-fined to include nephilids, metids, and tetrag-nathids. As mentioned above, theridiosomatidgenera were typically first described as theridiidsand then, as their orb webs were discovered,transferred to the araneids. Earlier authorstended to include all orb-weaving araneoid spi-ders in a single taxon—Araneidae or Argiopidae,defined by the orb web. However, all currentevidence suggests that the orb web of uloboridsand dinopids is homologous to that of the ara-neoid orb weavers (Coddington, in press). Giventhat Araneoidea is itself monophyletic, as is Di-nopoidea, the orb web is therefore a synapomor-phy for the two superfamilies and, therefore, byoutgroup comparison also primitive for Araneo-idea. Consequently the occurrence of an orbweb, per se, is no justification for uniting any orbweavers within the Araneoidea, specifically Ara-neidae and Theridiosomatidae.

Even though one classic group (Araneidae,sensu lato) based on symplesiomorphic behav-ioral characters turns out to be paraphyletic,behavioral features still seem to be the mostuseful character system for placing theridioso-matids with their closest araneoid relatives.These characters are discussed briefly here andat more length in Coddington (in press).

INNER SS LOOP LOCALIZATION.—Orb-weav-ing spiders have basically two ways that theycontact the innermost sticky silk (SS) loop duringthe construction of the sticky spiral (Eberhard,1982; Coddington, in press). Uloborids, dino-pids, and araneids (Araneinae, Gasteracanthinae,at least) use a lateral tap of the outside first legto touch the outermost SS loop prior to connect-ing the SS segment they are currently spinning.Because the Dinopoidea (Uloboridae and Dino-pidae) are the sister taxon of the Araneoidea(Coddington, in press), that method would seemto be plesiomorphic for Araneoidea. Metids, te-tragnathids, theridiosomatids, symphytognath-ids, anapids, and mysmenids use a differentmethod, a forward tap of the inside first leg (seeEberhard, 1982, for figures). Eberhard (1982)found surprisingly little homoplasy in this

character, and once seen, the difference is verystriking and consistent. Hence the feature seemsto define a subsidiary monophyletic group ofaraneoid spiders containing the taxa listed above.

Two problems must be mentioned at thispoint. First, "metid-tetragnathids" are almost cer-tainly a paraphyletic group (Coddington, inpress). Palp structure supports the monophyly oftetragnathids {Tetragnatha, Mimognatha, Pachyg-natha, Glenognatha, Azilia, at least), but the char-acters mentioned by Levi and Coddington (1983)(elongate chelicerae, "modified paracymbium")are poorly defined and weak synapomorphies, atbest, for a group including Meta and tetragnath-ids. Almost no published information is availableon "metids," so the problem can only be acknowl-edged until more research is completed. An in-side first leg forward tap is one of the very fewcharacters systematically investigated across allaraneoid lineages and it does support the mono-phyletic group defined above.

Second, tetragnathids are haplogyne spiders,whereas all the other Araneoidea are entelegyne.Haplogyny and entelegyny are supposed to befundamentally different conditions, and the for-mer primitive with respect to the latter based onoutgroup comparison to Mesothelae and the hy-pochiloid taxa (Brignoli, 1975; Platnick, 1975;Opell, 1979). Haplogyny in tetragnathids thusimplies that they are the sister group of all otheraraneoids, not part of the subsidiary araneoidgroup defined above. The behavioral and mor-phological data conflict. However, as Opell(1979) suggested for Uloboridae, entelegyny canbe independently derived within monophyleticlineages. Forster and Platnick (1984) reached thesame conclusion for Palpimanoidea. The conflictcould be resolved by a detailed study of theentelegyne condition in araneoid lineages. Thebehavioral evidence suggests that it has arisentwice, and thus one might expect to find twokinds of entelegyny in Araneoidea. Another pos-sibility is that tetragnathids are secondarily hap-logyne.

No detailed accounts of theridiid, linyphiid, ornesticid web building have been published. Some

NUMBER 422 15

theridiids, certainly, sequentially connect stickysilk lines to non-sticky lines, and thus might bescored for the manner in which they "measure"the spacing of the sticky silk line. At any rate,the condition for this character in any of theabove three lineages is unknown.

At least three characters, however, link liny-phiids with Araneidae (sensu stricto), and so lin-yphiids may also be discounted as a potentialsister group of theridiosomatids. For example,only linyphiids and araneids possess gnathocoxalsexual glands (Lopez, 1977). Likewise, only lin-yphiids and araneids possess a radix in the malepalp. By radix I mean a sclerite articulated to thetegulum and bearing the embolus and a "termi-nal apophysis." (The "radices" of theridiids (e.g.,Levi, 1961) or uloborids (Opell, 1979) do notfulfill this classical definition and thus are prob-ably autapomorphies of each of those taxa, re-spectively.) The "terminal apophysis" of liny-phiids and araneids is also synapomorphic for thetwo taxa. By terminal apophysis I mean a scleriteof the embolic division (thus not inserting on thetegulum) that inserts on the radix. Alterna-tively, the composite character "complex embolicdivision" could be considered a single, complexsynapomorphy for linyphiids and araneids. More-over, no convincing characters linking theridio-somatids to linyphiids have been discovered, andso linyphiids can be dropped from further con-sideration.

Two morphological characters do link thetheridiid-nesticid lineage with Theridiosomati-dae and symphytognathoids (i.e., Anapidae, Mys-menidae, and Symphytognathidae). These char-acters are cheliceral denticles (Figure 2) and elon-gate median claws (Figure 3). The former char-acter, however, occurs in at least Hyptiotes (Pe-ters, 1982) and Nephila clavipes (R.R. Forster,pers. comm.; pers. obs.) and thus cannot be asynapomorphy for theridiosomatids, symphytog-nathoids, and theridiid-nesticids.

An elongate median claw is ubiquitous in nes-ticids but unknown in theridiids. Nesticidae maybe a paraphyletic assemblage of primitive theri-diid genera or, at best, the sister group of theri-

diids (Coddington, in press). If the former case,elongate median claws could indicate the mono-phyly of theridiid-nesticids, theridiosomatids,and symphytognathoids.

Symphytognathoids themselves are monophy-letic by one very consistent behavioral featureand, possibly, two additional morphological fea-tures. After the completion of the sticky spiral,the spider adds numerous "accessory" radii,which extend from the hub to the frame lines(described in Eberhard, 1981; photographs inCoddington, in press). Among the web-spinningmysmenids, the genus Mysmena is autapomorphicand has lost the behaviors concerned with acces-sory radii ( Coddington, in press). These acces-sory radii cross the sticky spiral but are notcemented to it, whereas the sticky spiral is ce-mented to each structural radii it crosses. (Bystructural radii I mean the radii constructed dur-ing the discrete behavioral sequences defined asframe and radius construction in Coddington (inpress), not that structural radii necessarily con-tribute more to the strength of the web.) Eber-hard (1981) felt that the silk composing the ac-cessory radii was finer than that of the structuralradii, hence perhaps from a different gland al-together.

All symphytognathoids lack paracymbia. If thesister group relationship between Theridioso-matidae and symphytognathoids is accepted, thisparticular lack of a paracymbium is unique andunreversed in these taxa (see remarks about pal-pimanoids above). Nearly all other araneoidshave some sort of paracymbium (Heimer, 1982).Second, the male palpi of symphytognathoidshave, at most, one tegular apophysis, whereasother araneoids usually have two, the medianapophysis and the conductor. Whether the re-maining apophysis on the symphytognathoidpalp is a conductor or a median apophysis ismoot. Theridiosomatids have two tegular apo-physes. By the same logic applied to the evidenceof the paracymbium, the single tegular apophysisof symphytognathoids may be an additional syn-apomorphy for the group.

The sister taxon of Theridiosomatidae appears


to be the symphytognathoid taxa (Mysmenidae,Anapidae, Symphytognathidae) as a whole. Fivecharacters support this relationship—two strongbehavioral synapomorphies, one which is debat-able, and two morphological features that re-quire more research before their significance canbe fully substantiated.

HUB LOOP CONSTRUCTION AFTER SS CON-

STRUCTION.—Most araneoids (Coddington, inpress) either do not modify the hub region at allafter completing the sticky spiral, or they biteout the hub center and either leave it as a holeor fill it in with irregular lines around or acrossthe hole. Theridiosomatids, anapids, mysmenids,and symphytognathids always not only bite outthe hub but add a regular outward spiral of loopsto the hub region, usually completely replacingthe hub (Eberhard, 1981). The behaviors in-volved (Coddington, in press) closely resemblethe same behaviors used to construct the non-sticky spiral that all orb weavers construct priorto beginning the sticky spiral itself. Their ap-pearance in theridiosomatids, anapids, mysmen-ids, and symphytognathids in a completely differ-ent and later part of the overall web-buildingsequence is a uniquely derived character unitingthese families.

EGGSAC DOUBLY ATTACHED.—As far as I

know, most symphytognathoids (Symphytognathaglobosa is an exception; Hickman, 1931) retainthe eggsac near the hub of the web, and it isattached to the web by two lines, hence "doublyattached" (photographs in Coddington, in press).The more primitive theridiosomatid genera dothe same. Double attachment in Epeirotypus (Fig-ure 44) and Ogulnius (Figure 104) arises becausethe spider begins eggsac construction on a trans-verse silk line, rather than from the bottom of asingle pendant line (pers. obs.). Often the eggsacbecomes elongate along the axis of the silk lineas well. Even if the spider later cuts one of theattachment lines to the eggsac so that it is sec-ondarily pendant (e.g., Plato, Figure 13, someOgulnius, Figure 106, some Epeirotypus, Figure68) the slight remnant of the second attachmentline visibly persists on the lower apex of theeggsac.

Placement and attachment of eggsacs in me-tids, tetragnathids, nesticids, and theridiids isdiverse within and between taxa. In any case,none of those lineages behave in a manner con-sistently similar to that uniting theridiosomatidsand symphytognathoids.

RADII LENGTHENED.—The third behavioral

similarity is ambiguous. Theridiosomatids (ex-cept Ogulnius, Epeirotypus, and probably Naatlo),anapids, mysmenids, and symphytognathids cutand lengthen all structural radii built duringframe and radius construction. They do so afteradding the secondary hub loops described above.Theridiosomatids also join the lengthened struc-tural radii, thus "anastomosing" them (Figures157, 159, 192, 194). Symphytognathoids cut and

join their accessory radii to the radii built duringframe and radius construction, but the latterradii are not joined to each other.

For several reasons the two kinds of radialanastomoses, one joining structural radii to-gether, the other joining accessory to structuralradii, seem too dissimilar to infer homology atthis point. First, however similar the two kindsof radius-joining behaviors may be, they act onnon-homologous substrates, i.e., accessory versusstructural radii, and in slightly different ways.The inference of homology in a process such asbehavior is more convincing if the substrate onwhich the behavior acts is also homologous, andin this case it is not. Second, symphytognathoidsdo have structural radii, and don't anastomosethem, although no obvious engineering or func-tional reason prevents it. On the other hand,theridiosomatids lack accessory radii entirely, soit is unknown whether, if they had the opportu-nity, they would anastomose those accessory ra-dii. Third, among theridiosomatids, Epeirotypusand very probably Naatlo do not cut, lengthen,or anastomose structural radii. If, as seems likely,these genera compose the sister taxon to the restof the theridiosomatids, "radial anastomosis" mayhave been derived independently in the remain-ing theridiosomatids and in the symphytognath-ids. Otherwise, one could presume secondary lossof radial anastomosis in Epeirotypinae, and, con-sequently, that the anastomosis of radii, whether

NUMBER 422 17

structural or accessory, is also synapomorphic fortheridiosomatids and symphytognathids.

FEMALE PALPAL CLAW ABSENT.—As far as Iknow, only the females of theridiosomatids, mys-menids, anapids, and symphytognathids amongaraneoids consistently lack claws on their palpi.Of course, the palp of female anapids is usuallysomewhat reduced in size, and that of symphy-tognathids is reduced to a single segment orcomplete absence, but, nevertheless, the palplacks a claw. Mysmenidae appears to be the sistergroup to anapids and symphytognathids, and theclawless palps of mysmenid females are quitecomparable in size and development to those oftheridiosomatids.

RESERVOIR SWITCHBACK IN MALE PALP.—Asmentioned above, the primitive course of thereservoir in araneoids is one or two unreversedspirals. In Theridiosomatidae (see above, andFigure 176) and at least Anapidae and Mysmen-idae of the symphytognathoids, the reservoirstarts a spiral in one direction but loops back onitself almost immediately and thereafter followsa reasonably consistent spiral course in the otherdirection. I have not studied the routing of thereservoir in symphytognathids, but in view of theother characters linking the group with Theri-diosomatidae, the similarity is suggestive—pos-sibly an additional synapomorphy.

One reason telling against this feature as asynapomorphy for theridiosomatids and symphy-tognathids is the complexity of the reservoirrouting in nephilids, metids, and tetragnathids(Levi, 1980b, 1981). Although no convincingdetailed similarities in reservoir routing betweenthe former and the latter taxa have emerged, allthese routings are complex and will certainlyyield useful phylogenetic information if furtherstudied. One feature does speak against the ho-mology of the reservoir route complexity in theabove taxa. In metids, tetragnathids, and nephil-ids, the complexity occurs immediately proximalto the insertion of the embolus on the tegulum,whereas in theridiosomatids the reservoir com-pletes the first switchback and circuit of thetegulum before becoming markedly complex(Figures 147, 176). Thus the "complexity" arises

in somewhat different regions of the reservoir.In summary, the tetragnathids, metids, theri-

diosomatids, and symphytognathoids, and possi-bly also theridiid-nesticids, form a natural groupwithin the Araneoidea (Coddington, in press).With respect to the first four lineages, one com-ponent unites theridiosomatids and symphytog-nathoids, and another unites tetragnathids-me-tids to that pair of taxa. The placement of Ther-idiidae-Nesticidae, however, is uncertain; the lin-eage may eventually be placed at any of the fivepossible positions within the following threetaxon statement (metid-tetragnathids, (theridio-somatids (symphytognathoids))).

INTERGENERIC RELATIONSHIPS

A cladistic analysis of the family indicates foursubfamilies, Platoninae, Epeirotypinae, Ogulni-inae, and Theridiosomatinae (Figure 1). Char-acters used in the cladogram draw on a largermatrix of about 150 characters routinely scoredfor theridiosomatid taxa during the course ofthis revision. The majority have been omittedfrom Figure 1 because they are (1) synapomor-phies at subgeneric levels; (2) autapomorphic forgenera; (3) characters initially thought to be in-formative but later found to be too vaguely de-fined to be usable; (4) known only for a smallsubset of the studied taxa; or (5) important be-havioral features. Behavioral features are ofcourse mentioned throughout, but insofar as amajor goal of this revision has been to analyzetheridiosomatid morphology in preparation fora test of behavioral phylogenetic hypotheses, onecannot first include the behavior to construct atree, and then use the tree to analyze the behav-ior. Even if behavioral features are added to thedata set presented in Figure 1, no clear resolutionof the basal trichotomy results.

The cladogram of Figure 1 is the shortestpossible tree by the following argument. No char-acters substantially refute the monophyly of Pla-toninae, Epeirotypinae, or the sister group rela-tion between Ogulniinae and Theridiosoma-tinae. Thus the cladistic issue at the subfamilylevel is a three-taxon problem. I used the PHY-LIP and PHYSYS Wagner algorithms to ascer-


Cymbium pointed at tip

Alveolus margin pointed

Conductor with thick mesal ap<

PC T-shaped

MA with long, recurved tip

HA round with notch

PC A broadly attached blade

Epigynum a bulging done

Tegulum laterally expanded

Conductor with brown stripe

Epigynum with scape

MA with denticles

MA subrectangular

MA with deep trough

ED parts broad and blunt

Copulatory bursae winged

Cop. ducts collapsed, plicate

Eggsac with suture

Tegular spur

Eggsac singly attached

Tegular striae/denticles

Row of bristles on lamella

Epig. with lateral pits

MA with trough or groove

F-R junction dentate

Cop. ducts insert mesally

Switchback #2 in reservoir

Switchback #3 in reservoir

Conductor covers whole ED

Embolic apophysis present

Embolus a thin lamina

Embolic pore basal

Copulatory ducts convoluted

Cymbial lamella present

Rod/tube in reservoir

ST transparent over fundus

Abd, Ceph, Ster, unicolorous

Sternum papillate

Sternum truncate

Epig. with transverse groove

Sternal pits

Conductor hood shaped

Long Tb IV trichobothria

Central pit on epig. plate

Course of reservoir

Palp conformation

Spermathecae connate

II II II

PLATONINAE

EPEIROTYPINAE-THERIDIOSOMATINAE

OGULNIINAE

Fioi'RF. I.—Cladogram of theridiosomatid genera. Consist-ency index 0.82; autapomorphies of genera omitted; char-acters discussed in text. (Open squares = primitive states,closed = derived.)

tain that the lengths of each possible solutionwere equal, thus proving that a basal trichotomyis the appropriate representation of the data. Ina like manner, I specified all 105 possible rootedtrees for the five-taxon statement (Ogulnius,Theridiosoma, Baalzebub, Epilineutes, Wendil-garda), with the root always basal to Ogulnius.Figure 1 is the shortest tree, with a consistencyindex of 0.76. The three implied trees do differin F ratio. Of these, a sister group relationshipbetween Platoninae and Ogulniinae-Theridioso-matinae affords the least variance between treeand distance matrix (F = 9.4 vs. F = 10.1 and10.3 for the other two trees). In practice andtheory, however, F statistics offer scant reason toprefer one tree to another.

Observation of theridiosomatid web construc-tion shows that the more bizarre webs, for ex-ample those of Ogulnius (Figures 103, 105) andWendilgarda (Figure 202), employ only a subsetof behaviors that by outgroup comparison areprimitively part of the orb-web construction al-gorithm. Ogulnius and Wendilgarda, for example,lack non-sticky spiral construction, radius con-struction, frame construction, and hub modifi-cation (see Coddington (in press) for brief de-scriptions). Computer algorithms obviously don'tdistinguish between secondary loss and primitiveabsence. If these and other behavioral charactersare added to the data set, the consistency indexdrops considerably, but the form of the tree doesnot change, because of the massive evidence forthe monophyly of Theridiosomatinae.

PLATONINAE.—Platoninae includes the generaPlato and Chthonos. Five derived characters sup-port the monophyly of the subfamily, all featuresof the male palpus. First, the cymbium of allChthonos and all except one Plato (the exceptionis undescribed) are distinctly pointed; second, thedistal margin of the alveolus is also pointed (Fig-ures 10, 29). Corresponding features in othertheridiosomatid genera and family outgroups arerounded. Third, the conductor has a thick, re-curved, ventral apophysis, very similar in bothtaxa (Figures 10, 29). Outgroups to the familylack the particular form of the theridiosomatid

NUMBER 422 19

conductor in general, and the conductor apoph-ysis in particular. Theridiosoma species may haveapophyses on the conductor (Figures 133, 156),but they originate from different points on theconductor and are neither so recurved nor sorobust. Fourth, the paracymbium of bothChthonos and Plato is somewhat T-shaped (Figures12, 32). It hooks toward the tip of the cymbium,as do the paracymbia of other theridiosomatidgenera, but also has a backwardly directed lobethat gives it the form of a T. Fifth, the distal tipof the median apophysis is long, recurved, ro-bust, and quite sclerotized (Figures 10, 29). Themedian apophysis of Theridiosoma may be atten-uate and recurved, but it is a weakly sclerotizedstructure with a dorsal groove and pointed tip.That ofOgulnius is a much broader lamellar platewith differently shaped apophyses.

Observations of webs of Plato bruneti, newcombination, in Trinidad and P. guacharo, newcombination, and P. miranda, new combination,in Venezuela confirm that the genus does anas-tomose radii in the same manner as Theridiosomaand Epilineutes (e.g., Figures 159, 192). On theother hand, Plato species do not isolate a primaryradius for use as a tension line as do Epeirotypus,Naatlo, and Theridiosoma (Figures 45, 69, 157).In that respect, Plato is more similar to Epilineutes(Figure 192). Moreover, Epilineutes and Platooften sit at the periphery of their webs exertingnoticeable tension on a radial line. Plato is moresimilar to Epeirotypus in that they construct nu-merous closely spaced non-sticky spiral loops dur-ing frame and radius construction, whereas inTheridiosoma and Epilineutes the ratio of framesand/or radii constructed during non-sticky spiralconstruction is much higher (see Coddington (inpress) for definitions and descriptions of thesebehaviors). Outgroup comparison to very primi-tive orb weavers (Dinopidae, Uloboridae) indi-cates that a low ratio is primitive, but moreclosely related taxa (Anapidae, Mysmenidae)have as high a ratio as theridiosomatines.

The taxonomic affinity of Chthonos, of course,has always been controversial, but Chthonos iswithout doubt a theridiosomatid by palp and

epigynal characters. Chthonos are specialized spi-ders that do not spin webs, and their body form(Figures 33, 34) certainly differs from that ofPlato (Figures 14, 23). Female genitalia of bothgenera are very similar, but the similarities areall primitive features of the family in general.

EPEIROTYPINAE.—This subfamily includesEpeirotypus and Naatlo. Four derived featuresdefine the group. First, the median apophysis isa round, notched disk in both genera, uniqueamong theridiosomatids (Figures 42, 70). Sym-phytognathoids lack median apophyses alto-gether. Second, the paracymbium of both generais a flat, blade-like apophysis, broadly attached tothe cymbium (Figures 53, 72). Other theridio-somatid paracymbia are T-shaped (see above) orelse rounded hooks. Third, the epigynum in bothgenera is a convex bulging dome, whose poste-rior margin is closely appressed to the abdominalventral wall (Figures 50, 58, 85). The epigyna ofother theridiosomatids and symphytognathoidsare usually flat plates (or occasionally concave inlateral view), rather different from Epeirotypusand Naatlo. Fourth, the tegula are much ex-panded on their lateral faces (Figures 54, 65, 83).

The female genitalia also provide synapo-morphic features, probably related functionallyto the proportions of the palpi in the males. Theroute of the copulatory ducts is short, and theymake a simple acute turn. The distinction be-tween copulatory bursae and ducts is not marked.Details of surface structure and pigmentation,difficult to illustrate, are also very similar in thetwo genera.

Behavioral features also support the mono-phyly of Epeirotypinae. The webs of Epeirotypusand Naatlo are qualitatively indistinguishable,having many radii, no radial anastomosis, andhubs with two or more persistent hub loops (Fig-ures 67, 69). I have not seen Naatlo species build,but in Epeirotypus these loops are always addedafter sticky spiral construction. Finally, membersof both genera have tension lines that they useto distort their webs into cones (Figures 66, 69).The large number of radii must increase theforce required to distort the web; Naatlo and


Epeirotypus species have short, thick legs and"shoulders" on the carapace (Figures 46, 48, 77,86), presumably because of the increased me-chanical advantage provided and because of themuscle mass required to reel in the tension lines.The latter two features also occur in those Ther-idiosoma species that distort their webs exces-sively. A large number of radii is of course char-acteristic of symphytognathoids.

OGULNIINAE.—The subfamily includes thesingle genus Ogulnius. It is placed in its ownsubfamily because they are peculiar spiders quitedifferent from other theridiosomatids in bodyform and web architecture. Ogulnius might havebeen included in Theridiosomatinae, but placingit in its own subfamily makes each subfamilybiologically and morphologically more homoge-neous; a single subfamily, although monophy-letic, would be heterogeneous and would com-plicate discussions of theridiosomatid biology.Ogulnius is also a large and diverse genus; placingit in its own subfamily anticipates the advisabilityof splitting the genus into more manageable unitsat some time in the future. Synapomorphies de-fining the genus are listed in the systematic sec-tion.

No known species of Ogulnius spin completeorbs, merely rudimentary networks of non-stickysilk with often very short lengths of sticky silkhaphazardly draped across them (Figures 103,105). Outgroup comparison with all other theri-diosomatid genera suggests that the absence of acomplete orb is due to secondary loss, and thatimplication is confirmed by behavioral studies(Coddington, unpubl. data).

THERIDIOSOMATINAE.—This subfamily, in-cluding Baalzebub, Epilineutes, Theridiosoma, andWendilgarda, is monophyletic by numerous char-acters. First, the palp has rows or groups ofdenticles on the tegular surface beneath the con-ductor (Figures 130, 197), which are absent inother theridiosomatid genera. Second, a row ofshort, regular bristles occurs on the cymbium atits junction with the cymbial lamella (Figures153, 186, 211), also absent in other theridioso-matids. Third, the female epigyna are provided

with a pair of pits on the lateral parts of theepigynal plate (Figures 151, 173,213, but absentin Baalzebub). These pits are difficult to see ex-cept via compound microscopy of mounted epi-gyna.

Fourth, the median apophysis in all cases issomewhat elongate and has a trough or groovealong its upper surface (Figures 134, 163, 189,196). In Baalzebub, Epilineutes, and Wendilgardathe groove is deeper, whereas in Theridiosomathe groove is shallow and the median apophysisitself less rectangular and more attenuate. Fifth,the junction of the reservoir and the fundus isrugose or perhaps dentate. This feature of coursecan only be seen on cleared material with com-pound microscopy, but thus far it is believed tobe unique to these three genera. Sixth, the cop-ulatory ducts insert on the spermathecae mesally(Figures 145, 174, 215), not laterally or ventrallyas in other theridiosomatid genera. Seventh andeighth, the course of the reservoir in the tegulumhas two additional complexities, termed "switch-back 2" and "switchback 3" in Figure 1, whichare unique to these genera. Last, the eggsacs aresingly, not doubly attached, and close examina-tion of the distal tip of eggsacs indicates thatthese sacs are not doubly attached when they areconstructed, because they lack any trace of thesecond attachment line (see above for more ex-planation). These inferences have been con-firmed by observation of eggsac construction inEpilineutes globous, Theridiosoma gemmosum, andseveral other undescribed Epilineutes and Theri-diosoma species.

Ogulniinae and Theridiosomatinae are sistertaxa. They share an embolic apophysis mesal tothe embolus proper (Figures 101, 117, 162, 190,198), a basal, not distal, opening of the ejacula-tory duct (Figure 100), and the embolus itself isshort and tubular ending in a spatulate lamina,not a long, heavily sclerotized blade as in Platon-inae and Epeirotypinae. In addition the copula-tory ducts in the females are more convolutedthan the simple turns in the latter two subfami-lies.

Four characters support the monophyly of

NUMBER 422 21

Baalzebub, Epilineutes, and Wendilgarda. They allhave wrinkled, plicate copulatory bursae (Figure174, not detailed in Figures 184, 215) with lat-eral wings (Figures 174, 184, 215). The embolicdivision is divided into short, blunt elements(Figures 162, 190, 198). The median apophysishas a broad, deep trough (Figures 163, 189,196).

Wendilgarda and Epilineutes are sister taxa bythe epigynal scape (Figures 173, 213), the sub-rectangular form of the median apophysis (Fig-ures 189, 196), the denticles on its distal surface(barely visible in Figure 197), and a distal brownstripe on the conductor, apparently the result ofsclerotized thickening of that region. Baalzebublacks the derived features uniting Wendilgardaand Epilineutes and thus is the sister group tothat pair of taxa.

Wendilgarda make derived webs no longer re-sembling orbs (Figure 202, Coddington and Val-erio, 1980), but those of Epilineutes (Figure 194)and all Theridiosoma species (Figures 157, 159)are rather conventional, with few radii, a widelyspaced sticky spiral, and radial anastomosis. Thebuilding behavior of Epilineutes resembles thatof Theridiosoma species. Like Epeirotypus the lat-ter taxa also construct hub loops after stickyspiral construction, but the ensuing process ofradial anastomosis usually destroys all traces ofthe added hub loops. Baalzebub webs have asingle persistent hub loop in the web (Figures165, 167, 168), and radial anastomosis is absent,but that absence is probably due to secondaryloss.

Despite the relatively straightforward cladisticstructure explained above, the data do not re-solve the inter-relationships of Platoninae, Epei-rotypinae, and Ogulniinae-Theridiosomatinae,although possible solutions are restricted tothree, rather than 15 possibilities, because of themonophyly of Ogulniinae and Theridiosoma-tinae.

With respect to that unresolved three-taxonproblem, the evidence of individual characters isoften straightforward—for example, Plato andall ogulniine-theridiosomatines have a cymbial

lamella adjacent to the paracymbium, as well asa very small rod or tube in the wall of thereservoir, but, discordantly, the subtegulum inPlato and epeirotypines is transparent over theregion of the subtegulum. This character mayalso occur in other genera, but its appearanceseems to depend on how long the specimen hasbeen preserved. If further study invalidates thetransparent subtegulum as a reliable character,the trichotomy could well resolve in favor of(Epeirotypinae (Platoninae (Ogulniinae, Theri-diosomatinae))), a solution supported by the pres-ence of radial anastomosis in Plato and Theridio-somatinae. On the other hand, the form of thecymbial lamella in Plato (a vestige of it may existin Chthonos, although the character is not codedas present in Figure 1) does differ from that inOgulniinae-Theridiosomatinae. To repudiatethe possibility of homology between the struc-tures would nevertheless be disingenuous.

The few characters linking Chthonos and Ogul-nius are convergences, for example, the papillateand truncate sternum. Spiders of these generaare some of the smallest and most heavily scle-rotized theridiosomatids. Neither sternum char-acter is reinforced by evidence from any otherindependent character system. Likewise the pres-ence of a uniform pigment layer (i.e., no abdom-inal or cephalothoracic "pattern") in Plato andWendilgarda is also probably convergent. Speciesof other genera are often brightly and character-istically marked. Plato species usually live incaves, and the lack of abdominal pattern mayreflect their troglophilic habit, but that explana-tion does not explain the lack of pattern in thestrictly epigean Wendilgarda.

The epigyna of Platoninae, Epeirotypinae, andOgulniinae have a distinct transverse groove(Figures 25, 37, 50, 78, 124). Probably the malemedian apophysis engages this groove duringmating, although that might be the role of theparacymbium instead. If the former possibility,the presence of a scape and the absence of sucha groove in most theridiosomatines becomes in-telligible, because the female epigynal scape(functionally defined) in spiders usually engages


the male median apophysis. That explanationdoes not account for the absence of either scapeor groove in Theridiosoma, however. Symphytog-nathoids lack scapes (except, perhaps, Mysmena),grooves, and median apophyses. Either the epi-gynal groove is a primitive feature of theridio-somatids lost in the more derived genera (gainedonce, lost once), or it is convergent in Ogulniusand the Platoninae-Epeirotypinae (gained twice).Enough variability in the form and extent of theepigynal groove exists to preclude confidence ineither of these resolutions. The epigynal groovecharacter is the major evidence supporting asister group relationship between Platoninae andEpeirotypinae, one possible resolution of the tri-chotomy in Figure 1.

Finally, both Epeirotypinae and Theridioso-matinae have the tip of the mesal arm of thetegulum produced into a spur (Figures 43, 70,133, 187, 197). Ogulnius lacks such a lobe orspur altogether, but, as pointed out above, Ogul-nius is substantially autapomorphic. This tegularspur is the major evidence speaking against asister group relationship between Platoninae andOgulniinae-Theridiosomatinae.

In summary, the phylogenetic analysis of thefamily has identified seven components definingmonophyletic groups, each supported by the evi-dence of several characters, but it fails to resolvea single, and crucial, three-taxon problem—therelationships of Platoninae, Epeirotypinae, andOgulniinae-Theridiosomatinae. On the wholethe behavioral data are as equivocal as the mor-phological data. Certainly the two classic featuresof theridiosomatid webs, radial anastomosis andthe use of a tension line, do not unambiguouslydefine subsidiary monophyletic groups within thefamily. Given this discordance in both behaviorand morphology, it seems wiser to represent thecladistic relationships among the various lineagesof the family as unresolved. Perhaps closer studyof each genus in turn will reveal the solution. Iidentified this recalcitrant cladistic problem earlyin the study of the family and repeatedly tried tosolve it by restudying discordant characters andgathering more information. In the former case,

I either confirmed the initial coding of charactersor, as in the case of the homology of the cymbiallamellae of Platoninae and Ogulniinae-Theridio-somatinae, I could reach no confident resolutionof the conflict. Additional samples of informa-tion, surprisingly, yielded further sets of discor-dant characters in about equal numbers, thusimplying that the trichotomy is, in some sense, a"parametric" representation of relationship.Should the trichotomy persist, reconstructions ofcharacter evolution in the family will have to beconsidered from the point of view of each ofthree possible trees.

THERIDIOSOMATIDAE Simon

THERIDIOSOMATINI Simon, 1881:24 [type genus by mono-typy Theridiosoma O. Pickard-Cambridge, 1879].

ACTINAE McCook, 1889a: 180; 1889b: 195 [type genus ActisMcCook, 1889a: 180; genus name proposed and synony-mized with Theridiosoma in same publication].

THERIDIOSOMATEAE.—Simon, 1895:913.

THERIDIOSOMATINAE.—F.O. Pickard-Cambridge, 1902:

412.—Roewer, 1942:967.—Bonnet, 1959:4435.THERIDIOSOMATIDAE.—Vellard, 1924:132.

NOTE.—The genus name Actis, and the familygroup name Actinae based on it, are not listed inBonnet (1959), Roewer (1942), or Brignoli(1983). McCook (1889a) is a record of the pro-ceedings of a meeting of the Philadelphia Acad-emy of Natural Sciences at which McCook spokeand proposed the names (with McCook listed asauthor of the publication). It seems that McCook(1889a) does fulfill the criteria for valid publica-tion and availablity (ICZN, articles 8, 11, 12, 16),and thus ought to be listed in the above synon-ymy. McCook (1889a) did state that Actis was asynonym of Theridiosoma, but he continued touse the family group name Actinae; e.g., inMcCook (1889b), a usage conflicting with therequirement that any family group name, whenfirst proposed, must be based on a valid genusgroup name. As far as I know, however, neithername was ever used subsequently by anotherauthor writing about spiders.

DIAGNOSIS.—Theridiosomatids can be distin-guished from all other spiders by the presence of

NUMBER 422 23

pits on the prolateral margins of the sternum inboth sexes (Figure 140, but absent in Chthonos,see below), the conformation of the bulb androute of the reservoir of the male palp (e.g.,Figures 30, 115, 148, and see above discussion),and the connate spermathecae (e.g., Figure 113).The 4th tibial dorsal trichobothria (Figure 141),which are usually 2 to 4 times longer than thetibial diameter, also diagnose the family but arecomparatively short in Wendilgarda.

DESCRIPTION.—Small ecribellate, colulate, en-telegyne spiders of superfamily Araneoidea. To-tal length 0.5 to 3.0 mm (usually less than 2.5mm). Eight subequal eyes in two rows, lateralsjuxtaposed, AME separation V2 to 1 diameter,PME separation up to twice their diameter orjuxtaposed (Figures 111 vs. 172). Laterals sepa-rated from medians by about their diameters.Secondary eyes with full canoe tapeta. Seen fromabove, posterior row straight or procurved (Fig-ure 142), anterior row recurved. AME and some-times ALE, PLE on slight tubercles in adultmales. Clypeus height variable, usually more than'V'2 AME diameter. Carapace glabrous (or withfew scattered bristles), pear-shaped, length andwidth subequal, cephalic region frequently ele-vated (Figure 175). Chelicerae robust, no knownstridulatory surfaces; wide range in tooth size inindividual animals, from tiny denticles in fangfurrow (Figure 2) to larger, often bicuspid teethirregularly distributed—not usually two simplerows. Endites wider distally, inclined over la-bium. Labium indistinctly rebordered, widerthan long, sternal suture usually distinct. Ster-num about as long as wide, smooth or papillate,sparsely bristled; margin smooth or slightly in-cised, occasionally sharply truncate or even in-dented behind.

Legs in order of length 1-2-4-3 (4-1-2-3 inOgulnius), of variable proportions, short andthick in Epeirotypus, Naatlo, and Ogulnius (Fig-ures 48, 77, 109), longer and more slender inPlato, Epilineutes, and Wendilgarda (Figures 16,171, 205), intermediate in Theridiosoma andBaahebub (Figures 141, 181). Femur diameter V:<.carapace width in some genera (Epeirotypus, Naa-

tlo). Legs of adult males proportionately longerand more slender than in conspecific females.Macrosetae slender, serrate. Irregular group ofserrate setae on ventral tarsi, especially fourth,variably developed. Trichobothria grouped ontibiae and single on proximal surface of metatarsi(none on metatarsus IV or femora). Third andfourth tibiae with trichobothria in 3 or 4 rows,much longer than tibial diameter (Figures 77,141, 181). Lateral claws with 0-3 slight teethbasally, median claw prolonged beyond lateralclaw tips, attenuate, and recurved at tip (Figure3). Female palp without claw.

Abdomen smoothly ovoid (e.g., Figure 169)or with variously placed tubercles (Figures 34,47, 121) sparsely bristled, usually soft (leatheryin Chthonos), without scuta, usually higher thanlong or wide (distance from pedicel to spinneretsvery short), attached near its middle to cephalo-thorax, and to a greater or lesser extent over-hanging cephalothorax. Color uniform or withtransverse silvery or white bands, or mottledblotches sometimes merging to chevrons poste-riorly. Respiratory system diversity largely un-known (see Lamy (1902) for Theridiosoma gem-mosum), but booklung covers and single medianspiracle just anterior to colulus always present.Colulus relatively large, fleshy, bristled. Spinner-ets with 16 large spigots and numerous smallerspigots as in most araneoids (Mikulska, 1966;Wasowska, 1966, 1970; Kovoor, 1972). Anteriorspinneret in adult females (at least in Theridio-soma gemmosum) with single major ampullate andnumerous piriform gland spigots (Figure 7); me-dian with minor ampullate, one cylindrical, andseveral aciniform spigots (Figure 8), posteriorwith two aggregate, one flagelliform, two cylin-drical, and several aciniform gland spigots (Fig-ure 9). (The assignment of gland types to theexternal morphology illustrated in Figures 6-9is based on literature descriptions and has notbeen verified histologically.)

Epigynum usually a flat or domed, sclerotizedplate, plate often with central pit (Figure 37)rarely with scape protruding from beneath pos-terior margin (Figures 173, 206). Copulatory


FIGURES 2, 3.—Ogulnius gloriae from Puerto Rico: 2, cheliceral denticles on left chelicera,retrolateral; 3, elongate median claw.

FIGURE 4.—Cubical eggsac from Guatemala, IZABAL (not associated with any specimen,possibly made by Chthonos). Note opening on lower left facet.

FIGURE 5.—Mating in Wendilgarda clara from Puerto Rico; note reflected epigynal plateof female.

Stale lines: 2, 3, 10/xni; 4, 1 mm; 5, 0.5 mm.

NUMBER 422 25

FIGURES 6-9.—Theridiosoma gemmosum (L. Koch) from Alabama, spinnerets: 6, leftspinning field; 7, left anterior lateral spinneret; 8, left posterior median spinnerets; 9,left posterior lateral spinneret.

Scale lines: 10 /im.


bursae well developed (Figure 207); ducts ini-tially capacious, lightly sclerotized, abruptly nar-rowing just before junction with spermathecae(Figure 26). Coiling of ducts simple (Figure 26)or convoluted (Figure 215). One pair of sper-mathecae, always juxtaposed, connate (thus shar-ing median wall, e.g., Figure 20), short fertiliza-tion ducts exit from spermathecae laterally (Fig-ure 95). Wall of copulatory ducts adjacent tospermathecae often heavily sclerotized in narrowband (Figure 20).

Male palp with T- or hook-shaped paracym-bium (Figures 32, 164), sometimes with addi-tional cymbial lamella (Figure 211), and withbasal hematodocha, subtegulum (containing fun-dus), median hematodocha, and tegulum bearingmedian apophysis, conductor, and embolus, al-ways in consistent configuration or conformationwith respect to one another (e.g., Figure 10).Conductor usually thin, translucent plate moreor less covering embolic division. Conductor withsmall hematodocha on lateral side. Embolus asingle, long, robust sclerite (Figure 43) or shorttube with variously modified mesal apophysis,herein termed "embolic apophysis" (Figures 100,131).

NATURAL HISTORY.—Theridiosomatids livealmost exclusively in wet or humid, shaded foresthabitats. Plato is troglophilic, and Theridiosomaalso is common around cave entrances. All spe-cies seem to prefer dark situations.

Web form is diverse, ranging from completeorbs (Epeirotypus, Naatlo, Baalzebub, Figures 67,69, 165), through forms with anastomosed radii(Plato, Theridiosoma, Epilineutes, Figures 159,194), to sparse networks (Ogulnius, Wendilgarda,Figures 105, 202), or none (Chthonos). The spac-ing of the sticky spiral is nearly always greaterthan the body length of the spider (e.g., Figures105, 192), thus contrasting with that of mostother orb-weaving spiders. The web commonlyhas a line more or less at right angles to the planeof the web, herein termed the "tension line,"which the spider reels in to distort the web intoa conical form (Figures 45, 66, 157). The occur-rence of tension lines throughout the family is

sporadic (e.g., present in some Epilineutes, allTheridiosoma, Naatlo, and Epeirotypus; absent inBaalzebub, Ogulnius, Wendilgarda, and, report-edly, in Plato). Species that distort their webs viatension lines do so by gripping the hub region ofthe web with their hind four legs and reeling inthe tension line with their first four legs. Theri-diosomatids consequently have robust anteriorlegs. The slack of the tension line is either heldbeneath the spider by the pedipalps or, rarely,piled on top of the eye group. If the plane of theweb is perpendicular to the ground, the spiderusually tenses the web parallel to the ground(thus normal to the web plane), and in most casessits dorsal side up at the hub when the web istensed. The distorted position of the web repre-sents a "prey-ready" posture for the spider. In-dividuals maintain the posture for hours at atime, and most impacts of prey on the web sur-face occur with the web in a tensed position.Casual observations of Epeirotypus chavarria sug-gest that this species, at least, responds less rap-idly to prey vibrations if the impact occurs on aplanar as opposed to a distorted web. Also, theactual force in the tension line appears to varyfrom 50 to 300 dynes (roughly equivalent tomasses of 50 to 300 mg). Theridiosomatids weighno more than 3 mg, so they are exerting consid-erable force to tense their webs. The tension isall the more surprising because no morphologicalstructure that could ease the energetic cost ofthis activity has been found (for example, a cuti-cular catch mechanism at the femoral-trochan-teral joint). Webs typically occur near ground inlow vegetation or leaf litter.

Eggsacs are diverse (Figures 4?, 68, 104, 106,193, 200, 201): usually characteristic of genera,either papery or covered with flocculent silk;often retained near the web hub; cubical, sphe-roidal, pear-shaped, or fluted; with or withoutcaps or sutures at the proximal tip; and sus-pended from substrate by a long thread. Nodeliberate investigation of eggsac contents wasundertaken, but Wiehle (1931) examined 15 sacsthat each contained 20-35 eggs. Some smallOgulnius lay as few as 4 eggs per sac.

NUMBER 422 27

Prey, as far as known, is primarily nematocer-ous Diptera or other similarly weak-flying insects.The family may specialize on this prey type.

Males do not spin webs equipped with stickysilk (Wendilgarda clara is an exception, pers.obs.), but frequently they do spin the non-stickyscaffolding of the web, complete with frames,radii, hub loops, and, if characteristic of thespecies, tension lines. Males are also found "at-tending" females, i.e., on sparse silk networksjuxtaposed and often attached to the web of thefemale. The nearly ubiquitous presence of"plugs" in the copulatory bursae of females indi-cates that they mate almost immediately after thefinal molt.

Mating has been described only for Theridio-soma gemmosum (Gerhardt, 1933). Other genera(Epeirotypus, Wendilgarda, pers. obs.) seem to fol-low the same general pattern and sequence ofbehaviors. In at least the genus Epeirotypus, amuscle extends from the epigynal plate to theregion of the pedicel, presumably enabling thefemale to reflect the posterior margin of theepigynal vault during mating (see Figure 5). In-terestingly, no muscle that would serve as anadductor to clamp or close the epigynal openingseems to be present.

SPECIES.—Number of species in the family is

largely unknown. Chthonos and Plato are leastcollected, the former because it is nocturnal anddoes not spin a web, the latter because it seemsto live only in caves or cave-like situations. Prob-ably many more neotropical species of both gen-era await discovery. Ogulnius and Theridiosomaare the largest genera, both in terms of describedand undescribed species.

Wendilgarda assamensis Fage, 1924, is incertaesedis. It is a theridiosomatid, but, as Brignoli(1981) pointed out, it has little to do with Wen-dilgarda. It does not even share the synapomor-phies of Theridiosomatinae, but rather resem-bles some Plato species. Wendilgarda assamensisdoes not fit any of the generic diagnoses and,cladistically, would be placed at the bottom nodeof the cladogram in Figure 1. Rather than erecta new genus, it seems sensible to let the problemlie until more material is in hand.

RANGE.—Chiefly cosmotropical. At presentChthonos, Epilineutes, and Plato are restricted tothe Neotropics, the other genera occurring inboth the Old and New World Tropics. Few spe-cies reach the cold temperate regions (T. gemmo-sum in Europe and North America, T. epeiroidesin Korea and/or Japan). Theridiosomatids areapparently absent from western North America.

Key to the Genera of Theridiosomatidae

1. First and second tibiae, metatarsi, and tarsi with prolateral row of evenlyspaced, long, strong macrosetae [Figure 33] Chthonos

First and second legs without prolateral row of macrosetae 22. Fourth legs longer than first or, in males, subequal Ogulnius

First legs longer than fourth 33. Legs relatively short and thick [Figures 48, 77] 4

Legs long and slender [Figures 16, 141, 171, 181, 205] 54. Epigynum with closely fitting, anteriorly hinged flap covering copulatory

bursae [Figures 78, 88, 93]; male with striated embolus; tip of embolusblunt, rounded, sawtoothed [Figures 70, 71, 73, 97] Naatlo

Epigynum a domed vault, hind rim appressed to venter [Figures 50, 58];embolus smooth, tip acute [Figures 42, 43] Epeirotypus

5. PME separation 1 diameter or more [Figures 17, 208] 6PME separation '/* diameter or less [Figures 142, 172, 182] 7

6. Scape protruding from beneath entire (not notched) posterior epigynal


margin [Figures 206, 213, 219]; palp embolic division with mesalbristle protruding from beneath conductor [Figures 198, 217] (shortin mexicana only) Wendilgarda

Epigynum without scape, usually a blunt knob on a plate with a transversegroove [Figures 18, 23]; conductor with ventral apophysis [Figures 11,22] Plato

Posterior epigynal rim blunt, not pointed [Figures 143, 151], not aprotruding scape; embolic division fragmented into bristle-like parts,not blunt lobes; median apophysis attenuate, slightly grooved [Figures132, 134] Theridiosoma

Epigynum either triangular [Figures 183] or flat, medially indented platewith short scape protruding [Figure 173]; median apophysis small[Figure 163] or subrectangular in outline [Figure 188], grooved ineither case 8

Epigynal plate triangular [Figure 183]; median apophysis small but deeplycleft [Figure 163] Baalzebub

Epigynum a medially indented plate with apiculate or blunt scape pro-truding [Figure 173]; median apophysis with dorsal spur [Figure 189]

Epilineutes

PLATONINAE, new subfamily

DIAGNOSIS.—The subfamily contains the gen-era Plato and Chthonos. The following derivedfeatures define it: paracymbium a T-shaped lobe(Figures 12, 32), cymbium and distal alveolusmargin pointed or bifid median; apophysis witha long, recurved tip; conductor with a thick ven-tral apophysis (Figures 10, 29).

Platoninae may be distinguished from Epeiro-typinae by having slender, proportionatelylonger legs (Figures 16, 33, vs. 48, 77), less-compact body form, and proportionately smallermale palpi. The subfamily is distinguished fromOgulniinae by the absence of an embolic apoph-ysis encircling the male palpal bulb, and largerbody size, and from Theridiosomatinae by thesimple embolic division and grooved posteriorepigynal margin.

Plato, new genus

TYPE-SPECIES.—Plato troglodita, new species(see below).

ETYMOLOGY.—The genus name is masculineand honors the Greek philosopher in referring

to the strikingly cubical egg sacs that characterizethe genus, as well as their partiality for caves.

DIAGNOSIS.—Members of the genus Plato aredistinguished from other theridiosomatids by thefollowing combination of characters: fleshy spursextending from the lateral margins of the epi-gynal opening toward the midline (Figures 19,26), notches on the distal, mesal margin of thecymbium (Figure 10), cubical eggsacs suspendedby a long thread from one vertex (Figure 13, butsee Figure 4 and remarks under Chthonos), and astrongly curved ventral apophysis on the conduc-tor (Figure 11). The genus Chthonos has a ventralapophysis, but it is shorter and blunt. Plato lacksthe strong spines on legs I and II that character-ize Chthonos. In addition, the embolus tip in Platohas a thin, spine-like projection on the side near-est the conductor (Figures 10, 27).

DESCRIPTION.—Carapace rather low for atheridiosomatid (Figures 14, 24), pale tan headregion not especially elevated. Clypeus 3 timesA ME diameter. Eyes subequal or laterals veryslightly smaller, AME separated by 7L> their di-ameter, PME separation at least their diameter.Sternum tan or slightly darker, as wide as long,convex, rounded behind. Legs long and slender,

NUMBER 422 29

tarsi-metatarsi shorter than tibiae-patellae, paletan, no annulations. Abdomen ovoid, higherthan long or wide, sparsely setose, of uniformcolor, whitish gray or tan. Abdominal muscleapodemes not visible.

Conductor of male palp less extensive than inmore derived genera, barely covering embolusin contracted palp (Figure 11). Median apophysisvariable, basically a lobe tapering distally, fre-quently bifid, with recurved tip (Figure 10). Em-bolus single strong sclerite, duct opening distal.

NATURAL HISTORY.—Very little is known. Allspecies known to date live in caves or other darkplaces but do not seem to have marked troglob-itic modifications. The webs of P. bruneti, P.guacharo, P. miranda (pers. obs.), and an unde-scribed species from Colombia (W.G. Eberhard,pers. comm.) are loose orbs with anastomosedradii and no tension line, rather like those ofEpilineutes (Figures 192, 194).

SPECIES.—As Brignoli (1974) noticed, the spe-cies he described (1972a) in the genus Wendil-garda (miranda and guacharo) are closely relatedto Maymena bruneti Gertsch, 1960; all three spe-cies are here transferred to Plato (Plato bruneti,new combination, Plato miranda, new combina-tion, Plato guacharo, new combination). In addi-tion, Wendilgarda bicolor Keyserling has thefleshy lateral spurs on the posterior lip of theepigynal rim and therefore is also transferred toPlato (Plato bicolor, new combination). Other, asyet undescribed, species occur in the Neotropics.

RANGE.—At present the genus is exclusivelyneotropical; known from Trinidad, Venezuela,Colombia, Ecuador, and Brazil. (But see previousremarks about W. assamensis.)

Plato troglodita, new species

FIGURES 10-12, 23-28; MAP 1

TYPE.—6 holotype from Ecuador, Morona-Santiago Province, Los Tayos (78°12'W,3°06'S) in MCZ. Label states "24 July 1976,78°12'W, 3°06'S, Los Tayos, #783. Clefts bydry stream bed by main cave entrance. 20.00."

Collected by joint Ecuadorean-British Los TayosExpedition.

ETYMOLOGY.—The specific name is a femi-nine noun in apposition and means "cave-dweller" in Spanish.

DIAGNOSIS.—Plato troglodita is distinguishedfrom other Plato species in the female by thethick rounded epigynal rim with a deep centralpit in the midline and the denticulate process justanterior to it (Figure 25), and in the male by theform of the median apophysis (Figure 11) andthe tip of the embolus (Figure 10).

DESCRIPTION.—Female: Paratype collectedwith male holotype. Overall appearance as in P.bruneti (cf. Figures 14, 15). Total length 2.4 mm.Cephalothorax 1.03 mm long, 1.26 mm wide,0.89 mm high. Sternum 0.65 mm long, 0.64 mmwide. Carapace, sternum uniform yellow tan.Abdomen 1.3 mm long, 1.3 mm wide, 1.7 mmhigh; whitish tan, smoothly ovoid, sparsely setose.AME slightly smaller than PME, AME separation% their diameter, PME separation 1 diameter.ALE, PLE subequal, separated from AME, PMEby ~1 diameter. Clypeus 3.3 times AME diame-ter. Legs yellow tan, more reddish distally. Epi-gynum a smooth plate with thick posterior rim,rim with deep fossa or pit in midline and denti-culate ridge just anterior to it (Figure 25). Dorsalcleared view as in Figure 26.

Leg lengths of female described above (±0.02mm).

FemurPatellaTibiaMetatarsusTarsus

Total

11.460.521.140.810.694.62

111.240.460.930.690.653.97

III0.880.400.650.520.482.93

IV1.120.400.770.600.463.35

Male: Paratvpe, specimen #582, from samelocality as holotype. Similar in overall appearanceto female, slightly smaller (Figures 23, 24). Totallength 2.2 mm. Cephalothorax 0.95 mm wide,1.00 mm long, 0.83 mm high. Sternum 0.52 mmwide, 0.53 mm long. Abdomen 1.3 mm wide,1.2 mm long, 1.4 mm high. Clypeus 3.6 timesAME diameter. Eye proportions and spacing as


FIGURES 10-12.—Plato troglodita, new species, from Ecuador, left male palp: 10, mesal,enibolic division exposed; 1 1, ventral, enibolic division enclosed; 12, lateral.

FIGURE 13.—Plato bruneti ((iertsch) from Trinidad, eggsac.Scale lines: 100 //in, except 13, 1 mm.

NUMBER 422 31

in female, but AME projecting on a slight tuber-cle (Figure 24). Leg lengths proportionatelylonger than in female. Color of legs, carapace,sternum, and abdomen as in female. Palp as inFigures 10-12, 27, 28.

Leg lengths of male described above (±0.02mm).


Total

I1.340.451.150.830.624.39

II1.170.430.980.690.573.84

III0.860.340.840.520.463.02

IV0.980.340.760.520.453.05

VARIATION.—Females (12 specimens) range inlength from 2.2 mm to 3.1 mm, males (4 speci-mens) from 2.1 to 2.2 mm.

NATURAL HISTORY.—Mostly unknown. Labeldata indicate that the species occurs deep insidecaves (e.g., 88 m from entrance in one instanceor "bottom of main shaft" in another), as well asaround the cave entrance. One label specified"in orb web with eggsacs." Eight cubical eggsacswere present in the type series, all with a finepoint on the vertex opposite the suspension lineindicating that during construction, at least, thesac is suspended at both ends.

RANGE.—Known only from the type-locality

r

'r'] K,

MAP 1.—Distributions of Plato troglodita, new species, andP. bruneti (Gertsch).

(Map 1). Los Tayos is the name of the cavesystem. It does not appear on any map of Ecua-dor available to me. The species occurs in at least2 caves.

RECORDS.—ECUADOR, MORONA-SANTIAGO:Los Tayos caves, 7 vials from "main cave" at78° 12'W, 3°06'S, and 2 vials from "commandocave" at 78°12'W, 3°10'S; all vials with collec-tion numbers, collected by joint Ecuadorean-British Los Tayos Expedition, deposited in MCZ.

Plato bruneti (Gertsch), new combination

FIGURES 13-22; MAP 1

Maymena bruneti Gertsch, 1960:37, figs. 65-67 [$].—Brig-noli, 1974:226; 1983:377 [9 holotype from Trinidad, inAMNH, examined].

NOTE.—For evidence justifying the removalof bruneti from Maymena and its transfer to Plato,see above.

DIAGNOSIS.—P. bruneti is distinguished fromother Plato species by the distinctive epigynum,which presents a rounded tubercle on a trans-verse ridge in the female (Figure 18), and by theform of the median apophysis and the ventralconductor apophysis in the male (Figure 22).

DESCRIPTION.—Female: From Trinidad, ST.GEORGE, Simla. Total length 2.6 mm. Cephalo-thorax 1.17 mm long, 1.10 mm wide, 0.98 mmhigh. Sternum 0.65 mm long, 0.64 mm wide.Carapace, sternum, legs uniform tan. Abdomen1.5 mm long, 1.5 mm wide, 1.8 mm high; grayishwhite, smoothly ovoid, dorsum covered withstrong bristles. AME slightly larger than PME,AME separation Vi their diameter, PME separa-tion 1 diameter (Figure 17). ALE, PLEsubequal,separated from AME, PME by 1 diameter. Cly-peus height 3 times AME diameter. Epigynum asmooth plate anteriorly, produced posteriorlyinto transverse ridge with protuberant roundedtubercle medially. In posterior view, dorsal mar-gin of epigynal plate incised laterally, defining amedial lobe (Figure 19).

FIGURES 14-22.—Plato bruneti (Gertsch) from Trinidad. Female: 14, 15, habitus; 16, leftlegs; 17, eye group, frontal view; 18, epigynum, ventral; 19, same, posterior; 20, same,dorsal, cleared. Left male palp, expanded: 21, lateral; 22, mesal.

FIGURES 23-28.—Plato troglodita, new species, from Ecuador. Male: 23, 24, habitus; 25,epigynum, ventral; 26, same, dorsal, cleared. Left male palp, expanded: 27, lateral; 28,mesal.

Scale lines: 0.1 mm, except 14-16, 23-24, 0.5 mm.

NUMBER 422 33



Total

I1.430.501.170.830.654.58

II1.220.450.980.720.623.99

III0.710.400.690.520.502.82

IV1.080.400.860.600.483.42

Male: From Trinidad: ST. GEORGE, ArimaValley, Simla. Similar in overall appearance tofemale, slightly smaller. Total length wide, 1.08mm long, 0.96 mm high. Sternum 0.55 mm wide,0.57 mm long. Abdomen 1.3 mm wide, 1.5 mmlong, 1.6 mm high. Clypeus 3.3 times AME di-ameter. Eye proportions and spacing as in fe-male, but AME carried on an indistinct tubercleprojecting slightly beyond clypeus. Leg lengthsproportionately longer than in female. Color pat-tern of legs, carapace, sternum, and abdomen asin female. Palp as in Figures 21, 22.



Total

I1.380.451.120.890.554.39

II1.320.431.000.860.534.14

III0.860.340.640.690.312.84

IV0.980.310.790.580.453.11

VARIATION.—At present only 7 females and 1male are known. Females range in length from2.6 to 2.7 mm.

NATURAL HISTORY.—Like other Plato species,bruneti is apparently a troglophile. All collectionsto date are from caves. The the eggsac has thecharacteristic cubical form of the genus (Figure13). A small point on the opposite vertex fromthe suspension line of the eggsac indicates thatduring construction this end of the sac is alsoattached to supporting silk lines, as in Ogulniusand Epeirotypus, but later the lower attachmentis cut.

RANGE.—Apparently endemic to Trinidad(Map 1).

RECORDS.—TRINIDAD, ST. GEORGE, Lopi-

not (East) Cave, 220445B, P.C.J. Brunet, no date(9 type and paratypes, AMNH); Mt. El Cerro delAripo, Cave #1 (670 m), P.C.J. Brunet, no date(9, AMNH); Arima, cave nr jet. Simla Rd. andBlanchisseuse Rd., L.N. Sorkin, 23.vii.1979 (9,AMNH); Arima Valley, Simla, J.G. and B.L.Rozen, 7.iii.l968 (6\ 9, AMNH); Arima Valley,Simla, J. Rozen, 8.ii.l965 (eggsacs). ST. PATRICK,2 mi south of Fullerton, U.S. Navy base (9,AMNH).

Plato miranda (Brignoli), new combination

Wendilgarda miranda Brignoli, 1972a:375.

Plato guacharo (Brignoli), new combination

Wendilgarda guacharo Brignoli, 1972a:372.

Plato bicolor (Keyserling), new combination

Wendilgarda bicolor Keyserling, 1886:131.

Chthonos, new name

Tecmessa O. Pickard-Cambridge, 1882:433.—Bonnet,1959:4264.—Roewer, 1942:951.—Archer, 1953:6.—Brignoli, 1983:249. [Type-species by monotypy T. pecto-rosa O. Pickard-Cambridge, 1882:434, pi. 31: fig. lOa-d

(?)•]

NOTE.—The name Tecmessa O. Pickard-Cam-bridge, 1882, is preoccupied by Tecmessa Bur-meister, 1878 (Lepidoptera); the new nameChthonos is chosen to replace it.

ETYMOLOGY.—Chthonos is feminine andmeans "of the earth," referring to the habits andhabitat of this genus.

DIAGNOSIS.—Chthonos is distinguished fromother theridiosomatid genera by the lateral rowsof strong macrosetae on the first and secondtibiae, metatarsi, and tarsi; the sinuate curve ofthe first and second legs; the protuberant ster-num; the well-developed transverse ridges on theepigynum; and the form of the palp (Figures 29,32, 33, 37, 39).


FIGURES 29-32.—Chthonos sp. from Mexico, VERACRUZ, left male palp: 29, mesal, embolicdivision exposed; 30, ventral, embolic divison enclosed; 31, apical, embolic divisionexposed; 32, lateral.

Scale lines: 100 /mi.

NUMBER 422 35

DESCRIPTION.—Carapace tan or reddishbrown, cephalic region not higher than thoracicregion, clypeus 1 to 2 times AME diameter (Fig-ure 33). Sternum reddish brown, papillate, con-vex, truncate and indented posteriorly; lateralmargins deeply incised, projecting ventrally be-low level of coxae (Figure 33). AME and PMEdiameter subequal or AME somewhat larger;AME, PME separation '/a their diameter. Abdo-men color overall light yellow to reddish brown,uniform or with whitish guanine blotches; withsparse larger dark sclerotized flecks symmetri-cally arranged as well as numerous smaller punc-tate sclerotized flecks; wider above than long orhigh, tapering below to conical spinneret mound;dorsum with lateral and/or median tubercles,posterior creased; surface leathery (Figures 34-36). Spinnerets on sclerotized ring. Legs uniformbrown or light tan; patellae and tibiae sinuate;tibiae, metatarsi, and tarsi of legs I and II withdistinctive lateral single rows of long, strongspines (Figure 33). Epigynum a flat, transverselyridged plate with deeper transverse groove justanterior to posterior rim, also a dark spot or pitcentrally; transverse ridge toothed in region ofspot or pit (Figure 37). Palp with distally pointedcymbium, T-shaped paracymbium; tegulum ofonly moderate size; conductor covering embolus;embolus as one entire, strong sclerite; duct open-ing distal; median apophysis a flat plate withbackwardly directed prong (Figures 29-32). Sex-ual dimorphism slight.

NATURAL HISTORY.—Poorly known. Collec-tors have found the animals walking in leaf litteror on single dry threads between foliage (theanimal may have spun the thread, or it may haveused the drag line of another spider). ApparentlyChthonos are wandering predators that do notspin prey-catching webs. The form of the eggsacis also unknown, but both W.G. Eberhard (pers.comm.) and I have seen pink cubical eggsacs(Figure 4) much smaller than those of Plato (Fig-ure 13), unassociated with residence webs of anyspider, in habitats where Chthonos is known tooccur. In view of the other characters supportinga sister group relationship between Chthonos and

Plato (see "Intergeneric Relationships") and thecubical eggsac characteristic of Plato, thesesmaller cubical sacs may well be constructed byChthonos species. The leg macrosetae of Chthonosmay be analogous to those of mimetids. I col-lected one Chthonos individual adjacent to theweb of another spider; perhaps like mimetids,Chthonos also eats other spiders. No direct evi-dence, however, supports either of these specu-lations.

SPECIES.—Chthonos at present is an exclusivelyneotropical genus, but has been very poorly col-lected. Unsorted material from the AMNH andMCZ collections show fairly uniform genitaliaunder light microscopy but diverse combinationsof tubercles on the abdomen. The latter featuremay be a good indicator of species limits (and istaken to be so here), or may simply be intraspe-cific variation. Available series of specimens aretoo small to settle the question. Five Tecmessaspecies have been described, four properlyplaced in Chthonos: pectorosa Pickard-Cambridge,1882, peruana Keyserling, 1886 (= T. peruvianaof Bonnet, 1959:4264; unjustified emendation),tuberosa Keyserling, 1886, and quinquemucronataSimon, 1893a. The fifth, tetrabuna Archer,1958, belongs in Ogulnius (hence 0. tetrabuna(Archer), new combination).

RANGE.—AMNH and MCZ collections in-clude Chthonos individuals from the Antilles,Mexico, Guatemala, Costa Rica, Panama, Vene-zuela, and Brazil. In addition, C. peruana Key-serling, 1886, is from Peru.

Chthonos pectorosa (O. Pickard-Cambridge),new combination

FIGURES 33-38

Tecmessa pectorosa O. Pickard-Cambridge, 1882:434, pi. 31:fig. lOa-d [9].—Bonnet, 1959:4264.—Roewer,1942:951. [Holotype (?) 9 from "Mr. Traills Amazoncollection," in HDEO, examined. No locality informationin vial.]

DIAGNOSIS.—Chthonos pectorosa is distin-guished from other Chthonos species by the twopairs of lateral tubercles on the dorsum of the


abdomen (Figure 34). Judging from the figuresaccompanying the original species descriptions,it appears that peruana Keyserling (1886, pi. 11:figs. 144, 144a-c, 9) has only one pair of lateraltubercles, whereas tuberosa Keyserling (1886, pi.20: figs. 303, 303a, 9) and quinquemucronataSimon (1893a, figs. 1001-1003, 9) have anteriorand posterior median tubercles in addition to

lateral tubercles. Chthonos pectorosa lacks mediantubercles.

DESCRIPTION.—Female: Holotype. Abdo-men and cephalothorax separated. Total length1.9 mm (estimated). Cephalothorax 0.74 mmlong, 0.69 mm wide, 0.81 mm high. Sternumreddish brown, papillate, 0.46 mm long, 0.48mm wide, margins deeply incised, projecting be-

FlGURES 33-38.—Chthonos pectorosa (O. Pickard-Cambridge), type female: 33, frontal;34, dorsal; 35, abdomen, lateral; 36, same, posterior; 37, epigynum, ventral; 38, same,ventral, cleared.

FIGURES 39-41.—Chthonos sp. from Puerto Rico, left male palp: 39, ventral; 40, expanded,lateral; 41, expanded, mesal.

Scale lines: 0.1 mm, except 33-36, 0.5 mm.

NUMBER 422 37

low level of coxae in lateral view. AME and PMEdiameter subequal, AME, PME separation Vatheir diameter. Clypeus height 1.5 AME diame-ter. Abdomen 1.27 mm long, 1.82 mm wide,0.81 mm high. Overall color light yellow (prob-ably bleached) with sparse dark flecks symmetri-cally arranged (Figures 34-36), and four lateralpointed tubercles (Figure 34). Epigynum a flat,transversely ridged plate with deeper transversegroove just anterior to posterior rim; also a darkspot or pit centrally, with its transverse ridgetoothed in region of spot or pit. Legs light tan,tarsi longer than metatarsi.

Leg lengths of holotype (±0.03 mm).


Total

I0.910.260.740.400.482.79

110.720.260.600.340.462.38

III0.460.190.400.220.311.58

IV0.690.190.410.290.261.84

Male: Unknown.VARIATION.—Unknown.NATURAL HISTORY.—Unknown.RANGE.—Unknown. In his description, Pick-

ard-Cambridge cites "Amazon" as type-locality.

Chthonos peruana (Keyserling), newcombination

Tecmessa peruana Keyserling, 1886:20.

Chthonos tuberosa (Keyserling), newcombination

Tecmessa tuberosa Keyserling, 1886:252.

Chthonos quinquemucronata (Simon), newcombination

Tecmessa quinquemucronata Simon, 1893a:324.

EPEIROTYPINAE Archer

NOTE.—Archer (1953:5) included Phricotelus,Colphepeira, and Epeirotypus, with the last namedas type genus. The former two genera are not

theridiosomatids (see above, under "TaxonomicHistory"). The subfamily is here relimited toinclude Epeirotypus and Naatlo, new genus.

DIAGNOSIS.—The subfamily is defined by thefollowing derived features: the median apophysisis round with a dorsal notch (Figures 42, 55, 64,70, 82), not elongate, subrectangular, or a thinplate; the epigynum is a bulging dome (Figures50, 58, 78, 85) with a groove on the posteriorrim; the paracymbium is a broadly attachedblade; the tegulum is expanded laterally (Figures53, 72).

Epeirotypines may be recognized more easilyby the following combination of characters:larger, strongly built theridiosomatid spiders;PME separated by at least a diameter (someNaatlo species have PME separated by '/a diame-ter); short, stout legs (Figures 48, 77); humps orshoulders lateral to the fovea on the carapace;epigyna either with a closely fitting flap (Figure78) or with the posterior rim of the bulging domeclosely appressed to the abdominal wall; largepalpi (Figures 52, 61, 81) with the tegulum ex-panded laterally (Figure 54), and the emboli assingle, robust sclerites (Figures 43, 71). The webshave at least 2 hub circles, often 3 or 4, usuallymore than 12 radii, tension lines, no radial an-astomosis, and in Epeirotypus the eggsacs are at-tached to lines in the web. During the day, thespiders usually maintain their webs in the tensedposition.

Epeirotypus O. Pickard-Cambridge

Epeirotypus O. Pickard-Cambridge, 1894:134.—Bonnet,1956:1706.—Roewer, 1942:968.—Archer, 1953:5.—Brignoli, 1983:239. [Type-species by monotypy E. brevipesO. Pickard-Cambridge, 1894:134, pi. 18: figs. 11,1 Ia-d(?); 1896:161, pi. 19: figs. 14, 14a-e (<J). The name ismasculine.]

DIAGNOSIS.—Epeirotypus males are distin-guished from all other theridiosomatids by thepalp morphology: a very large tegulum, a simplereduced conductor that only partially covers thetip of the embolus, and a long smooth stoutembolus with a simple acute tip (Figures 42, 43).Chthonos and Plato have long stout emboli and


FIGURE 42.—Epeirotypus sp. from Costa Rica, PUNTARENAS, left male palp, embolic divisionexposed, mesal.

FIGURE 43.—Epeirotypus brevipes O. Pickard-Cambridge, from Guatemala, IZABAL, leftmale palp, embolic division enclosed, submesal view.

FIGURES 44, 45.—Epeirotypus chavarria, new species, from Costa Rica, HEREDIA: 44,incomplete web show ing outer sticky spiral, non-sticky "spiral" as two peripheral circles;note eggsacs doubly attached to web lines; 45, lateral view of web, showing tension line,spider ready posture, and web distortion.

Scale lines: 42, 43, 100 /xm; 44, 45, 10 mm.

NUMBER 422 39

conductors that only partially cover the embolus,but their median apophyses differ, and they donot have such short stout legs. Naatlo is verysimilar to Epeirotypus in overall appearance, butthe embolus tip of the former genus is complexand truncate (Figure 73), not simple and pointed.Some Ogulnius species superficially resembleEpeirotypus, but Ogulnius are much smaller andtheir genitalia are very different.

The epigyna of female Epeirotypus are alsodiagnostic: a strongly sclerotized vault with atransverse groove along the posterior rim, thevault closely appressed to the abdominal venter(Figures 50, 58). The ventral surface of the cop-ulatory bursae is also strongly ridged in all speciesexamined thus far (Figure 59). Naatlo epigynaare similar, but they possess a closely fitting flap.

DESCRIPTION.—Total length 1 to 3 mm. Car-apace light yellow to almost black, head regionnot elevated, lateral regions of carapace posteriorto head region strongly developed as "shoulders."Eyes subequal, AME separation ~xh their diam-eter; PME separation usually 1 diameter, occa-sionally almost juxtaposed. Sternum smooth,convex, margins slightly incised around coxae,bluntly rounded behind. Legs short and stout(Figure 48), light tan or dark and annulate, fe-mora often Vfi-V'2 carapace width (Figure 57).Tarsi with irregular group of ventral serratedhairs. Fourth tibial trichobothria 2 times tibialdiameter. Abdomen ovoid, with (Figure 47), orwithout posterior lateral tubercles (Figure 56),pointed anteriorly or rounded, usually with somedorsal color pattern, often species specific. Epi-gynum a broad vault, almost planar in front andgently rounded behind, closely appressed to ven-tral abdominal wall; posterior margin with dis-tinct, transverse groove (Figure 50). Copulatoryducts with single bend (Figure 60) like Naatlo.Male palp large; paracymbium a broadly attachedlamella (Figure 53); median apophysis, embolus,and conformation as in Figures 54, 55, 64, 65.

NATURAL HISTORY.—Habitats preferred byEpeirotypus species are shrubs or, more com-monly, tree trunks or buttresses in shaded, low-land to montane wet forest. Epeirotypus brevipes

has been taken in limestone karst hollows inGuatemala that become quite xeric by the endof the dry season. All known Epeirotypus websare alike and similar to Naatlo webs, with a ten-sion line, profuse (15-25) radii, fairly closelyspaced SS loops, and all radii inserting on thehub (lacking radial anastomosis). The hub regionhas 2-5 hub loops, which in E. brevipes and E.chavarria are constructed after the completionof the SS spiral (compare Figures 44, 67). Thenon-sticky "spiral" has two peripheral circles inall species examined thus far (Figure 44). Judgingfrom measurements made on brevipes, chavarria,and an undescribed species from Costa Rica andMexico, it appears that the adult females exert aforce about 100 to 200 times their body weightto tense the web into a cone. Eggsacs are keptnear the hub, attached to or pendant from thetension line. The sacs are covered with sparse ordense flocculent silk (Figure 68), and lack anysuture such as occurs in Theridiosoma.

SPECIES.—Bonnet (1956) and Roewer (1942)list only two Epeirotypus, the type-species E. brev-ipes and E. gloriae Petrunkevitch (1930). Epeiro-typus gloriae belongs in Ogulnius (see below).However, at least 5 other species are known fromLatin America, and possibly Malaysia, all unde-scribed.

RANGE.—Antilles, Central and South Amer-ica, possibly Malaysia.

Epeirotypus brevipes O. Pickard-Cambridge

FIGURES 43, 46-55, 68; MAP 2

Epeirotypus brevipes O. Pickard-Cambridge, 1894:134, pi.18: figs. 11, lla-d [$]; 1896:161, pi. 19: figs. 14a-e[<5].—Bonnet, 1956:1706.—Roewer, 1942:968. [Typespecimens (BMNH) lost.]

NOTE.—The original description cites speci-mens from Senahu, ALTA VERAPAZ, Guatemala,but none of the specimens in the BMNH arefrom that locality. Pickard-Cambridge's secondlisting of the species (1896:161) cites specimensfrom Teapa, TABASCO, Mexico, and some of theBMNH vials do include a Teapa label. It seems

NUMBER 422 41

that the original type series is missing, but theavailable specimens were indeed identified byCambridge. Epeirotypus brevipes is a well-markedspecies, easily identified by the abdominal colorpattern, and thus there is no doubt about thetaxon to which the name refers. Consequentlyno neotype is designated.

DIAGNOSIS.—Males and females of brevipesmay be distinguished from all other species ofEpeirotypus by the white dorsum of the abdomenwith a black dot in the middle (Figure 47). Thegenitalia of both sexes (Figures 50, 51, 54, 55),however, are similar to those of other species inthe genus.

DESCRIPTION.—Female: From Guatemala,IZABAL. Total length 2.8 mm. Cephalothorax1.26 mm long, 1.03 mm wide, 0.88 mm high;dark overall, with parallel light streaks on eitherside of midline (Figure 47). Sternum duskybrown, 0.58 mm long, 0.55 mm wide. Abdomen1.7 mm long, 1.7 mm wide, 1.9 mm high. Ab-dominal dorsum white with numerous whitesmall guanine dots and central black patch, dor-sum bordered by black; sides black, interruptedby light dots ventrally; venter dark (Figures 46,47). Abdomen flat above, with posterior lateraltubercles; anterior margin bluntly pointed (Fig-ure 47). PME 5/4 times AME diameter, separatedby slightly less than their diameter. AME sepa-rated by 74 their diameter. ALE, PLE subequalto AME, separated from AME, PME by % di-ameter. Clypeus slightly more than AME diam-eter. Posterior margin of epigynum with trans-

FIGURES 46-55.—Epeirotypus brevipes O. Pickard-Cam-bridge, from Guatemala, IZABAL. Female: 46, 47, ha-bitus; 48, left legs; 49, eye group, frontal view; 50,epigynum, ventral; 51, same, dorsal, cleared. Male: 52,habitus; 53, left male palp, dorsal; 54, same, ventral;55, mesal.

FIGURES 56-65.—Epeirotypus chavarria, new species,from Costa Rica, HEREDIA. Female: 56, 57, habitus;58, epigynum, ventral; 59, same, ventral, partiallycleared; 60, same dorsal, cleared. Male: 61, habitus;62, left male palp, expanded, sublateral; 63, same,ventral; 64, left male palp, mesal; 65, same, ventral.

Scale lines: 0.1 mm, except 46-48, 52, 56, 57, 61, 0.5mm.

verse groove, rim closely appressed to the surfaceof abdomen (Figure 50).



Total

I0.860.380.560.500.312.61

II0.730.460.540.470.302.50

III0.680.410.540.430.252.31

IV0.450.320.280.340.271.66

Male: From Guatemala, IZABAL. Total length2.2 mm. Cephalothorax 1.24 mm long, 0.95 mmwide, 0.74 mm high. Sternum 0.53 mm long,0.47 mm wide. Abdomen 1.6 mm long, 1.4 mmwide, 1.3 mm high. PME $A AME diameter,separated by slightly more than their diameter;AME separated by xh their diameter, projectingbeyond clypeus on slight tubercle (Figure 52).ALE, PLE subequal to PME, separated fromAME, PME by "A diameter. Clypeus height 2times AME diameter. Color pattern of legs, car-apace, sternum, and abdomen as in female. Legsproportionately longer, thinner than in female.Palp as in Figures 54, 55. In ventral view palpdiameter almost equal to carapace diameter.



Total

I0.900.400.610.590.392.89

II0.710.350.590.550.322.52

III0.480.270.300.370.261.68

IV0.500.370.470.430.272.04

VARIATION.—Females range in length from2.6 to 3.1 mm, males from 1.9 to 2.4 mm.Intensity of pigmentation varies from very lightto almost black.

NATURAL HISTORY.—The web resembles thatshown in Figures 66, 67. Females hang eggsacsfrom the tension line by a single thread. Whenthe eggsacs are made, however, they are doublyattached as in E. chavarria, but in the course ofweb replacement and repair, brevipes femalessever one of the connecting lines so that the


FIGURES 66, 67.—Web of Epeirotypus sp. from Guatemala, IZABAL: 66, lateral; 67, frontal,showing persistent hub loops.

FIGURE 68.—Eggsac of Epeirotypus brevipes O. Pickard-Cambridge, from Guatemala,IZABAL.

FIGURE 69.—Web of Naatlo fauna (Simon) from Panama, PANAMA, frontal view; notetension line, hub loops.

Scale lines: 10 mm, except 68, 1 mm.

NUMBER 422 43

MAP 2.—Distribution of Epeirotypus brevipes O. Pickard-Cambridge.

eggsacs are pendant (Figure 68). The speciesseems to prefer tree buttresses or rock crevicesas web sites, with the tension line facing inward.Males are found at the periphery of the femaleweb.

RANGE.—The species is recorded from Mex-ico, Guatemala, Honduras, and Costa Rica, butprobably occurs throughout Central America(Map 2).

RECORDS.—MEXICO. CHIAPAS: Palenque,Mayan ruins (d\ 99, MCZ). TABASCO: Grutas deAguas Blanca, 7 km S of Rio Tulija, 17°39'N,92°26'W (99, MCZ); Teapa (99, 6, BMNH).GUATEMALA, PETEN: Tikal, forest aroundMayan ruins (9, MCZ). ALTA VERAPAZ: Purulha(99, MCZ). IZABAL: Santo Tomas (66, 99, MCZ).HONDURAS, ATLANTIDA: 6 km S of Tela, Lan-cetilla, along base of mts. (9, MCZ). COSTARICA, HEREDIA: Puerto Viejo, Organization forTropical Studies field station Finca La Selva (66,99, MCZ).

Epeirotypus chavarria, new species

FIGURES 44, 45, 56-65

TYPE.—9 holotype and one 6, three 9 para-types from Costa Rica, Heredia Province, nearPuerto Viejo, Organization for Tropical StudiesFinca La Selva, 50 m, J. Coddington, coll.,25.iii.1979, in MCZ.

ETYMOLOGY.—The name is a noun in apposi-tion, honoring Sr. Rafael Chavarria, the late

administrator of Finca La Selva, of Puerto Viejode Sarapiqui. The ending of "chavarria" conflictswith Recommendation 31a of the ICZN, but inthis case an exception seems warranted to easepronunciation of the name.

DIAGNOSIS.—Males and females of chavarriamay be distinguished from all other species ofEpeirotypus by the color pattern: an off-whiteabdomen with numerous very white guaninespeckles. The genitalia are very similar to thoseof other Epeirotypus species.

DESCRIPTION.—Female: Paratype. Totallength 2.4 mm. Carapace whitish yellow, cephal-othorax 0.98 mm long, 0.83 mm wide, 0.86 mmhigh; white, head region slightly elevated, lateralportions strongly developed into "shoulders"(Figure 57). Sternum 0.52 mm long, 0.45 mmwide, whitish yellow. Abdomen 1.3 mm long, 1.4mm wide, 1.7 mm high; smoothly ovoid, uni-formly white with numerous white small guaninedots on dorsum and sides, absent on venter (Fig-ures 56, 57). AME slightly smaller than PME,separated by V2 their diameter. PME separatedby slightly less than their diameter. ALE, PLEsubequal to AME, separated from AME, PMEby V2 their diameter. Clypeus 4A AME diameter.Epigynum with central pit; posterior margin withtransverse groove, groove bisected medially byridge; posterior rim closely appressed to surfaceof abdomen (Figures 58, 59). Copulatory ductsridged, simple (Figures 59, 60).



Total

I0.670.320.460.380.322.15

II0.670.310.410.370.302.06

III0.500.310.320.270.201.60

IV0.630.350.440.330.191.94

Male: From type-locality. Total length 1.9mm. Cephalothorax 0.86 mm long, 0.76 mmwide, 0.83 mm high; head region elevated, lat-eral portions strongly developed into "shoulders"(Figure 61). Sternum 0.47 mm long, 0.41 mmwide. Abdomen 1.1 mm long, 1.1 mm wide, 1.3


mm high. AME slightly larger than PME, sepa-rated by xh their diameter. PME separated by 4Atheir diameter. ALE, PLE subequal, slightlysmaller than AME, separated from AME, PMEby '/> their diameter. Clypeus 3 times AME di-ameter. Color of carapace, sternum, legs, andabdomen as in female. Palp as in Figures 62-65.



Total

I0.680.330.600.470.322.40

II0.620.320.440.430.302.11

HI0.350.240.250.280.241.36

IV0.500.240.460.320.221.74

VARIATION.—Females range in length from2.0 to 2.8 mm, males from 1.4 to 2.0 mm.Coloration may be slightly yellowish in life, butnever very dark.

NATURAL HISTORY.—The species occurs onlyin primary, wet lowland rain forest, making itswebs within 2 m of the ground, on shrubs orother woody substrates. The web is shown inFigures 44,45. Females suspend eggsacs betweenthe tension line and other silk lines or the sub-strate. One female observed for about 2 monthsconstructed a total of 6 eggsacs. Eggsacs in col-lections contain from 6 to 12 eggs. The speciesseems to prefer tree buttresses, palm roots, andwind falls as web sites, with the tension line facinginward towards the substrate. Males are found atthe periphery of the female web.

RANGE.—Thus far, the species is known onlyfrom the type-locality.

RECORDS.—COSTA RICA, HEREDIA: nrPuerto Viejo de Sarapiqui, OTS Finca La Selva(66, 9$, numerous records, MCZ). The locality isthe same as the southernmost dot on Map 2, E.brevipes.

Naatlo, new genus

TYPE-SPECIES.—Naatlo sutila, new species.ETYMOLOGY.—Na'atlo is the name given by

Navaho Indians to "cat's cradle" string figures,

an art taught to them by a deity known as Spi-derwoman. The name is taken to be feminine.

DIAGNOSIS.—Naatlo is diagnosed by the pres-ence of the epigynal flap hinged anteriorly andcovering the copulatory bursa in the females(Figures 78, 88, 93) and the reduced conductorand the more complex embolic tip in the males(Figures 71, 73, 83, 97). Somatic morphologymost resembles that of Epeirotypus, but Naatlo isreadily distinguished by these diagnostic fea-tures.

DESCRIPTION.—Total lengths 1.7 to 3.0 mm.Cephalothorax glabrous, light tan, sternumslightly darker. Head region (narrower and lessdistinct in males) only somewhat higher thanthoracic region and set off by deep depressionor groove (Figure 74). Lateral region of carapaceopposite fovea high, forming "shoulders" (Figure86). AME % PME diameter, separation $A theirdiameter and similarly from ALE. PME ovoid,separation xh their diameter and similarly fromPLE. ALE, PLE juxtaposed, subequal to PME.Male eye group more compact, AME projectingslightly beyond clypeus (Figure 81). Cheliceraerobust, 2 to 3 prolateral and retrolateral, discreteteeth. Sternum longer than wide, rounded be-hind. Legs light tan proximally, darker reddishtan distally. Legs I, II much more robust thanIII, IV; female femur I width about 'A carapacediameter. Tarsi with numerous serrated bristlesventrally. Male legs I, II less robust, proportion-ately longer and thinner than in female. Abdo-men ovoid, without prominent humps. Speciesdistinguishable by abdominal shape and colorpatterns or, in doubtful specimens, by genitalicdetails.

NATURAL HISTORY.—Preferred habitat is typ-ical of the family, wet or humid shaded forest,from near sea level to montane forest (e.g., 2300m in Colombia). The webs are similar to those ofEpeirotypus, with 14-20 radii and a similar num-ber of SS loops. All radii reach the hub (Figure69). At least two hub loops and a tension line arepresent. Judging from the robust first and secondlegs, these spiders also exert considerable forceto tense their webs into a cone (cf. remarks under

NUMBER 422 45

Epeirotypus). Eggsac form apparently is diverse.One eggsac accompanying a specimen of splen-dida from Colombia was a ball of fluffy silkattached directly to a leaf. An undescribed spe-cies from Queensland, Australia, however, had apapery sack suspended from two points (cf. re-marks under Ogulnius, Epeirotypus).

SPECIES.—Theridiosoma fauna Simon, 1897b,T. sylvicola Hingston, 1932, and T. splendidum(Taczanowski, 1873) are here transferred toNaatlo. The identity of sylvicola is somewhat un-certain, because no Hingston specimens are ex-tant. He did provide a detailed description, inparticular mentioning a "subtriangular" epi-gynum (1932:376). Specimens from Guyana inthe MCZ and AMNH agree with Hingston's de-scription, and thus may be referred to sylvicola.Archer (1953:12, fig. 25) also provided a briefdescription and figure of the epigynum, as wellas identifying several specimens as T. sylvicola(currently in the AMNH). Both authors over-looked that the subtriangular structure on theepigynum is merely the ridge on the epigynalflap, not the epigynum itself. A neotype for syl-vicola should be designated when Naatlo is re-vised. Undescribed species are known fromSouth America, Australia, and New Guinea.

RANGE.—The genus occurs at least in Centraland South America, Australia, and New Guinea.

Naatlo sutila, new species

FIGURES 70-72, 74-83; MAP 3

TYPE.—9 holotype and 6 paratype from Col-ombia, Valle Province, near Saladito (1700 m).III. 1976 (W.G. Eberhard, #1064); in MCZ.

ETYMOLOGY.—The name is a Spanish adjec-tive meaning "refined" or "delicate," and refersto the overall morphology of the species in com-parison to other members of the genus.

DIAGNOSIS.—Males and females of sutila maybe distinguished from other Naatlo species by theabdomen, which is higher than long, the equa-torial, unsilvered, light band on the sides andanterior of the abdomen (incomplete behind),and the dorsal abdominal color pattern (Figures

74, 75). Naatlo splendida has a lateral silver stripeon the abdomen, and Naatlo fauna has a differentdorsal abdominal color pattern. In addition, thewidth of the epigynal flap in sutila is 0.8 or lessthe width of the sternum (Figure 76), the epi-gynal flap is less sclerotized than in splendida andfauna, and the copulatory bursae are morepointed than in splendida and fauna (Figure 80).The copulatory bursa seen when the epigynalflap is reflected (Figure 79) lacks the prominentlateral lobes characteristic of splendida andfauna. Thus far the palpal bulbs of the threespecies appear indistinguishable; abdominalcolor pattern generally will separate males.

DESCRIPTION.—Female: From Colombia,VALLE. Total length 2.0 mm. Carapace yellowtan, dusky around eyes and on carapace midline;cephalothorax 0.91 mm long, 0.69 mm wide,0.62 mm high. Sternum brownish yellow; 0.43mm long, 0.38 mm wide, acutely rounded behind(Figure 76). AME V4 PME diameter, separationslightly less than their diameter, PME separationabout xh their diameter. Clypeus height subequalto AME diameter. Abdomen 1.2 mm long, 1.1mm wide, 1.3 mm high. Dorsum of abdomendark with indistinct bilateral pattern of obliquelighter marks (Figure 75); sides and anterior withwide, unpigmented, equatorial band speckledwith sparse white dots, band incomplete behind(Figure 74); venter dark laterally, with darkpatch between spinnerets and epigynum (Figure76). Abdomen flattened above, higher than long.Legs light tan, darkening to a reddish browndistally. Epigynal flap with posterior, marginal,transverse groove (Figure 78), flap 2.1 timeswider than long, flap width 0.6 the sternumwidth. In dorsal view, copulatory bursae pointedand spermathecae four-chambered (Figure 80).

Leg lengths of a female described above (±0.02 mm).

I 11 III ivFemur 0.71 0.57 0.38 0.50Patella 0.31 0.31 0.20 0.31Tibia 0.47 0.41 0.25 0.35Metatarsus 0.47 0.44 0.30 0.36Tarsus 0.30 0.27 0.22 0.28

Total 2.26 2.00 1.35 1.80


FIGURES 70-72.—Naatlo sutila, new species, from Colombia, VALLE, left male palp: 70,submesal, embolic division exposed; 71, subapical, embolic division exposed; 72, cym-bium and paracymbium.

FIGURE 73.—Naatlo splendida (Taczanowski), from Colombia, META, left male palp, apical.Scale lines: 100 /mi.

NUMBER 422 47

Male: From Colombia, VALLE. Total length1.7 mm. Cephalothorax 0.79 mm long, 0.64 mmwide, 0.64 mm high. Sternum 0.34 mm long,0.34 mm wide. PME % AME diameter, separa-tion V4 their diameter, AME separation % theirdiameter. Clypeus height 1.8 AME diameter.Abdomen 1.0 mm long, 0.9 mm wide, 1.0 mmhigh. Color pattern of carapace, sternum, legs,and abdomen as in female. Palp with large te-gulum (Figure 83), embolic blade striated (Figure71), distal margin saw-toothed (Figure 83).



Total

I0.670.270.590.550.322.40

II0.530.270.440.500.251.99

III0.320.210.220.270.221.24

IV0.320.220.350.350.221.46

VARIATION.—Females range in length from1.6 to 2.4 mm, males (3 specimens) from 1.2 to2.0 mm. In gravid females the abdomen is occa-sionally longer than high, but the abdomen usu-ally does not project as far behind the spinneretsin lateral view as in splendida and fauna. Thecolor pattern varies in intensity of pigmentation,lighter specimens as in Figure 75, darker oneswith the light marks merging to form chevronsposteriorly.

NATURAL HISTORY.—Nothing known, butcollecting records indicate the preferred habitatis montane wet forest.

RANGE.—Panama, Colombia, and Surinam(Map 3).

RECORDS.—PANAMA. Illegible label, notmapped (99, MCZ). COLOMBIA, VALLE: nr Sa-ladito, 1700-1800 m (99, 6, MCZ); Central Hi-dalgo Anchicaya, 400 m (6, MCZ). HUILA: 18 kmE Santa Leticia, 2300 m (99, MCZ). SURINAM.SARAMACCA: Voltzberg Raleigh Nature Preserve(9, MCZ).

Naatlo splendida (Taczanowski), newcombination

FIGURES 73, 84-90; MAP 3

Theridium (Phrurolithrum) splendidum Taczanowski, 1873:5[$ holotype from "Uassa" (probably Brazil, Estado Amapa,Rio Uafa), in PAN, not examined].

Theridiosoma splendida.—Keyserling 1884:221, pi. 10: figs.134, 134a-c[9].

Theridiosoma splendidum.—Bonnet 1959:4435.—Roewer1942:970.

NOTE.—Although it has been impossible toborrow the holotype from the Polish Academy,Taczanowski described the abdominal propor-tions for Naatlo and the lateral longitudinal sil-very stripe diagnostic for splendida. In addition,Keyserling (1884) figured specimens from War-saw, presumably part of the type series.

DIAGNOSIS.—Males and females of splendidamay be distinguished from other Naatlo speciesby the longitudinal silvery stripes on the abdo-men (Figures 84-86). Naatlo sutila has a similarlateral stripe, but it lacks the silver pigment, theabdomen is usually higher than long, and theepigynal flap is narrower than the sternum width.Naatlo fauna also lacks the silver band and has adorsal abdominal color pattern, whereas the dor-sum in splendida is usually uniformly dark (Fig-ure 84). The lateral lobes inside the copulatorybursa in splendida are blunt (Figure 89), notpointed, and the bursae are indented in theirmedian sides in dorsal, cleared view (Figure 90).

DESCRIPTION.—Female: From Colombia,META. Total length 2.4 mm. Cephalothorax 0.98mm long, 0.72 mm wide, 0.69 mm high. Sternum0.46 mm long, 0.43 mm wide. Abdomen 1.51mm long, 1.08 mm wide, 1.17 mm high. AMEslightly smaller than PME, separated by SA theirdiameter. PME separation V-> their diameter.ALE, PLE subequal to PME, separation fromAME, PME '/a their diameter. Clypeus height VaAME diameter. Dorsum of abdomen uniformlydark, sides with prominent thin longitudinal sil-very stripe on dark background (Figure 86). Ven-ter light with dark patches laterally and between


9 0

NUMBER 422 49

• Naatlo sutila \Y~'1^^<V—SYVf\ ( \ ^1

A Naatlo splendida \\J f" ••«."?-'(/" 7>A ' ^ v

MAP 3.—Distributions of Naatlo sutila, new species, AT. splendida (Taczanowski), and N. fauna(Simon).

spinnerets and epigynum (Figure 85). Abdomenrounded above, longer than high. Posterior mar-gin of epigynal flap with groove laterally, medi-ally with triangular mound or ridge (Figure 88);

FIGURES 74-83.—Naatlo sutila, new species, from Col-ombia, VALLE. Female: 74-76, habitus; 77, left legs;78, epigynum, ventral; 79, same, posterior, flap re-moved; 80, same, dorsal, cleared. Male: 81, habitus;82, left male palp, mesal; 83, same apical.

FIGURES 84-90.—Naatlo splendida (Taczanowski), fromColombia, META, female: 84-86, habitus; 87, eyegroup, frontal; 88, epigynum, ventral; 89, same pos-terior, flap removed; 90, same, dorsal, cleared.


flap 2.8 wider than long, subequal to sternalwidth.



Total

I0.740.340.540.490.312.42

II0.630.310.530.440.302.21

III0.340.210.310.320.241.42

IV0.500.300.450.370.271.89

Male: From Colombia, META. Total length2.1 mm. Cephalothorax 1.07 mm long, 0.74 mm


wide, 0.65 mm high. Sternum 0.41 mm long,0.40 mm wide. Abdomen 1.1 mm long, 0.9 mmwide, 1.1 mm high. Eye group and color patternof carapace, sternum, abdomen, and legs as infemale. Abdomen more ovoid than in female.Palp morphology (Figure 73) essentially the sameas in fauna and sutila.



Total

I0.740.310.580.650.382.66

II0.610.270.530.530.342.28

III0.340.220.270.330.271.43

IV0.440.270.380.420.271.78

VARIATION.—Females range in length from2.0 to 2.7 mm, males from 1.6 to 2.1 mm.Intensity of pigmentation varies but specimensof splendida are usually darker than those ofeither sutila or fauna. The ratio of the width ofthe epigynal flap to the width of the sternum alsovaries, and overlaps completely that in fauna, butit is never so small as that in sutila.

NATURAL HISTORY.—This species apparentlyoccurs in lowland as well as cloud forest. Oneweb (photograph supplied by W.G. Eberhard)had 2 hub loops, 19 radii, no radial anastomosis,and a tension line at an angle to the plane of theweb. One eggsac was made of fluffy silk and wasattached to a leaf, rather than to the tension lineor suspended by a silk line. That form of eggsacwould be unique among theridiosomatids and,indeed, among symphytognathoids; perhaps theattribution of the eggsac to splendida was a mis-take.

RANGE.—Colombia, French Guiana, Peru,Bolivia, and Brazil (Map 3).

RECORDS—COLOMBIA, META: 15 km SWPuerto Lopez, Hacienda Mozambique 200-500m (9$, 66, numerous records, MCZ). FRENCHGUIANA. GUYANA: Cayenne (9, AMNH).PERU, UCAYALI: Huanuco, Divisoria (probablyon Ucayali border between Tingo Maria andPucallpa) (99, AMNH). HUANUCO: Tingo Maria

(99, AMNH). BOLIVIA, BENI: Rio Benicito (9?,AMNH); Rio Benicito, Chacobo Indian village(9, AMNH). BRAZIL, AMAPA: Rio Uaca (9,PAN?).

Naatlo fauna (Simon), new combination

FIGURES 69, 91-98; MAP 3

Theridiosoma fauna Simon, 1897a:483, figs. 1, 2, 7, 8 [websand eggsacs].—Bonnet, 1959:4433.—Roewer, 1942:970.[A syntype series of 1 male and 19 females from SanEsteban, CARABOBO, Venezuela, in MNHN, Paris, exam-ined. Female lectotype here designated.]

DIAGNOSIS.—Males and females of fauna maybe distinguished from other Naatlo species by theabdominal color pattern (Figures 91, 92). Naatlosplendida has a lateral silver stripe on the abdo-men, and no paired spots on the dorsum. Theepigynal flap of sutila is narrower than the ster-num, whereas in fauna it is subequal or wider.The lateral lobes in the copulatory bursa arepointed, not blunt (Figure 94).

DESCRIPTION.—Female: From Colombia,VALLE. Total length 2.2 mm. Cephalothorax0.89 mm long, 0.77 mm wide, 0.77 mm high,head region not markedly elevated, carapace op-posite fovea higher, forming "shoulders" (Figure92). Sternum 0.50 mm long, 0.45 mm wide.AME slightly less than PME diameter, separatedby % their diameter. PME separation Vs theirdiameter. ALE, PLE slightly larger than AME,separated from AME, PME by xh their diameter.Clypeus height 2 times AME diameter. Abdomen1.3 mm long, 1.2 mm wide, 1.1 mm high. Dor-sum of abdomen dusky with distinct large mar-ginal light blotches, blotches paired or formingtransverse bands posteriorly (Figure 91); sidesdark, venter dark laterally, dark patch betweenspinnerets and epigynum. Abdomen roundedabove, longer than high, extending posteriorlyfar beyond spinnerets (Figure 92). Epigynal flapwith posterior transverse groove (Figure 93), flap2.6 times wider than long, subequal to sternalwidth. Bursae rounded (Figure 95).

NUMBER 422 51

FIGURES 91-98.—Naatlo fauna (Simon), from Colombia, VALLE. Female: 91, 92, habitus;93, epigynum, ventral; 94, same, posterior, flap removed; 95, same, dorsal, cleared.Left male palp, expanded: 96, lateral; 97, ventral; 98, apical, showing reservoir route.

Scale lines: 0.1 mm, except 91, 92, 0.5 mm.



Total

10.830.350.610.560.362.71

II0.700.310.540.490.362.40

III0.380.200.270.300.241.39

IV0.830.280.450.410.242.21

Male: From Colombia, VALLE. Total length2.2 mm. Cephalothorax 0.91 mm long, 0.72 mmwide, 0.74 mm high, head region not markedlyelevated, carapace lateral to fovea higher, form-ing "shoulders." Sternum 0.44 mm long, 0.41mm wide. Eye group as in female, clypeus height

% AME diameter. Abdomen 1.3 mm long, 1.1mm wide, 1.2 mm high. Color of carapace, ster-num, abdomen and legs as in female. Abdomenmore ovoid in lateral view than in female. Unex-panded palp as in other members of the genus,expanded as in Figures 96-98.



Total

I0.900.330.670.720.383.00

II0.750.280.570.590.332.52

III0.400.210.270.340.271.49

IV0.550.260.440.430.281.96


VARIATION.—Females range in length from2.3 to 2.9 mm, males from 1.7 to 2.2 mm. Thecolor pattern varies in intensity of pigmentation,and in lighter specimens the diagnostic lightblotches may be open laterally. Females fromPanama are very light, and the abdomen dorsumis almost a uniform dusky color. Naatlo splendidaand N. fauna overlap in many characters, butsplendida always has silvery stripes on the sidesof the abdomen.

NATURAL HISTORY.—The species occurs overa wide altitudinal range (50-1800 m), apparentlyin wet forest or cloud forest. Web (Figure 69)with numerous radii, hub loops, no radial anas-tomosis, and a tension line.

RANGE.—Costa Rica, Panama, Colombia,Venezuela, Trinidad, Ecuador, and Brazil (Map3).

RECORDS.—COSTA RICA, SAN JOSE: Qui-zarra, 9 km SW San Isidro del General, 2500 m(9, MCZ). PANAMA. PANAMA: Lake Gatun,Barro Colorado Island (99, MCZ). COLOMBIA.VALLE: 28 km E of Buenaventura, nr Saladito,1800 m (99, 66, numerous records, MCZ); RioSan Juan, tributary of Rio Dagua nr Queremal,1300 m (9, MCZ); above Atuncela, 1800 m ($9,MCZ); Central Hidalgo Anchicaya, 400 m (9,MCZ). VENEZUELA, ARAGUA: Chorom (9,AMNH). CARABOBO: San Esteban (99, 6,MNHN). TRINIDAD, ST. GEORGE: Arima Rd.,gap to Blanchisseuse (9, AMNH). ECUADOR.PICHINCHA: Rio Palenque, 47 km SW SantoDomingo, on road to Queredo (99, MCZ). BRA-ZIL, RIO DE JANEIRO: Teresopolis, (6, MCZ).

Naatlo sylvicola (Hingston), new combination

Theridiosoma sylvicola Hingston, 1932:376.

OGULNIINAE, new subfamily

DIAGNOSIS.—Ogulniinae contains only the ge-nus Ogulnius and is diagnosed by the same char-acters diagnosing Ogulnius (see below).

Ogulnius O. Pickard-Cambridge

Ogulnius O. Pickard-Cambridge, 1882:432.—Bonnet,1958:3157.—Roewer, 1942:968.—Archer, 1953:7.—Brignoli, 1983:239. [Type-species by monotypy 0. obtectusO. Pickard-Cambridge, 1882:433, fig. 9a-d [9]. The nameis masculine.]

DIAGNOSIS.—Ogulnius can generally be rec-ognized by body proportions: the fourth legs arelonger than the first (Figures 109, 122 in females;some males with LI, LIV subequal); their smallsize; the abdomen greatly overbalancing cephal-othorax and even obscuring the eye group indorsal view (Figures 107, 120). The sternum ismarkedly truncate behind, often incised. In theembolic division of the male palp the mesalapophysis is forked, and its lateral arm is pro-longed into a long tapering bristle that encirclesthe palp (Figure 101). The epigynum resemblesEpeirotypus, Plato, or Chthonos in having a trans-verse groove sometimes interrupted by a longi-tudinal median ridge on the posterior margin(Figures 112, 124), but the vault is low and flat,not domed as in Epeirotypus.

DESCRIPTION.—Total length 0.5 to 2 mm(rarely to 2.4 mm). Carapace longer than wide,cephalic area much elevated; dark brown to lightyellow-tan, or with light central area and periph-ery marked by characteristic dark figure (Figure110). AME smaller than or subequal to PME,separated by V2 their diameter, PME separatedby at least V* diameter, never juxtaposed (Figures111, 123). Clypeus usually 2 times AME diame-ter. Sternum smooth or papillate, tan to darkreddish brown, sparsely bristled, always truncatebehind, often incised. Legs yellow-tan or annu-late dark and light, short and stout. Fourth legslongest, relatively slender, tibial trichobothriaquite long. First femur diameter '/> sternumwidth. Abdomen smoothly ovoid or with anteriorand/or posterior tubercles, higher than long orwide, often wider than long, always overhangingcarapace; light yellow to dark reddish brown.

Epigynum a flat vault with posterior marginalgroove. No scape, lateral spurs, or lateral pits.Interior ducts convoluted (Figure 113). Palp as

NUMBER 422 53

FIGURES 99-102.—Ogulnius gloriae (Petrunkevitch), from Puerto Rico, left male palp,embolic division exposed: 99, mesal; 100, apical; 101, sublateral; 102, cymbial lamellaand paracymbiuni.

Scale lines: 100 /mi.


FIGURES 103, 104.—Ogulnius sp. from Guatemala, ALTA VERAPAZ: 103, web; note threedimensional trajectories of radii, sticky spiral; 104, eggsac.

FIGURES 105, 106.—Ogulnius gloriae (Petrunkevitch), from Puerto Rico: 105, web; notetwo-dimensional trajectories of radii, sticky spiral; spider at upper center; 106, eggsac.

Scale lines: 103, 105, 10 mm; 104, 106, 1 mm.

NUMBER 422 55

in Figures 99-102, median apophysis a relativelyextensive thin plate, often elongate, with one ortwo apophyses. Routing of reservoir simpler thanin other genera (Figures 118, 119).

NATURAL HISTORY.—Ogulnius prefer verywet habitats; they usually require a water-satu-rated humid environment. Usually found within0.2 m of ground, often in leaf litter, web attach-ment sites tend to be stationary and inflexible,such as stones, thick branches, or plant stemforks. All species observed construct simple,sparse, non-sticky silk networks on which the SSspiral winds in an irregular trajectory (Figures103, 105), only vaguely resembling orbs. Framelines absent, radii always attached directly tosubstrate, joining each other toward hub in anirregular series of trivalent nodes. Overall shapeof web may be planar (Figure 105) or threedimensional (Figure 103). Dimensionality of non-sticky scaffolding strongly influences regularityof SS spiral. Spiders sit at the center of the webor at the end of a radius, face away from hub,and usually tense that radius slightly.

Eggsacs various, either double pointed penta-gons or hexagons (Figure 104) or pendant tear-drop-shaped leathery sacs (Figure 106). If theformer, usually attached to web at both ends ofsac, if the latter, suspended from a single silkline. Pendant eggsacs have a minute point oppo-site the suspension line, indicating that even pen-dant eggsacs are doubly attached when con-structed (Figure 106).

Prey, as far as known, are nematocerous fliesor other weak flying ectomorphic insects, oftenmuch larger than the spider (e.g., tipulids). Gen-eral behavior of spider unusually sluggish andslow, never exhibiting the rapid attack behaviorcharacteristic of other genera. Probably rely onextremely viscid, slack silk lines to catch long-legged, relatively weak prey that cannot escapeonce trapped. As in other genera, males oftenremain adjacent to female webs in their ownsmall, non-sticky tangles.

SPECIES.—Ogulnius is at present second toTheridiosoma in numbers of species. It also iscircumtropical, diverse, and undercollected.

About 10-15 species, described and unde-scribed, are known from the Neotropics, someapparently endemic to various Antillean islands.The Old World fauna is also diverse. Species aredifficult to distinguish because of small size andindividual variation (Figures 99-100). Epeiroty-pus gloriae Petrunkevitch (1930) (see below) andTecmessa tetrabuna Archer (1958) belong in Ogul-nius. The following Ogulnius species also seemproperly placed in the genus: agnoscus Strand,1918, pullus Bosenberg and Strand, 1906, clarusKeyserling, 1886 (type examined), cubanusArcher, 1958 (type examined), infumatus Simon,1897 (type examined), obscurus Keyserling, 1886(type examined), pallisteri Archer, 1953 (typeexamined), gertschi Archer, 1953 (type exam-ined), fulvus Bryant, 1945a (type examined), la-tus Bryant, 1948 (type examined), and yaginumaiBrignoli, 1981. Some of the taxa may, however,be synonyms of each other; the matter needsfurther research.

RANGE.—Circumtropical. Ogulnius pullus ap-parently extends its range into north temperateAsia.

Ogulnius obtectus O. Pickard-Cambridge

FIGURES 107-119; MAP 4

Ogulnius obtectus O. Pickard-Cambridge, 1882:433, figs. 9,9a-d [$].—Bonnet, 1956:1706.—Roewer, 1942:968.—Brignoli, 1981:12-13, figs. 1-2 [lectotype $]. [Lectotype9 and paralectoty pe 9 designated by Brignoli (1981) fromBrazil, "Amazon," in HDEO, examined.]

DIAGNOSIS.—Ogulnius obtectus may be recog-nized by the combination of lateral dark markson the margin of the carapace (Figures 107, 110),the PME separated by twice their diameter (Fig-ure 111), and the absence of posterior lateraltubercles on the abdomen. Ogulnius tetrabunahas the lateral dark marks, but the PME arecloser together and the abdomen has posteriorlateral tubercles. Ogulnius gloriae may have amottled carapace, but the pattern is indistinct,and the background color of the carapace ismuch darker. In addition, gloriae also has poste-rior lateral tubercles, but obtectus does not.


FIGURES 107-119.—Ogulnius obtectus O. Pickard-Cambridge. Female: 107, 108, habitus;109, left legs; 110, carapace; 111, eye group, frontal view; 112, epigynum, ventral;1 13, same, dorsal, cleared. Male: 114, habitus; 115, left palp, mesal; 116, same, lateral;1 17, same, ventral; 1 18, same expanded, mesal; 1 19, same, lateral.

Scale lines: 0.1 mm, except 107-1 10, 0.5 mm.

NUMBER 422 57

DESCRIPTION.—Female: From Colombia.Total length 1.5 mm. Cephalothorax 0.71 mmlong, 0.87 mm wide, 0.55 mm high; light yellow,mottled around periphery (Figures 107, 110).PME % AME diameter, PME separation twicetheir diameter, AME separation 'A their diame-ter. ALE, PLE % PME diameter, separated bytheir diameter from AME, PME. Clypeus height1.2 AME diameter. Sternum 0.32 mm long, 0.37mm wide, reddish brown, smooth, evenly trun-cate behind. Abdomen large for cephalothoraxsize, 1.2 mm long, 1.3 mm wide, 1.3 mm high;greatly overhanging carapace, almost obscuringeyes (Figure 107). Abdominal dorsum evenlyreddish brown. Legs short, thick, yellowishbrown, tibia I with retrolateral distal light patch,4th leg longer than 1st.



Total

10.360.170.190.200.171.09

II0.350.180.200.170.181.08

HI0.300.160.180.200.161.00

IV0.490.210.260.270.221.45

Male: From Peru, UCAYALI. Total length 1.2mm. Cephalothorax 0.64 mm long, 0.59 mmwide, 0.60 mm high. Eye proportions as in fe-male. Clypeus height 1.7 AME diameter. Ster-num 0.38 mm long, 0.35 mm wide. Abdomen1.0 mm long, 1.0 mm wide, 0.9 mm high. Ab-dominal shape as in female, with prolateral lightstreak (Figure 114). Color pattern of carapace,sternum, and legs as in female. Legs I, IV sube-qual in length. Palp as in Figures 115-119.



Total

I0.390.160.260.250.201.26

II0.370.160.220.250.211.21

III0.250.130.170.200.190.94

IV0.400.160.250.240.211.26

VARIATION.—Females range in lengths from1.3 to 1.6 mm, males from 0.9 to 1.2 mm (two

specimens). Abdominal coloration varies from auniform reddish brown to forms with bilaterallight patches defining a central dark line on theabdomen. All specimens have the characteristicpattern on the carapace. Archer (1953:19) statedthat obtectus "has a series of pits on either side ofthe carapace." Neither the type series nor anyother specimens referable to obtectus have pits onthe carapace; presumably Archer mistook pig-ment for pits.

NATURAL HISTORY.—Nothing known. Col-lecting labels said "beaten from vegetation."

RANGE.—Colombia, Peru, and Brazil (Map 4).RECORDS.—COLOMBIA, MAGDALENA: Sierra

Nevada de Santa Marta, San Pedro-San Javier,1500 m (JAK)- PUTUMAYO: Rio Putumayo, nrPuerto Asis (probably 00°30'S, 76°31'W) (9,MCZ). PERU, UCAYALI: "Colonia Calleria, RioCalleria," 15 km from Ucayali (9, AMNH) (prob-ably Callaria at 8°08'S, 74°35'W); Huanuco,Divisoria (probably on UCAYALI border betweenTingo Maria and Pucallpa 9°05'S, 75°46'W) (9,AMNH). cuzco: Machu Picchu above ruins,2060 m (99, 6, AMNH); Torentoy Canyon, baseof Machu Picchu 2000-2200 m (probably To-rentoy, 15°10'S, 72°30'W) (9, AMNH). BRA-ZIL, AMAZONAS: exact locality unknown, notmapped.

NOTE.—The type series is from Brazil, but thelocality is inexact ("Amazon"). Occurrence ofother individuals in southeastern Colombia andnortheastern Peru suggests that Mr. Traill col-lected the animal at the headwaters of the Ama-zon rather than farther downstream. The speciescovers a wide altitudinal range (150-2600 m),and abdominal color pattern varies considerably,but throughout this diversity the female genitaliaand carapace color pattern remain constant.

Ogulnius gloriae (Petrunkevitch), newcombination

FIGURKS 99-102, 105, 106, 120-129; MAP 4

Epeirotypus gloriae Petrunkevitch, 1930:307, figs. 180-183[?], fig. 184 [web].—Bonnet, 1956:1706.—Roewer,1942:968. [Holotype 9 from Ponce. Coamo Springs (nrCoanio), Puerto Rico, in PMNH, examined.]


Ogulnius obtectus

MAP 4.—Distributions of Ogulnius gloriae (Petrunkevitch) and 0. obtectus O. Pickard-Cam-bridge.

DIAGNOSIS.—Ogulnius gloriae uniquely pos-sesses three closely grouped tubercles (two above,one below) on the posterior of the abdomen(Figures 120, 121).

DESCRIPTION.—Female: From Puerto Rico,HUMACAO. Total length 1.9 mm. Cephalothorax0.89 mm long, 0.65 mm wide, 0.69 mm high;

chestnut brown, mottled around periphery, elon-gate ridge between PME (Figures 120, 121, 123).PME slightly smaller than AME, PME separationslightly more than their diameter, AME separa-tion 'A their diameter. ALE, PLE % PME diam-eter, separated by 'A' their diameter from AME,PME. Clypeus height 1.5 AME diameter. Ster-

NUMBER 422 59

FIGURES 120-129.—Ogulnius gloriae (Petrunkevitch), from Puerto Rico. Female: 120,121, habitus; 122, left legs; 123, eye group, frontal view; 124, epigynum, ventral; 125,same, dorsal, cleared. Left male palp: 126, sublateral; 127, lateral; 128, expanded,lateral; 129, expanded, subniesal.

Scale lines: 0.1 mm, except 120-122, 0.5 mm.


num 0.38 mm long, 0.38 mm wide, reddishbrown, papillate, sharply truncate and notchedbehind. Abdomen very large for cephalothoraxsize, 1.7 mm long, 1.2 mm wide, 1.6 mm high;greatly overhanging carapace, almost obscuringeyes (Figure 120). Abdominal dorsum mottled,light flecks on dark brown background; two largeanterior lateral tubercles, two indistinct posteriorlateral tubercles, and three grouped small tuber-cles on posterior midline (Figures 120, 121).Legs short, thick (1st femur diameter ~'/2 ster-num width), annulate with dark and light bands,4th leg longer than 1st (Figure 122).



Total

I0.490.270.270.210.211.45

II0.400.270.250.300.211.43

III0.340.210.220.250.191.21

IV0.560.260.320.280.211.63

Male: From Puerto Rico, HUMACAO. Totallength 1.2 mm. Cephalothorax 0.64 mm long,0.55 mm wide, 0.55 mm high; cephalic ridge asin female. PME 3/» AME diameter, separation 1diameter, AME separation lh their diameter.ALE, PLE % PME diameter, separated by Vatheir diameter from AME, PME. Clypeus height1.5 AME diameter. Sternum 0.34 mm long, 0.34mm wide. Abdomen 0.7 mm long, 0.8 mm wide,1.0 mm high. Color of carapace, sternum, ab-domen and legs as in female. Leg lengths 1-2-4-3, proportionately longer and more slender thanin female. Palp as in Figures 127-129.



Total

10.410.210.280.220.191.31

110.370.200.270.210.191.24

III0.270.160.210.150.170.96

IV0.380.180.240.180.131.11

VARIATION.—Females range in length from1.6 to 2.4 mm, males from 0.9 to 1.2 mm. Color

tan to almost black. Posterior lateral abdominaltubercles indistinct in some specimens.

NATURAL HISTORY.—The species occurs inhumid shaded forest habitats or similar microcli-mates (for example, between shaded window barsor under roadside ferns). It seems less restrictedto deep primary forest habitats than is typical oftheridiosomatids. The web (Figure 105) lacks ahub, the number of radii rarely exceeds 10, andthe sticky spiral spacing much exceeds the bodylength of the spider. The web is roughly planarand oriented vertically. Although the radii ap-pear "anastomosed" as in those of Theridiosomaor Epilineutes, the anastomosis arises because theanimals never make a well-formed hub duringthe construction of the non-sticky scaffolding.Thus not only does "radial anastomosis" in Ogul-nius occur before, rather than after sticky spiralconstruction (in contrast to Theridiosoma and Epi-lineutes), but the anastomosis also occurs in a verydifferent way. The two aspects of web architec-ture are only superficially similar in the finalproduct, not homologous. The spider usually sitsat the periphery of one of the upper radii, facesaway from the web center, and slightly tensesthat radius.

The eggsac is hard, leathery, pear-shaped, andsuspended from nearby vegetation by a long silkline whose end nearer the sac is thickened. Spi-derlings apparently escape through a hole theymake on the side of the sac. A slight point op-posite the suspension line (Figure 106) indicatesthat during construction eggsacs are doubly at-tached, and the lower end later cut free.

Prey items collected from webs in a windowframe comprised mostly tipulids whose bodylength and certainly leg span much exceeded thatof the spiders or the sticky spiral spacing of theirwebs. Forest spiders caught mainly midges andmosquitos.

RANGE.—Apparently endemic to Puerto Rico(Map 4), although similar, undescribed speciesoccur on Cuba, Hispaniola, and Jamaica.

RECORDS—PUERTO RICO. GUAYAMA:Cidra (immature, AMNH). HUMACAO: La Gloria,Luquillo Range (9 paratype, YALE); El Yunque

NUMBER 422 61

(99, 6, MCZ, AMNH); Rio Piedras (9, AMNH);Luquillo Forest, aviary (99, 66, MCZ); trail tosummit of El Toro (99, 6, MCZ); Rt. 191, road-side (99, MCZ). MAYAGUEZ: Bosque Estado deMaricao, 2400' (66, AMNH); Mayaguez (o\AMNH); Mayaguez, University campus, PONCE:

Coamo Springs Hotel, on plants near spring (9,holotype, PMNH).

Ogulnius tetrabuna (Archer), new combination

Tecmessa tetrabuna Archer, 1965:130.

THERIDIOSOMATINAE Simon

DIAGNOSIS.—Theridiosomatinae includesBaalzebub, Epilineutes, Theridiosoma, and Wendil-garda, and is diagnosed by the following derivedcharacters: embolic division divided into bristle-like parts or broad, blunt apophyses (Figures 131,136, 162, 190, 198); median apophysis with dor-sal groove or trough (Figures 135, 163, 189,196); denticles on tegular margin beneath distallip of conductor (Figures 130, 135, 197); row ofshort, uniform bristles on cymbial margin oppo-site cymbial lamella (Figures 147, 153,186,211);epigynum with lateral pits (Figures 151, 173,213, but absent in Baalzebub); routing of reser-voir inside tegulum more complex than that ofother genera (Figures 146, 147); junction ofreservoir and fundus dentate.

Theridiosoma O. Pickard-Cambridge

Theridiosoma O. Pickard-Cambridge, 1879:193.—Bonnet,1959:4432.—Roewer, 1942:969.—Archer, 1953:6.—Brignoli, 1983:240. [Type-species by monotypy T. argen-teolum O. Pickard-Cambridge, 1879:194, pi. 12: fig. 8a-g ($) (=T. gemmosum (L. Koch)). Female from Bloxworth,Dorset, England, in BMNH, apparently lost. The name isneuter.]

Microepeira Emerton, 1884:320, pi. 34: fig. 7 [?], pi. 38: figs.1 [<5], 2-4 [9] [type-species by monotypy Epeira radiosaMcCook, 1881; objective synonym of Theridisoma ra-diosum (McCook) (= T. gemmosum (L. Koch))].

Actis McCook, 1889a: 180 [type-species by monotypy Epeiraradiosa McCook, 1881 (see above); objective synonym

of Theridiosoma radiosa (McCook) (= T. gemmosum(L. Koch))].

Andasta Simon, 1895:918 [type-species by monotypy A. sem-iargentea Simon; female from Ceylon, in MHNH, Paris,examined]. [New synonymy.]

NOTE.—Andasta semiargentea does not differsubstantially from the generic diagnosis givenbelow, hence the generic name is considered ajunior synonym of Theridiosoma. Without doubtsome species groups of Theridiosoma will meritgeneric status, but much more research is re-quired to assure the monophyly of those groups.Until the cladistic relationships within the genusare better understood, it seems best to adopt abroad definition.

DIAGNOSIS.—Theridiosoma differs from allother theridiosomatids by the form of the em-bolic division—a short, tubular embolus with theembolic apophysis fragmented into several longbristle-like parts (Figures 131, 133, 137). Theembolus of Plato, Chthonos, Epeirotypus, andNaatlo are strongly sclerotized tubes longer thanthose of Theridiosoma, and they have no embolicapophyses. The embolic division of Ogulnius ismore similar to that of Theridiosoma in having anembolic apophysis, but it is at most bifid, notfragmented, and much longer.

The embolic divisions of Baalzebub, Epili-neutes, and Wendilgarda also have a short tubularembolus and embolic apophyses divided distally,but the apophyses are broad and blunt, three-parted, and not fragmented into many bristle-like parts. The embolic division is normally hid-den beneath the conductor, but its form—whether broad and blunt or fragmented andbristle-like—can be discerned without dissectionbecause the conductor is usually translucent (cf.Figures 149, 155, 178, 187).

In addition, the median apophysis of Theridio-soma is consistently a curved lobe attenuate dis-tally with a slight groove along its upper longaxis (Figures 130, 132, 135). The median apo-physes of Baalzebub, Epilineutes, and Wendilgardaalso are grooved along their upper surfaces, butthe groove is deeper. The median apophyses ofEpilineutes and Wendilgarda are square or sub-


FIGURES 130, 131.—Theridiosoma goodnightorum Archer, from Costa Rica, HEREDIA, leftpalp: 130, ventral, embolic division enclosed; 131, apical, embolic division exposed.

FIGURES 132, 133.—Theridiosoma sp. from Costa Rica, LIMON, left palp: 132, mesal,embolic division enclosed; 133, ventral, embolic division exposed.

Scale lines: 100 /xm.

NUMBER 422 63

rectangular and they do not taper to a fine pointas does that of Theridiosoma. The median apoph-ysis of Baalzebub somewhat resembles that ofTheridiosoma, but it is more heavily sclerotized,deeply cleft, and not so attenuate.

Female Theridiosoma have relatively firm, ro-bust copulatory ducts leading from the bursa tothe spermathecae (Figures 145, 152). Ducts ofBaalzebub, Epilineutes, and Wendilgarda are deli-cate, plicate, and usually collapsed, and so thelumen of the duct is difficult to discern. Thelatter genera also have some sort of functionalscape—either a spiniform or blunt projection ofthe ventral epigynal wall, or the entire wall isprolonged posteriorly into a triangle.

The posterior median eyes of Theridiosomaspecies arejuxtaposed, nearly or in fact touching.The PME of Baalzebub and Epilineutes are alsojuxtaposed, and the genera cannot be separatedby this character. Some Naatlo and Epeirotypusspecies have PME separated by as little as 'Adiameter, but these species have "shoulders" lat-eral to the fovea, and their legs are shorter andthicker.

DESCRIPTION.—Total length 0.5 to 2.5 mm.Carapace dark brown to very light tan, occasion-ally with distinctive marks around eye region orfovea. Eyes nearly subequal, AME separated byabout V'2 a diameter, PME separated by at most'A» diameter (Figure 142). Chelicerae typical offamily. Sternum smooth, sparsely bristled, usu-ally with a dusky margin (Figure 140). Legs shortor long, metatarsi usually shorter than tibiae(except leg III), uniform tan or segments annu-late distally. Trichobothria as in family descrip-tion, 4th trichobothria often 3-4 times tibialdiameter.

Abdomen ovoid, higher than long or broad,dorsum very often with transverse thin light bandand or symmetric light spots. Sides light gradingto dark ventrally, venter dark.

Epigyna often with lateral sclerotized spursprojecting toward the median (T. savannum, T.nechodomae, T. davisi, T. argenteolunulatum, notillustrated), or a simple, flat or domed vault(Figures 143, 151). No scape or transverse

groove on posterior epigynal lip. At least somespecies with lateral pits or depressions (Figure151) in the epigynal plate. Copulatory duct rout-ing moderately complex (Figures 145, 152).

Sexual dimorphism slight, males usually simi-lar to female in color pattern, slightly smaller instature, legs proportionately longer. The portionof the tegulum beneath the distal point of theconductor usually is a denticulate, smoothlycurved surface (Figures 130, 133, 135).

NATURAL HISTORY.—Habitats preferred byTheridiosoma species are typical for the family,i.e., wet, shaded forest understory. All speciesknown to date construct simple orbs with theradii joined together, or anastomosed, beforethey reach the hub. The 4-5 radial lines that doreach the hub are referred to as "hub radii." Thespider anastomoses radial lines and constructsthe hub radii after completion of the sticky spiral.The web usually has a tension line approximatelynormal to the plane of the orb, and by reeling itin the spider distorts the web into a cone (Figure157). In a typical prey-ready posture the spiderfaces away from the hub, dorsal side up, the backfour legs holding onto groups of radii or hubloops, the front four to the tension line, and theslack of the tension line piled on the eye groupor held by the palpi. In at least some webs nohub loops exist, so it is only the grip of thespider's back legs on the hub radii that holds thehub together. If disturbed, spiders may "drop"one or more hub radii, and the hub portion thenhas to be rebuilt before the web can be tensedagain. Consequently, many figures of Theridio-soma webs (McCook, 1889a,b; Wiehle, 1931) areof webs with rebuilt or repaired hub regions.

The eggsacs of all known species are paperyor leathery in texture, pear-shaped, and sus-pended from twigs, leaves, or stones by a longsilk line. Frequently the junction of the sac andline is marked by a distinct suture or cap—thesac dehisces at this point (Figure 158).

SPECIES.—Theridiosoma is the largest, mostwidespread genus in the family, mainly circum-tropical, but ranging into north temperate re-gions in Europe, North America (gemmosum), and


Korea {epeiroides), and into south temperate re-gions in Australia and New Zealand (species un-described). Approximately 20 species, describedand undescribed, are known from tropical Amer-ica, perhaps half that from the rest of the Trop-ics. Species can be distinguished by genitalic andcolor differences, but some are very similar (e.g.,the davisi group).

Past workers tended to place new theridioso-matid species in Theridiosoma: hence many de-scribed species belong elsewhere. Theridiosomafauna Simon, 1897, splendidum (Taczanowski,1873), and sylvicola Hingston (1932) belong inthe genus Naatlo; globosum (O. Pickard-Cam-bridge, 1896) in Epilineutes; albinotatum Petrunk-evitch, 1930, and brauni Wunderlich, 1976, inBaahebub, and nigrum (Keyserling, 1886) in Wen-dilgarda (where it was originally described). An-dasta semiargentea Simon, 1901, and A. geneven-sium Brignoli, 1972, belong in Theridiosoma ashere defined, but Andasta cyclosina Simon, 1907,is probably a theridiid.

The following species appear to be properlyplaced in Theridiosoma: argenteolunulatum Simon,1897 (type examined), benoiti Roberts, 1978, cir-culoargentum Wunderlich, 1976, davisi Archer,1953 (type examined), epeiroides Bosenberg andStrand, 1906 (figured by Brignoli, 1981), fascia-tum Workman, 1896 (type examined), goodnigh-torum Archer, 1953, (type examined), kikuyuBrignoli, 1979, lopdelli Marples, 1955, lucidumSimon, 1897 (type examined), nechodomae Pe-trunkevitch, 1930 (type examined), picteti Simon,1893 (figured by Brignoli, 1972b), and savannumChamberlin and Ivie, 1944 (type examined).Theridiosoma nelsoni Bryant, 1945 (type exam-ined), as Archer (1953) indicated, is a synonymof savannum.

The following species were not examined, andtheir placement in Theridiosoma cannot be con-firmed by the existing descriptions: blaisei Simon,1903b, concolor Keyserling, 1884, latebricolaLocket, 1968, nebulosum Simon, 1901, and obscu-rum Keyserling, 1886. The holotype and onlyspecimen of concolor is apparently lost. Simon(1903b:720) remarked that blaisei resembled

Ogulnius in body form. The reported small sizeof the type (1.5 mm) lends credence to his com-ment; perhaps blaisei belongs in Ogulnius.

RANGE.—Mainly circumtropical, occasionalextensions into temperate regions.

Theridiosoma gemmosum (L. Koch)

FIGURES 6-9, 134-137, 138-149, 157-160; MAP 5

Theridium gemmosum L. Koch, 1878:181, figs. 2 [eggsac], 6-8 [$, 6) [type series from Nuremberg, West Germany;possible syntypes exist at Vienna (NMW) and Oxford(HDEO)].

Theridiosoma argenteolum O. Pickard-Cambridge, 1879:194,fig. 8a-g [$] [type 9 from Bloxworth, Dorset, England,apparently lost (BMNH?, HDEO?)].

Theridiosoma gemmosum.—Simon, 1881:26.—Wiehle,1931:130-135, figs. 210-213 [$], 214, 215 [<5], 216, 217[web, eggsac].—Wiehle, 1967:195, fig. 54 [$].—Roewer,1942:971.—Bonnet, 1959:4817.—Archer, 1953:16-17,figs. 29, 30.

Epeira radiosa McCook, 1881:163, figs. 1-12 [all ofwebs].—Emerton, 1884:320, pi. 34: fig. 7 [9], pi. 38: figs.1 [<$], 2-4 [$]. [Type probably lost, but specimens fromMcCook's collection at ANSP examined.]

Theridiosoma radiosum.—McCook, 1889b: 195.Theridiosoma argentatum Keyserling, 1886:218, fig. 132 [im-

mature male holotype from Georgia, apparently lost(MNHN)]. [New synonymy.]

NOTE.—In his description, Koch based thename "gemmosum" on a pair of specimens, maleand female. Koch's collection is now much dis-persed. The BMNH has most of it, but no spec-imens that can be confirmed as types of gemmo-sum. Oxford has a series of gemmosum specimens,with correspondence documenting that Koch didsend specimens of his new species to Pickard-Cambridge soon after he described it. Theseseem to qualify as syntypes and have been solabeled. Likewise, Vienna, which also has part ofthe Koch collection, has a male and female withcatalog records indicating that they were col-lected in Nuremberg. These may also be syn-types, and have been so labeled. Because allavailable type specimens are gemmosum, no lec-totype is here designated.

No museum seems to have specimens that canqualify as the types of argenteolum. Perhaps Pick-

NUMBER 422 65

FIGURES 134-137.—Theridiosoma gemmosum (L. Koch), left palp: 134, mesal, embolicdivision enclosed; 135, ventral, embolic division enclosed; 136, apical (damaged),embolic division exposed; 137, subventral, embolic division exposed.

Scale lines: 100 jim.

NUMBER 422 67

ard-Cambridge, on receiving Koch's specimens,realized that his were the same and re-labeledthem. On the other hand, no specimens locatedto date bear a Bloxworth label.

McCook did not usually designate holotypesor syntype series when he described species. Onlyone vial labeled Theridiosoma radiosum (lackingany data on locality or date) was found in hiscollection at the Academy of Natural Sciences inPhiladelphia; it contains 12 gemmosum females inpoor condition. No evidence suggests that thesespecimens are the ones McCook described in1881. In fact McCook (1881) mentions havingfound the species at four localities in two statesas well as numerous others not specificallynamed, and so it is very unlikely that the availablespecimens are part of the type series. McCook(1881) gave the first locality as "in the vicinity ofPhiladelphia, June 14, 1881," and other speci-mens from that region in the MCZ and AMNHare gemmosum. Archer collected specimens helabeled as topotypes; these are also gemmosum.

The synonymy of radiosum with gemmosum has,however, been disputed. McCook (1889a: 180;1889b: 195) thought they were the same, andRoewer (1942) and Bonnet (1959) synonymizedthem, as did Kaston (1948), but the most recentauthor (Archer, 1953) maintained the taxa.Archer (1953:7) stated: "Indeed, there is no jus-tification for uniting the European species withT. radiosum (McCook) or any North Americanspecies [i.e., argentatum or savannum]. The gen-italia are readily distinguished in both sexes."After a careful study of series of North Americanmaterial identified by Archer as either radiosum

FIGURES 138-149.—Theridiosoma gemmosum (L. Koch).Female: 138, 139, habitus; 140, sternum; 141, leftlegs; 142, eye group, dorsal view; 143, epigynum,posterior; 144, same lateral; 145, same, dorsal, cleared.Left male palp: 146, expanded, subapical; 147, lateral;148, ventral (drawing by H.W. Levi); 149, subapical(drawing by H.W. Levi).

FIGURES 150-156.—Theridiosoma semiargentum (Simon).Type female: 150, habitus; 151, epigynum, ventral;152, same dorsal, cleared. Paratype male, left palp:153, dorsal; 154, lateral; 155, apical; 156, submesal.

Scale lines: 0.1 mm, except 138-141, 150, 0.5 mm.

or argentatum and of several European gemmosumkindly provided by Alex La Touche, G.H.Locket, and J. Wunderlich, I find no consistentdifferences to support a differential diagnosis.

The type of argentatum has also apparentlybeen lost. Only two theridiosomatid species occurin the southeastern USA, namely gemmosum andsavannum. The color pattern of savannum isstriking and distinctive, and Keyserling's verbaldescription fits gemmosum; hence argentatum is asynonym of gemmosum.

DIAGNOSIS.—Theridiosoma gemmosum is distin-guished from all other Theridiosoma by thestrongly sclerotized, protuberant, hood-shapedepigynum in the female (Figures 143-145), andthe form of the conductor in the male (Figures137, 148), which lacks the distal apophyses char-acteristic of savannum or goodnightorum.

DESCRIPTION.—Female: From USA, GEOR-GIA. Total length 1.9 mm. Cephalothorax 0.99mm long, 0.86 mm wide, 0.84 mm high. Sternum0.51 mm long, 0.49 mm wide. Cephalothoraxolive tan with few radiating dark streaks; legstan, segments darker distally. AME s/» PME di-ameter, AME separation '/•> their diameter, PMEseparation slightly less than Vs» their diameter.ALE, PLE subequal to PME, separated fromAME, PME by '/a their diameter. Clypeus heightslightly more than V2 AME diameter. Abdomen1.29 mm long, 1.26 mm wide, 1.22 mm high;smoothly ovoid, overhanging carapace, dorsum(Figure 139) with central dark blotches inter-rupted by white guanine dots, sides (Figure 138)peppered with white dots, venter dark with fewerdots. Epigynum a smooth domed hood (Figure143), protuberant in lateral view (Figure 138).



Total

I0.940.340.670.580.352.88

II0.810.350.570.530.332.59

III IV0.51 0.770.24 0.270.32 0.410.38 0.460.26 0.281.71 2.19

Male: From USA, NORTH CAROLINA. Total


FIGURES 157-160.—Theridiosoma gemmosum (L. Koch): 157, web, lateral view; notetension line; 158, opened eggsac; 159, web, frontal view; note radial anastomosis andlack of tension line; 160, web showing incomplete sticky spiral and non-sticky "spiral"as a few central loops and one peripheral circle.


NUMBER 422 69

length 1.9 mm. Cephalothorax 0.86 mm long,0.83 mm wide, 0.84 mm high. Sternum 0.48 mmlong, 0.45 mm wide. Eye group as in female, butAME on slight tubercle; clypeus height ¥2 AMEdiameter. Abdomen 1.26 mm long, 1.24 mmwide, 1.29 mm high. Color of carapace, sternum,abdomen, and legs as in female. Palp with con-ductor produced into strong point (Figures 135,148, 149) from which bristles of embolic apoph-ysis barely protrude in resting state.



Total

I1.090.380.900.740.353.46

II0.910.340.650.620.322.84

HI0.560.270.300.440.271.84

IV0.760.300.380.510.272.22

VARIATION.—Females range in length from1.9 to 2.4 mm, males from 1.3 to 1.9 mm. Invery light specimens the abdominal color patternis often only white guanine dots on a light back-ground; in dark specimens almost black. At leastsome males have paracymbia ending in sharpspines.

NATURAL HISTORY.—Theridiosoma gemmosumis common and widespread in eastern NorthAmerica (Map 5). The species requires a damphabitat such as ravines, swamps, stream banks,wet cliff faces, or lush undergrowth. It is mosttypical of old-growth forest, but I have taken itoccasionally in second-growth habitat. Onlyrarely is it found far from running water. Indi-vidual animals prefer web sites in dark, shadedsituations. Females move away from the wetplaces they choose as web sites toward drier sitesto construct eggsacs. Hence, as Kaston(1948:262) observed, the eggsacs often seemmore common than the spiders. The eggsac (Fig-ure 158) is pear-shaped, invariably suspended ona strong silk line at least 3-4 times the length ofthe eggsac itself. Frequently the suspension lineforks before it reaches the substrate. The sacopens at a specific spot at its upper end, the spotmarked in the intact sac by a suture.

In colder latitudes animals overwinter as su-badults, and so in New England adults can befound as early as March. The adult populationreaches a low in June and July in that region, andrises again in August and September. Hatchlingstake about 5 weeks to mature. Farther south(e.g., Georgia, Alabama) reproduction is prob-ably not seasonal.

Mating has been described by Gerhardt(1933). In New England males guard females(often even immatures) and will defend againstother males. I have observed adult males in webscontaining sticky silk—apparently they "rob"functional webs from females, as Eberhard et al.(1978) also observed in Metazygia. Males are fre-quently found on webs with guy lines, frames,and radii, but no sticky spirals, placed adjacentto those of females.

Theridiosoma gemmosum is usually considered anative European species (Locket and Millidge,1953; Wiehle, 1931). In view of the apparentsynonymy of radiosum and gemmosum, that infer-ence should be questioned. In North Americagemmosum has a large range, and it is both ubiq-uitous and common throughout. In Europe andthe British Isles its occurrence is reportedly spo-radic, despite the presence of suitable habitat.The sister taxa of the species will probably befound either in tropical America or eastern Asia(epeiroides is a very close relative). Although suchquestions cannot be conclusively settled, the evi-dence suggests that the species may be native toNorth America, and introduced to Europe. Thespecies in North America often occurs in damprock heaps or other such places, and might easilybe transported with ship's ballast.

Web construction has been described byWiehle (1931). The web (Figure 159) sometimesresembles that of Epilineutes globosus. The non-sticky spiral (Figure 160) has the form of twoconcentric circles. Most individuals spin webswith tension lines, but some construct completelyplanar webs. Some populations in the Appala-chian Mountains appear to be monomorphic foreither state. Individuals that spin tension lines atdawn will often remove them by mid-morning.


pL

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i

i Lj

K

i—.-(

tt I

)I. '\

I •"

4

0

/ -(

1

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MAP 5.—Distribution of Theridiosoma gemmosum (L. Koch)in the Western Hemisphere.

Rough measurements indicate that adult femalesimpose a force on the tension line of about 50times their body weight when in a prey-readyposture.

RANGE.—Eastern North America (Map 5)and, from Wiehle (1931) and Bonnet (1959),England, Ireland, France, Germany, Hungary,Poland, "Balkans," Greece, and Corfu. Saito(1932, 1933, 1935) listed gemmosum from thenorthern Kurile Islands (off Korea) but did notfigure the specimens; probably they are the verysimilar species epeiroides, which is native to thatregion.

RECORDS.—These records primarily includespecimens from the MCZ and AMNH collec-tions, and data are given only to county level.

USA. ALABAMA: BALDWIN (9, AMNH); COOSA

(99,66, AMNH); CONECUH (9, MCZ); JACKSON (9,AMNH); TUSCALOOSA (99, 66, MCZ). CONNECT-

ICUT: FAIRFIELD (99, 66, AMNH; see also Kaston,1948:263). DISTRICT OF COLOMBIA (99, AMNH).FLORIDA: ALACHUA (9, WJG); CALHOUN (99, 6,

MCZ); HIGHLANDS (9, MCZ); ORANGE (9, MCZ);

SEMINOLE (9, AMNH). GEORGIA: CHARLTON (99,MCZ); RABUN (9, MCZ); THOMAS (99, 66, MCZ);

UNION (9, MCZ). ILLINOIS: JACKSON (9, AMNH);KANE (9, AMNH); LAKE (9, AMNH). INDIANA:

LAWRENCE (99, 66, AMNH); MONTGOMERY (99,66, AMNH); PARKE (99,6, AMNH). IOWA: MONT-

GOMERY (9, AMNH). KENTUCKY: BREATHITT (99,66, AMNH); PERRY (?) (96\ AMNH); WOLFF (6,MCZ). MAINE: HANCOCK (66, MCZ, 96, AMNH);LINCOLN (9, MCZ). MASSACHUSETTS: BARNSTABLE

(9, AMNH); ESSEX (99, 66, MCZ); FRANKLIN (6,MCZ); MIDDLESEX (9, MCZ). MICHIGAN: CHEBOY-

GAN (99, 6, MCZ, AMNH); EMMET (99, MCZ);HILLSDALE (99, MCZ); MARQUETTE (99, 6, MCZ,AMNH); NEWAYGO (9, MCZ); OSCEOLA (6, MCZ).MINNESOTA: BLUE EARTH (9, MCZ); FREEBORN (9,

MCZ); WINONA (99, MCZ). MISSOURI: BOONE (9,

AMNH). NEW HAMPSHIRE: CARROLL (99, 66,MCZ); CHESHIRE (9, MCZ); coos (99, 66, MCZ);GRAFTON (99, 66, MCZ). NEW JERSEY: BERGEN

(99, 66, AMNH); HUNTERDON (99, 66, AMNH).NEW YORK: CORTLAND (9, AMNH); DUTCHESS

(99, 66, MCZ); ERIE (?) (9, AMNH); FRANKLIN (9,AMNH); FULTON (99, 66, MCZ, AMNH); HAM-ILTON (6, MCZ); NASSAU (99, 66, MCZ); ROCK-

LAND (eggsacs, AMNH); SUFFOLK (99, 66, MCZ,AMNH); TOMPKINS (99,66, MCZ, AMNH); WAR-

REN (6, AMNH); YATES (99, 66, AMNH). NORTHCAROLINA: BUNCOMBE (99, 66, MCZ, AMNH);HAYWOOD (9, MCZ); JACKSON (99, 66, MCZ,

AMNH); MACON (99, 66, AMNH); MCDOWELL

(99, 66, MCZ, AMNH); NANTAHALA (99, 66,UVM); SWAIN (99, 66, MCZ); YANCEY (99, 66,

MCZ). OHIO: HAMILTON (99, AMNH); HOCKING

(99, 66, AMNH); MEIGS (96, AMNH). PENNSYL-

VANIA: BERKS (immature, AMNH); BUCKS (99,66,AMNH); CENTER (99, MCZ); FRANKLIN (99, 66,MCZ); PHILADELPHIA (9, AMNH). SOUTH CAR-

OLINA: AIKEN (6, AMNH); OCONEE (9, WJG);

SPARTANBURG (9, AMNH); SUMTER (9, MCZ).TENNESSEE: ANDERSON (9, MCZ); BLOUNT (9, 6,

MCZ); GRUNDY (9, AMNH); MACON (9, WJG);

MORGAN (9, MCZ); OVERTON (9, MCZ); SEVIER(99, 66, MCZ, AMNH); SULLIVAN (9, MCZ); UN-

ICO (9, AMNH). VERMONT: ADDISON (6, MCZ);CHITTENDEN (99, 66, MCZ); LAMOILLE (99, 66,MCZ); WINDSOR (6, MCZ). VIRGINIA: AMHERST

(99, MCZ); AUGUSTA (99, MCZ); FAIRFAX (9,

MCZ); NANSEMOND (9, MCZ); PRINCE EDWARD(99, MCZ). WEST VIRGINIA: PLEASANT (99, MCZ);

POCAHONTAS (99, MCZ, AMNH); PRESTON (99,MCZ); RANDOLPH (9, AMNH). WISCONSIN:

DOUGLAS (96, MCZ); GRANT (96\ MCZ).

NUMBER 422 71

CANADA. NEW BRUNSWICK: EDMUNDSTON(CNC). NOVA SCOTIA: VICTORIA (CNC). ON-TARIO: ALGOMA ($$, AMNH); HALTON (CNC);HASTINGS (CNC); MIDDLESEX (CNC); MUSKOKA($, MCZ); NIPISSING ($9, AMNH); PETERBOR-OUGH (CNC); WENTWORTH (9$, AMNH). QUE-BEC: PERKINS (CNC).

Theridiosoma semiargenteum (Simon), newcombination

FIGURES 150-156

Andasta semiargentea Simon, 1895:918.—Roewer,1942:968.—Bonnet, 1955:320.—Archer, 1953:5.—Brignoli, 1972b:913-915, figs. 13-16 [9,8]. [Lectotypefemale (designation by Brignoli, 1972b) and paralectotypemale from Kandy, Sri Lanka, in MNHN, Paris, examined.]

DIAGNOSIS.—Theridiosoma semiargenteum isdistinguished from other Theridiosoma species bythe simple epigynal vault (Figure 151), differ-ently shaped from gemmosum, and in the palp bythe shape of the spur on the conductor (Figure156).

DESCRIPTION.—Female: Lectotype. Totallength 2.3 mm (estimated, abdomen and cephal-othorax separated). Cephalothorax 0.74 mmlong, 0.52 mm wide. Sternum 0.34 mm long,0.31 mm wide. Cephalothorax olive-brown, lightpatch around thoracic depression, legs yellow-tan, faint distal annulations. Abdomen 1.47 mmlong, 1.70 mm wide, 1.78 mm high; olive brown,transverse light stripe of faint white dots (Figure150). AME 3/4 PME diameter, separation '/a theirdiameter, PME juxtaposed. ALE, PLE subequalto AME, separated from AME by V2 AME di-ameter. Clypeus height % AME diameter. Epi-gynum a smoothly curved plate, with lateral pitsbarely visible under high magnification (Figure151). Copulatory ducts as in Figure 152.


FemurPatellalibiaMetatarsusTarsus

Total

I0.570.210.370.340.241.73

II0.500.210.340.330.241.62

III0.400.140.180.220.211.15

IV0.460.160.28--

Male: Not measured. Similar in overall ap-pearance to the female, slightly smaller (totallength 1.2 mm), legs proportionately longer, co-loration similar. Palp as in Figures 154, 156.

VARIATION.—Only the type series is known.NATURAL HISTORY.—Nothing known.RANGE.—Kandy, Sri Lanka. In his original

description, Simon also listed the species fromthe island of Taprobane, just north of Sri Lanka.

Theridiosoma genevensium (Brignoli, 1972),new combination

Andasta genevensium Brignoli, 1972b:911.

Baalzebub, new genus

TYPE-SPECIES.—Baalzebub baubo, new species(see below).

ETYMOLOGY.—Baalzebub is here taken to be amasculine noun and commemorates the earlySemitic deity. The Hebrew means literally "Lordof Flies."

DIAGNOSIS.—Baalzebub males have elongatetegula (Figure 186), small, pointed median apo-physes with a trough so deep that the scleriteappears two-pointed (Figures 163, 187). Baalze-bub females have epigynal plates shaped likebackward-pointed triangles (Figure 183; Pe-trunkevitch, 1930, fig. 179; Wunderlich, 1976,fig. 18) with a central sclerotized pit.

Epilineutes have a median apophysis with astrong dorsal spine projecting from it, and themedian apophysis is much larger and subrectan-gular. Wendilgarda also has a larger, subrectan-gular median apophysis. The trough of the me-dian apophysis in these genera is only moderatelydeep compared to that in Baalzebub. The embolicdivision of Baalzebub (Figure 162) resembles thatof Epilineutes in having four blunt, spatulateprocesses without the prolonged mesal bristlethat characterizes Wendilgarda. The tegulum ofBaalzebub is more elongate than in Epilineutes orWendilgarda (Figure 186; Wunderlich, 1976, fig.18); the mesal arm of the tegulum (bearing themedian apophysis) angles upward towards theconductor in mesal view (Figures 163, 187),


whereas in Epilineutes and Wendilgarda the te-gulum arm dips down or is more or less horizon-tal.

Baalzebub females lack the lateral pits on theepigynal plate common to Epilineutes and Wen-dilgarda. As far as is known, Baalzebub sperma-thecae are fused only at the tip (e.g., Figure 184;Wunderlich, 1976, fig. 24), and, like Epilineutes,the course of the ducts as they approach thespermathecae (Figure 184) is less complex thanin Wendilgarda. Baalzebub have juxtaposed PME;in Wendilgarda the PME are separated by at leasttheir diameter.

DESCRIPTION.—Total length 1 to 3 mm. Car-apace low in eye region (Figure 169), dark olive,dusky or tan, if light usually with dusky regionaround eye group. Eyes subequal, AME and PMEjuxtaposed or separated by at most V2 diameter(Figure 182). Sternum smooth, very convex,sparsely bristled, usually with dusky margin. Legsonly moderately long and somewhat stout.Fourth tibial trichobothria often 3-4 times tibialdiameter.

Abdomen ovoid, higher than long or broad,dorsum dark with symmetric light blotches (Fig-ure 179); sides dark with lighter marks (Figure182). Epigynum a backward-pointed triangle(Figure 183; Wunderlich, 1976, fig. 22). Sper-mathecae connate at tips, bursae thin-walled,wrinkled, with lateral wings or pockets (Figure184). Central but no lateral pits on epigynal plate,and no transverse groove on posterior epigynallip. Copulatory duct routing simple (Figure 184;Wunderlich, 1976, fig. 24).

Sexual dimorphism slight, male similar to fe-male in color pattern, often subequal in stature,legs longer, clypeus slightly higher (Figure 185).Palp shape and median apophysis distinctive (Fig-ures 161, 163, 187), and elements of embolicdivision quite short (Figures 162, 163). Conduc-tor lacks apophyses, not geniculate, and bluntlypointed.

NATURAL HISTORY.—Baalzebub prefer darkersites than Epilineutes. For example, B. baubo andrelated (but undescribed) neotropical species oc-cur in the interior of hollow logs, or under the

lee of large fallen trees in Costa Rica and Gua-temala. Baalzebub baubo has been collected incaves in Panama. Baalzebub albinotatus, however,does occur in somewhat sunnier habitats inPuerto Rico. Baalzebub don't build as close towater as do Epilineutes or Wendilgarda. Theirwebs are distinctive: complete orbs with a singlehub circle, and no radial anastomosis (Figures165, 167, 168). The single hub loop of bauboand albinotatus is constructed after the comple-tion of the sticky spiral, as in Epilineutes, but thelatter always removes the hub loop during theprocess of radial anastomosis. Tension lines arecompletely absent in baubo and albinotatus. Spi-ders do pull slightly on the radii when at the hub.

The eggsac of baubo is unknown, but that ofalbinotatus and an undescribed species from Gua-temala (Figure 166) resemble those of Epili-neutes—papery, pear-shaped, and suspendedfrom twigs, leaves or stones by a long silk line.The junction of the sac and line is also markedby a distinct suture, but not a cap.

SPECIES.—Theridiosoma albinotatum Petrunk-evitch, 1930, and T. brauni Wunderlich, 1976,belong in Baalzebub, hence Baalzebub albinotatus,new combination, and Baalzebub brauni, newcombination. Perhaps 10-20 species, all unde-scribed, have been collected.

RANGE.—Baalzebub is widespread, circum-tropical, but ranging into warm temperate re-gions in Australasia (brauni, and undescribedspecies). Thus far the genus is known from Mex-ico, Guatemala, Costa Rica, Panama, Colombia,Brazil, South Africa, Malaysia, New Guinea, Aus-tralia, and New Zealand.

Baalzebub baubo, new species

FIGURES 161, 162, 179-187; MAP 6

TYPE.—6 holotype from Costa Rica, HerediaProv. nr Puerto Viejo, Organization of TropicalStudies Field Station Finca La Selva, ~50 m, J.Coddington, 28.ii. 1982: 1, in MCZ. Two para-type 99 from same locality, J. Coddington, 15.xii.1981: 1, in MCZ.

DIAGNOSIS.—Baalzebub baubo can be distin-

NUMBER 422 73

FIGURES 161, 162.—Baalzebub baubo, new species, left male palp, embolic division ex-posed: 161,mesal; 162, ventral.

FIGURES 163, 164.—Baalzebub albinotatus (Petrunkevitch), left male palp: 163, mesal,embolic division exposed; 164, paracymbium and cymbial lamella.

Scale lines: 100 /*m.


guished by the shorter squatter palp proportionsfrom brauni and albinotatus (Figure 162). Baal-zebub albinotatus also has a median dorsal whitestripe on the abdomen and is restricted to PuertoRico (Petrunkevitch, 1930, figs. 177, 178).Other, undescribed species of Baahebub occur inat least Guatemala and Colombia in the Neotrop-ics, but their epigynal scapes are either moreapiculate or blunter than that of baubo.

DESCRIPTION.—Female: From Costa Rica,HEREDIA. Total length 1.6 mm. Cephalothorax0.69 mm long, 0.66 mm wide, 0.60 mm high.Sternum 0.41 mm long, 0.37 mm wide. Carapaceolive with dusky blotches, sternum dark yellow,dusky margin, legs yellow with faint annulations.Eye diameters subequal, AME, PME separationVa, 'A their diameter. ALE separation from AME'/> diameter, PLE from PME by diameter. Cly-peus height AME diameter. Abdomen 0.98 mmlong, 0.86 mm wide, 0.98 mm high; smoothlyovoid, dorsum with symmetric light blotches ondark background (Figure 179), sides and venteralso blotched. Epigynum as in Figures 183, 184.



Total

I0.680.300.380.400.302.06

II0.570.270.350.370.271.83

III0.380.210.220.270.241.32

IV0.560.240.340.300.241.68

Male: From Costa Rica, HEREDIA. Totallength 1.5 mm. Cephalothorax 0.77 mm long,0.65 mm wide, 0.69 mm high. Eye group as infemale, cephalic region slightly higher (Figure185). Clypeus height 2 times AME diameter.Sternum 0.38 mm long, 0.37 mm wide. Abdo-men 0.96 mm long, 0.86 mm wide, 1.00 mmhigh. Color of carapace, sternum, abdomen, andlegs as in female. Palp with elongate tegulum(Figure 186) and upturned mesa! arm of tegulum(Figure 187). Median apophysis deeply cleft, den-ticulate on distal surface.



Total

I0.860.310.690.480.292.63

II0.770.290.550.430.292.33

III0.480.220.290.310.221.52

IV0.600.220.400.330.241.79

VARIATION.—Females range in lengths from1.5 to 2.0 mm, males from 1.2 to 1.5 mm (3specimens). Overall coloration usually very dark,posterior abdominal light blotches merge to formchevrons. Males may be much lighter in color.Animals from Panama are larger and lighter.

NATURAL HISTORY.—Baahebub baubo are se-cretive, somewhat rare animals that live in darkplaces like hollow logs or under fallen limbs. Theweb is shown in Figure 168. Animals are usuallyat the hub at all times, occasionally at the periph-ery of one of the upper radii. The eggsac isunknown.

RANGE.—Costa Rica and Panama (Map 6).RECORDS.—COSTA RICA, HEREDIA: nr

Puerto Viejo, Organization for Tropical StudiesField Station La Selva ($9, 66, MCZ). PANAMA.PANAMA: Lake Gatun, Barro Colorado Island (99,6, MCZ); Chilibrillo Caves.

Baahebub albinotatus (Petrunkevitch), newcombination

Theridiosoma albinotatum Petrunkevitch, 1930:305.

Baahebub brauni (Wunderlich), newcombination

Theridiosoma brauni Wunderlich, 1976:117.

Epilineutes, new genus

TYPE-SPECIES.—Andrasta [sic] globosa O. Pick-ard-Cambridge, 1896:192, pi. 24: figs. 5, 5a-f(6), figs. 6, 6a-b (9), see below.

ETYMOLOGY.—Epilineutes is a masculineGreek noun meaning "one who catches with anet."

DIAGNOSIS.—Male Epilineutes have a median

NUMBER 422 75

FIGURES 165, 166.—Baalzebub sp. from Guatemala, ALTA VERAPAZ: 165, web, frontalview; note single hub loop and lack of radial anastomosis; 166, opened eggsac.

FIGURE 167.—Baalzebub albinotatus (Petrunkevitch), from Puerto Rico, close-up of hub.FIGURE 168.—Baalzebub baubo, new species, from Costa Rica, HEREDIA, web, frontal view;

note lack of tension line.Scale lines: 10 mm, except 166, 1 mm.

NUMBER 422 77

apophysis with a strong dorsal spine projectingfrom it (Figure 189), a complex embolic apoph-ysis with the median element prolonged (oftengreatly so, Figure 190), and a geniculate conduc-tor (Figures 188, 190). Female Epilineutes havethe posterior epigynal rim notched and a short,blunt or apiculate scape (Figure 174).

The median apophysis is squarish and has amoderately deep trough as in Wendilgarda. Theembolic division (Figure 190) resembles that ofWendilgarda, in having four blunt, spatulateprocesses, the lateral one containing the distal tipof the embolic duct, the more mesal three com-posing the embolic apophysis. The embolicapophysis of Epilineutes, however, lacks themesal, attenuate bristle characterizing Wendil-garda, and the processes of the embolic apophysisare broader and more blunt than in Wendilgarda.Female Epilineutes have a scape (Figure 173)reminiscent of that in Wendilgarda, but shorterand more pointed. Epilineutes spermathecae areconnate at the tip, and the course of the ducts asthey approach the spermathecae (Figure 174) isless complex than in Wendilgarda. Both sexes ofEpilineutes have juxtaposed PME; in Wendilgardathe PME are separated by at least their diameter.

The embolic division of Epilineutes resemblesthat of Baalzebub, but usually the embolic apoph-ysis elements are longer and more developed. Inaddition the embolus of Epilineutes does not pro-trude laterally from beneath the conductor. Themedian apophysis of Epilineutes is larger, squar-ish, and provided with a dorsal spine, rather thansmall and pointed as in Baalzebub. Epilineutes

FIGURES 169-178.—Epilineutes globosus (O. Pickard-Cambridge). Female: 169, 170, habitus; 171, left legs;172, eye group, frontal view; 173, epigynum, ventral;174, same, dorsal, cleared. Male: 175, habitus; 176,left male palp, expanded, sublateral; 177, left malepalp, mesal; 178, same ventral.

FIGURES 179-187.—Baalzebub baubo, new species. Fe-male: 179, 180, habitus; 181, left legs; 182, eye group,frontal view; 183, epigynum, ventral; 184, same, dor-sal, cleared. Male: 185, habitus; 186, left palp, lateral;187, same, submesal.


females all have a transverse, notched rim, not aposteriorly pointed, triangular epigynal plate asin Baalzebub.

DESCRIPTION.—Total length 0.5 to 2.5 mm.Carapace high in eye region (Figure 169), higherin male (Figure 175), light tan, usually duskyaround eye group. Eyes subequal, AME, PMEjuxtaposed or separated by at most 'A diameter(Figure 172). Sternum smooth, convex, sparselybristled, usually with dusky margin. Legs longand slender, metatarsi subequal or longer thantibiae (except leg III); femora distinctively long,robust proximally, slender distally (Figure 171);uniform tan. Trichobothria as in family descrip-tion, 4th trichobothria often 4-5 times tibialdiameter (Figure 171).

Abdomen ovoid, higher than long or broad;coloration various, dorsum with light guaninespeckles (Figure 173), dusky, or with brokentransverse light bands and symmetric lightblotches. Sides light, grading to dark ventrally(Figure 168), venter usually dark.

Epigynum often ridged, with blunt, notchedrim, and short, pointed median scape (Figure173). Spermathecae connate at tips, copulatorybursae thin-walled, wrinkled, with lateral wingsor pockets. Epigynal plate with lateral and centralpits (Figures 173, 174). No transverse groove onposterior epigynal lip. Copulatory duct routingsimple (Figure 174).

Sexual dimorphism slight, male color patternusually similar to female, stature slightly smaller,legs proportionately longer, eye group propor-tionately much higher (Figure 175). Palpi withdistinctive median apophysis (Figure 177), anddivided embolic division (Figure 190, usually vis-ible through translucent conductor, Figure 178).Conductor without apophyses, somewhat genic-ulate, prolonged towards tegulum (Figures 178,188).

NATURAL HISTORY.—Epilineutes often buildclose to or over water in the same habitat asWendilgarda, but they construct complete orbs(Figure 194). The central radii of the web areanastomosed, usually with 4-5 radii reaching thehub (Figure 192). The spider anastomoses radial


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MAP 6.—Distributions of Baalzebub baubo, new species, and Epilineutes globosus (O. Pickard-Cambridge).

lines and constructs the hub radii after comple-tion of the sticky spiral. Tension lines are rare,but do occur occasionally, especially among ju-veniles or in the first web of the day spun byadults. In webs without tension lines, the spidersoften "pull up" on one of the hub radii, althoughthe web is never distorted as much as in Theridio-soma, where the tension line is routinely normalto the web surface. In at least some webs of E.globosus no hub loops exist, and the spider holds

the hub together with its legs, as in many Theri-diosoma. Also like Theridiosoma, the non-stickyspiral is in the form of one peripheral circle(Figure 195).

The eggsacs of all known species are papery,pear-shaped, and suspended from twigs, leavesor stones by a single, long silk line. The junctionof the sac and line is marked by a distinct suture,but not a cap (Figure 193). The sac does open atthis point.

NUMBER 422 79

SPECIES.—Epilineutes is thus far believed to beneotropical. Perhaps 5 or 10 undescribed speciesin this genus occur in the American tropics.

Epilineutes globosus (O. Pickard-Cambridge),new combination

FIGURES 169-178, 188-195; MAP 6

Andrasta [sic] globosa O. Pickard-Cambridge, 1896:192, pi.24: figs. 5, 5a-f [6], figs. 6, 6a-b [9].—Bonnet,1955:321.—Roewer, 1942:970. [A series of 3d, 5? syn-types from Teapa, Tabasco, Mexico in BMNH, examined.Andrasta is a lapsus calami for Andasta Simon, 1895.]

Theridiosoma globosum.—F.O. Pickard-Cambridge, 1902:414. Simon, 1903a: 1007.—Bonnet, 1959: 4434.—Roewer, 1942:970.—Archer, 1953:11, fig. 16 [9].

NOTE.—Both F.O. Pickard-Cambridge (1902)and Simon (1903a) questioned whether globosuswas a synonym of Theridiosoma argentatum (= T.gemmosum). The ranges of the two species arecompletely disjunct, and their morphology verydifferent.

DIAGNOSIS.—Epilineutes globosus can be distin-guished by the large, ridged, protruding epigynalplate with a dark, pointed, short scape and thelateral dark spots on the epigynal plate in thefemale (Figure 173), and the very large palpalbulb with the embolic apophysis and medianapophysis as in Figures 175-178 in the male. Inparticular the middle division of the embolicapophysis frequently protrudes from beneath theconductor (Figure 190), and shows as a darkstripe under the translucent conductor (Figure178). Other Epilineutes species have scapes butthey are more blunt or the epigynal plate issmaller in proportion and not as ridged.

DESCRIPTION.—Female: From Costa Rica,HEREDIA. Total length 1.9 mm. Cephalothorax0.83 mm long, 0.71 mm wide, 0.55 mm high.Sternum 0.42 mm long, 0.40 mm wide. Carapacelight yellow, dusky around eyes; sternum darkeryellow, dusky margin; legs yellow with faint an-nulations. AME % PME diameter, AME, PMEseparation ]A their diameter. ALE, PLE subequalto PME, separated from AME, PME by theirdiameter. Clypeus height 2 times AME diameter.

Abdomen 1.29 mm long, 1.08 mm wide, 1.32mm high; smoothly ovoid, dorsum with guaninespeckles on light background (Figure 170), sidesand venter also light. Epigynum as in Figures173, 174.



Total

I0.900.340.630.540.332.74

II0.780.310.510.470.282.35

III0.470.210.270.320.241.51

IV0.690.250.380.390.241.95

Male: From Costa Rica, HEREDIA. Totallength 1.8 mm. Cephalothorax 0.74 mm long,0.72 mm wide, 0.98 mm high. Eye group morecompact than in female, cephalic region veryhigh (Figure 175). Clypeus height 4 times AMEdiameter. Sternum 0.41 mm long, 0.43 mmwide. Abdomen 1.08 mm long, 1.03 mm wide,1.20 mm high. Color of carapace, sternum, ab-domen, and legs as in female. Palp large, con-ductor extended, almost touching tegular mar-gin (Figure 178). Dorsal bristle on median apoph-ysis prominent.



Total

I1.070.310.740.690.383.19

II0.890.310.640.600.362.80

HI0.480.220.290.380.261.63

IV0.690.240.400.460.242.03

VARIATION.—Females range in length from1.7 to 2.6 mm, males from 1.5 to 1.9 mm. Invery light specimens the abdominal color patternis often only white guanine dots on a light back-ground; darker specimens have a dark back-ground with a pattern of light blotches. Animalsfrom Mexico, including the type series, aresmaller in stature than more southern popula-tions.

NATURAL HISTORY.—Epilineutes globosus is apredictable part of the streamside fauna through-


FIGURES 188-191.—Epilineutes globosus (O. Pickard-Cambridge), from Panama, PANAMA,left male palp: 188, mesal, embolic division enclosed; 189, ventral, embolic divisionenclosed; 190, subapical, embolic division exposed; 191, paracymbium and cymbiallamella.

Scale lines: 100 fim.

NUMBER 422 81

FIGURES 192-195.—Epilineutes globosus (O. Pickard-Cambridge): 192, hub region of web;193, unhatched eggsac; 194, web, frontal view; 195, incomplete web, showing non-sticky "spiral" as a few central loops and one peripheral circle.



out the lowland tropics of Central America.Adults prefer the dark, moist hollows created bystreamside cave-ins, or similar situations. Juve-niles make webs in more open sites, often onsnags in the middle of the stream. The web(Figures 192, 194, 195) resembles that of Theri-diosoma gemmosum. The eggsac is a pear-shapedpapery sac with a faint annulation at the junctionof the single suspension line and the sac itself,and is laid some distance from the female's web(Figure 193). The males are commonly found onlines at the side of the female web.

RANGE.—Mexico, Guatemala, Costa Rica,Panama, Colombia, Venezuela, Ecuador, Peru,and Brazil (Map 6).

RECORDS.—MEXICO, TABASCO: Teapa (99,66, BMNH). CHIAPAS: Palenque, Palenque ruins(99, 66, MCZ). GUATEMALA, IZABAL: SantoTomas; Livingston (99 ,66, MCZ). COSTARICA, HEREDIA: nr Puerto Viejo, Organizationfor Tropical Studies field station Finca La Selva(99,66, MCZ). LIMON: Cahuita (99, 6, MCZ). PUN-

TARENAS: San Vito, Organization for TropicalStudies field station Las Cruces (99, 66, MCZ).PANAMA. PANAMA:, Lake Gatun, Barro Colo-rado Island (99, 66, MCZ, AMNH); Gamboa (9,MCZ); El Valle (99, 66, MCZ). COLOMBIA.META: ~15 km SW Puerto Lopez, Hacienda Moz-ambique, 200 m (99, MCZ). CHOCO: betweenCucurripi and Nanama, Quebrada Docordo, RioSan Juan (6, AMNH). VALLE: Central HidalgoAnchicaya (99, 6, MCZ); Bajo Colima (on ColimaRiver), nr Puerto Patiao (6*, MCZ); Rio Querada,Descansion (not located, not mapped, 9, AMNH).ECUADOR, MORON A-SANTI AGO: LOS TayosCaves (99, MCZ). PERU, AMAZONAS: Montene-gro, 350 m (6, AMNH). BRAZIL, RIO DE JA-NEIRO: Teresopolis (99, MCZ, AMNH).

Wendilgarda Keyserling, 1886

Wendilgarda Keyserling, 1886:129.—Bonnet, 1959:4816.—Roewer, 1942:971.—Archer, 1953:14. [Type-species by original designation Wendilgarda mexicana Key-serling. 1886:130, pi. 15: fig. 208a-c (9). The name isfeminine.]

Enthorodera Simon, 1907:264 [type-species by monotypy E.atricolor Simon, 1907:265]. [New synonymy.]

Cyathidea Simon, 1907:265 [type-species by monotypy C.merula Simon, 1907:26]. [New synonymy.]

NOTE.—The holotype females of E. atricolorand C. merula have blunt scapes as in W. mexicana,one central pit and a pair of lateral pits on theepigynal plates, and eye group proportions, legproportions, and coloration as in Wendilgarda.Consequently the genera are judged synonymouswith Wendilgarda, even though web architectureand eggsac form are unknown.

DIAGNOSIS.—Wendilgarda is recognized by theorange-yellow of the cephalothorax combinedwith uniform abdomen color, the mesal bristleof the embolic apophysis protruding from be-neath the conductor and lying along the mesalside of the conductor itself (Figure 198), thecomplex looping of the copulatory ducts at thejunction with the spermathecae (Figure 215), thefluted, double-pointed form of the eggsac (Fig-ures 200, 201), and the web architecture (Figure202).

Wendilgarda has posterior median eyes sepa-rated by at least their diameter, whereas the PMEof both Epilineutes and Baalzebub are juxtaposed.Epilineutes has a squarish median apophysis likeWendilgarda, but the median apophysis has astrong dorsal spine. The median apophysis ofBaalzebub is much smaller, and more pointed,and the tegulum in lateral view is elongate. Thescapes of female Wendilgarda are longer andlarger than those of Epilineutes, and the sperma-thecae are connate throughout, not just at theirtips. In Baalzebub the epigynal plate is triangular.

Two behavioral characters also diagnose Wen-dilgarda: web architecture (Figure 202) and eggs-acs fluted in cross-section that taper to fine pointsalong the longitudinal axis (Figures 200, 201).The web architecture is unique, but at least oneundescribed species of Ogulnius makes a double-pointed, fluted eggsac. It, however, is both muchsmaller than those of Wendilgarda and also isattached at both ends, whereas the eggsacs ofWendilgarda are pendant from a single line.

DESCRIPTION.—Total length 1 to 3 mm. Car-

apace uniform yellow-orange, as long as broad.Cephalic region high in lateral view, clypeus 3

NUMBER 422 83

times the diameter of an AME (Figure 208).Sternum reddish brown to yellow, often withdusky margins, as broad as long, slightly convex,sparsely bristled. Posterior margin prolonged be-tween fourth coxae. Legs yellow-tan, slightlydarker distally, metatarsi often longer than tibiaeand abruptly tapered proximally. Tibial tricho-bothria long, in two or three irregular rows. Eyediameters subequal or AME largest. PME sepa-rated by at least a diameter, never juxtaposed.Abdomen uniform gray to black (sometimesbleached in preserved material) with no pigmentpattern; smooth, abdominal muscle apodememarks prominent. Abdomen wider than long,overhanging carapace, sometimes dorso-ven-trally flattened. Venter often with two indenta-tions just forward and lateral to spinnerets. Epi-gynum with scape (Figures 206, 213, 219), malepalp with complex embolic division; embolicapophysis composed of broad, spatulate lobes.

NATURAL HISTORY.—Wendilgarda are oftencommon along the margins of slow-moving, low-land tropical streams. They do not seem to occurabove an elevation of 500 m, in fast rapids, oralong stream courses that become dry for anextended period. Individuals are active at alltimes of day, except for an apparent siesta inmidday during the dry season. Web constructionis most frequent at dawn and dusk, althoughindividuals often re-spin their webs once or twicean hour. In appropriate habitats population dens-ities may reach 3-4 individuals/m of stream mar-gin. During heavy rains spiders remain over thestream and simply rise with the flood waters.Their webs are attached to overhangingbranches or snags in the stream itself. The webof Wendilgarda clara was first described by Simon(1895), and has been redescribed by Coddingtonand Valerio (1980) (both under the name W.theridionina). W.G. Eberhard (pers. comm.)found that at least some individuals of W. gala-pagensis do not attach their webs to water, unlikethe other members of the group whose web isknown. Prey records from clara, mustelina, andmexicana include mainly immature Gerridae andnematocerous flies. The web seems to functionin two distinct ways, by trapping insects actually

on the surface of the water, and by trapping low-flying insects aerially. Because most prey col-lected from webs were at the junction of thesticky silk vertical line and the water, rather thanmidway up the line, it seems that the formermode of capture is more frequent. Eggsacs are2-3 mm in diameter, 4-7 mm long, contain upto 15 eggs, and are suspended on single, rein-forced lines about 10 mm long. Unlike Theridio-soma eggsacs, there is no distinct cap or opercu-lum. In the appropriate habitat, eggsacs can becommon along streams in the upper portions ofoverhanging vegetation. The eggsacs are laidsome distance from the water, the most distant Ihave seen being almost 4 meters.

SPECIES.—Wendilgarda assamensis Fage, 1924,is incertae sedis (see above under family descrip-tion. Wendilgarda bicolor Keyserling, 1886, W.guacharo Brignoli, 1972a, and W. miranda Brig-noli, 1972a, belong in Plato. Keyserling (1886)originally described nigra in Wendilgarda, butSimon (1895) transferred the species to Theridio-soma. Examination of the type as well as othermaterial (NRS) confirms Keyserling's originalopinion and the species is here returned to Wen-dilgarda. Wendilgarda clara Keyserling, 1886,mustelina Simon, 1897, and galapagensis Archer,1953, are also properly placed in Wendilgarda.About 10 new species have been collected fromthe Neotropics.

RANGE.—Wendilgarda is predominantly neo-tropical; W. atricolor is African.

Wendilgarda mexicana Keyserling

FIGURES 199, 200, 203-211; MAP 7

Wendilgarda mexicana Keyserling, 1886:130, pi. 15: fig.208a-c [9] [a series of 9 syntype females from Mexico inBMNH, examined].—Roewer, 1942:971.—Bonnet.1959:4817.—Archer, 1953:14-15.

DIAGNOSIS.—Female W. mexicana may be dis-tinguished from other Wendilgarda species by theshort blunt scape of the epigynum, the differ-ently arranged internal ducts, and the sclerotizedtransverse bar bridging the spermathecae (Fig-ures 206, 207). Males are distinguished by the


FIGURES 196-198.—Wendilgarda clara Keyserling, left male palp: 196, mesal, embolicdivision exposed; 197, ventral, embolic division enclosed; 198, ventral, embolic divisionexposed.

FIGURE 199.—Wendilgarda mexicana Keyserling, from Mexico, YUCATAN, left palp, ven-tral.

Scale lines: 100 fim.

NUMBER 422 85

FIGURE 200.—Wendilgarda mexicanaKeyserling, from Mexico, YUCA-TAN, eggsac.

FIGURES 201, 202.—Wendilgardaclara Keyserling: 201, eggsac; 202,web attached to stream surface(note reflections).

Scale lines: 1 mm, except 202, 10mm.

relatively small size of the bulb (Figure 210) andthe relatively simple embolic division (Figure199). The conductor also lacks the transversedark stripe that characterizes the other Wendil-garda species. Wendilgarda mexicana tends to belarger, more reddish, and the abdomen more

flattened than other Wendilgarda species, andwith smaller PME.

DESCRIPTION.—Female: From Mexico, YU-CATAN. Total length 2.6 mm. Cephalothorax1.12 mm long, 1.08 mm wide, 1.01 mm high.Sternum 0.69 mm long, 0.65 mm wide. Carapace


220

NUMBER 422 87

uniform orange-yellow, legs proximally samecolor, darker distally. Abdomen 1.72 mm long,1.93 mm wide, 2.06 mm high; uniform reddishgray, smoothly ovoid, dorso-ventrally flattened,dorsal muscle scars as four distinct pits. AME 4hthe PME diameter, AME separation xfa their di-ameter, PME separation 1 diameter. ALE, PLEsubequal, separated from AME, PME by theirdiameter. Clypeus 3 times AME diameter. Epi-gynum a smooth plate, blunt scape barely visible(Figure 206), dorsal view as in Figure 207.



Total

I1.430.480.891.070.524.39

II1.290.430.770.950.483.92

III0.950.360.530.720.342.90

IV1.170.410.690.840.383.49

Male: From Cuba, VILLAS. Total length 2.0mm. Cephalothorax 0.89 mm long, 0.98 mmwide, 0.95 mm high. Sternum 0.60 mm long,0.53 mm wide. Abdomen 1.15 mm long, 1.15mm wide, 1.41 mm high, smaller, more ovoidthan female. AME 4A PME diameter, AME sep-aration VA their diameter, PME separationslightly more than 1 diameter, ALE slightlylarger than PLE, separated from AME, PME bytheir diameters. Clypeus 3 times AME diameter.Color of carapace, sternum, legs and abdomenas in female. Palp as in Figures 210, 211, rela-

tively small, ratio of palp width in ventral viewto sternum width 0.6-0.7; embolic division sim-ple (Figure 199).



Total

I1.380.400.911.000.454.14

II1.170.380.760.860.433.60

III0.830.310.460.520.342.46

IV0.980.360.570.720.292.92

VARIATION.—Females range in length from2.1 to 3.2 mm, males from 1.6 to 2.0 mm. Inmany cases the epigynal plate of the female isfused to the scape and thus incomplete medially.Populations on Cuba apparently are substantiallylarger than on the mainland.

NATURAL HISTORY.—The web is qualitativelyindistinguishable from that of clara, and theeggsac (Figure 200) is typical of the genus. Likeother Wendilgarda species, mexicana is alwaysfound over water. Two prey items taken withspecimens in Mexico were both immature Ger-ridae. One eggsac contained 10 eggs.

RANGE.—Mexico, Guatemala, Honduras, andCuba (Map 7).

RECORDS.—MEXICO, CHIAPAS: Triunfo (9,AMNH); 20 km N Huixtla (99, AMNH); SanCristobal de Las Casas (9, AMNH). COLIMA: Te-coman, Orilla, Rio Ameria ($9, AMNH); Bocade Pasquales (99, 66, AMNH). TAMAULIPAS: 1 mi

FIGURES 203-211.—Wendilgarda mexicana Keyserling.Female: 203, 204, habitus; 205, left legs; 206, epi-gynum, ventral; 207, same, dorsal, cleared; 208, eyegroup, frontal view. Male: 209, habitus; 210, left palp,ventral; 211, same, lateral.

FIGURES 212-217.—Wendilgarda clara Keyserling. Fe-male: 212, habitus, dorsal view; 213, epigynum, ven-tral; 214, same, ventral, cleared; 215, same, dorsal,cleared. Left male palp: 216, cymbium, ventral view;217, bulb, ventral.

FIGURES 218-220.—Wendilgarda atricolor (Simon), hol-otype female: 218, dorsal view; 219, epigynum, ven-tral; 220, same, dorsal, cleared.

Scale lines: 0.1 mm, except 203-205, 209, 212, 218, 0.5mm. MAP 7.—Distribution of Wendilgarda mexicana Keyserling.


SE Villa Juarez (immatures, AMNH); 3 mi SECiudad del Mais (99, AMNH). SAN LUIS POTOSI:

Pujal (immature, 9, AMNH); 13 km NE Xilitla(99, 66, AMNH); 10 km W San Joaquin (9,AMNH). YUCATAN: Chichen-Itza, Cenote Stoloc($9, 66, MCZ, AMNH); "Mexico," more exactlocality unknown, not mapped (99, BMNH).GUATEMALA, ESCUINTLA: Tiquisate, 60 m (9,AMNH). PETEN E end Lago Itza, Tikal (6,AMNH). HONDURAS. TEGUCIGALPA: NuevoRosaria, San Juancita (99, UCR). CUBA, VILLAS:

Soledad (nr Cienfuegos) (99, 66, AMNH, MCZ);Mina Carlota, Trinidad Mts (99, MCZ).

Wendilgarda clara Keyserling

FIGURES 196-199, 201, 202, 212-217; MAP 8

Wendilgarda clara Keyserling, 1886:132, pi. 15: figs. 210,210a-d [9] [female holotype from Amazonas Prov., Brazil(HDEO), examined].

Wendilgarda theridionina Simon, 1895:919, fig. 986 [<$] [aseries of 21 syntype males and 18 females from SanEsteban, Carabobo Prov., Venezuela (MHNH), exam-ined].—Simon, 1897a:484, pi. 13: figs. 4-6 [eggsac].—Roevser, 1942:971.—Bonnet, 1959:4817.—Archer,1953:16-17, figs. 29, 30. [Lectotype female here desig-nated.] [New synonymy]

Wendilgarda panamica Archer, 1953:15, fig. 23 [female hol-otype from Barro Colorado Island, Lake Gatun, Panama,Panama, in AMNH, examined]. [New synonymy.]

Wendilgarda hassleri Archer, 1953:16, figs. 24, 28 [maleholotype from upper Shudikar River, Guyana, in AMNH,examined]. [New synonymy.]

NOTE.—Keyserling (1886:133) said the singlefemale specimen of clara was in the O. Pickard-Cambridge collection. HDEO has a single femalelabeled Bellinda clara. The genus Bellinda is asynonym of Argyrodes (Theridiidae), but thename Bellinda clara is not listed in Bonnet(1955), Roewer (1942), or Brignoli (1983), andhence was apparently never published. Probablythis specimen was the one before Keyserling, andhis vial label simply disagrees with his publishedplacement of the species. The Bellinda clara spec-imen agrees in most details with Keyserling'sdescription and figures. It seems simplest to as-sume that this specimen is the type of clara and,

consequently, is the oldest available name for thespecies, even though theridionina has been morecommonly used.

The type series of theridionina contained twotheridiid species, Achaearanea nigrovittata (Key-serling) and Dipoena alta (Simon) (det. H.W.Levi), and one other theridiosomatid, Epilineutesglobosus (O. Pickard-Cambridge) as well as theri-dionina, which composed the bulk of the series;a lectotype female has been designated to ensurestability. The other specimens have been desig-nated as paralectotypes of theridionina.

No features were found that distinguishedeither hassleri or panamica from clara. Archerwas the only person to identify specimens besidesthe types as hassleri or panamica, and some of hisparatype material is Epilineutes globosus. Theranges of hassleri and panamica are also entirelycontained by that of clara. Because Archer of-fered no differential diagnosis of hassleri and saidonly that panamica was smaller in stature thanclara, the three taxa are synonymized.

DIAGNOSIS.—Wendilgarda clara is distin-guished from mexicana and mustelina by theshort, pointed scape in the female (Figure 213),and by the shape of the median apophysis, thetegular spur, and relative size of the palp in themale (Figure 217). The species is distinguishedfrom galapagensis by the lighter color of theabdomen (the abdomen of galapagensis is darkbrown or black).

DESCRIPTION.—Female: From Puerto Rico,HUMACAO. Total length 2.1 mm. Cephalothorax0.98 mm long, 0.91 mm wide, 0.86 mm high.Sternum 0.52 mm long, 0.52 mm wide. Carapacepale yellow, legs proximally yellow, more orangedistally. Abdomen 1.31 mm long, 1.43 mm wide,1.55 mm high; uniform pale grayish tan,smoothly ovoid, dorsal muscle scars as four dis-tinct pits (Figure 212). Eye diameters subequal,AME, PME separation VL> diameter. ALE, PLEseparated from AME, PME by slightly less thantheir diameter. Clypeus 3 times AME diameter.Epigynum a smooth plate, incised medially, withdark sclerotized spot on center, two lateral pits,and short, tapering scape truncate distally (Fig-ures 213, 214).

NUMBER 422 89



Total

I1.200.410.830.830.403.67

II1.080.400.650.970.343.44

III0.650.290.400.500.282.12

IV0.910.330.560.690.342.83

Male: From Puerto Rico, HUMACAO. Totallength 1.8 mm. Cephalothorax 0.95 mm long,0.89 mm wide, 0.83 high. Sternum 0.52 mmlong, 0.48 mm wide. Abdomen 1.20 mm long,1.20 mm wide, 1.26 mm high. Eye group as infemale. Clypeus 2 times AME diameter. Color ofcarapace, sternum, legs, and abdomen as in fe-male. Ratio of palp width to sternum width 0.9-1.0. Palp as in Figures 196-198, 216, 217.



Total

I1.260.410.980.860.463.97

II1.170.400.720.810.413.51

III0.710.260.430.520.312.23

IV0.860.310.520.640.292.62

VARIATION.—Females range in length from1.7 to 2.5 mm, males from 1.5 to 1.9 mm. Infresh specimens the abdomen is sometimes grayrather than tan, but never black. Very largespecimens superficially resemble W. mexicana,but can be distinguished by the genitalia. Wen-dilgarda clara seems to be the most widespreadof all Wendilgarda, occurring as least as far southas Ecuador and as far north as Puerto Rico.

NATURAL HISTORY.—The web (Figure 202) ispeculiar, consisting of a horizontal non-sticky linefrom which vertical sticky lines adhere with con-siderable tenacity to the surface film of flowingtropical streams (Coddington and Valerio, 1980).Wendilgarda clara is always found over water.The eggsac (Figure 201) is similar to that of W.mexicana. Live animals usually appear distinctlyorange.

RANGE.—Antilles, Guatemala south to Col-ombia, Venezuela, Guyana, and Brazil (Map 8).

RECORDS.—PUERTO RICO, AGUADILLA:

Maraicao, El Vivero (9, MCZ). GUAYAMA: Cidra,Treasure Island (9, d\ AMNH). HUMACAO: Car-ibbean National Forest, Luquillo Division (nu-merous records, 9$, 66, AMNH, MCZ). MAYA-GUEZ (numerous records, 99, 66, AMNH, MCZ).PONCE: Castaner (99, AMNH). SAN JUAN: San

Juan (9, AMNH). DOMINICAN REPUBLIC, ELSEIBO: forest between Hato Mayor and Sabanade La Mar (99, 6, AMNH). LA VEGA: Constanza(9, MCZ). PUERTO PLATA: Puerto Plata (6, MCZ).HAITI. Nr Aux Cayes (9, AMNH). GUATE-MALA, IZABAL: Santo Tomas (9$, 66, MCZ, nu-merous records). COSTA RICA, HEREDIA: nrPuerto Viejo, Organization for Tropical Studiesfield station Finca La Selva (99, 66, MCZ, nu-merous records), PUNTARENAS: Corcovado Na-tional Park (99, 66, MCZ, numerous records).PANAMA, COLON: Portobelo (66, MCZ); Pan-ama, Lake Gatun, Barro Colorado Island (99,66,MCZ, AMNH, numerous records). COLOM-BIA, VALLE: Central Hidalgo Anchicaya (9, 66,MCZ). VENEZUELA, AMAZONAS: upper Rio Pa-ciba (9, AMNH). GUYANA, ESSEQUIBO: RU-

punni River between Isherton and Dadawana (9,6, AMNH). ECUADOR, LOS RIOS: Juan Mon-talvo (6, AMNH). PICHINCHA: 4.7 km SW ofSanto Domingo de los Colorados, Rio Palenque(9, 6, MCZ). BRAZIL, AMAZONAS? (more exactlocality unknown, not mapped).

Wendilgarda atricolor (Simon), newcombination

FIGURES 218-220

Enthorodera atricolor Simon, 1907:265, no figs, [female hol-otype from "Roca. inf. d'Henrique," Principe (Republicof Sao Tome and Principe), in MCSN, Genoa, examined]

Cyathidea merula Simon, 1907:266, no figs, [female holotypefrom "Agua Ize, Ribeira Palnia, Sao Tome (Republic ofSao Tome and Principe), in MNHN, Paris, examined].[New synonymy.]

NOTE.—See remarks under the genus Wendil-garda for justification of synonymy of Enthorod-era and Cyathidea. The synonymy of atricolor andmerula is less certain, because only the holotype


MAP 8.—Distribution of Wendilgarda clara Keyserling.

females are known. The scape of atricolor isdefinitely slightly bifid, whereas that of merula ismore rounded, but on the whole the differencebetween the two specimens is very slight. Somaticmorphology seems indistinguishable. Shouldlarger series of males and females refute thesynonymy, merula would still be the sister taxonto atricolor by the posterior ventral pits on theabdomen. The abdomen of the atricolor typefemale is badly damaged and separated from thecephalothorax, and so Figure 218 is a reconstruc-tion. Wendilgarda atricolor is taken as the appro-priate name for the taxon because Simon used itfirst in his publication.

DIAGNOSIS.—Wendilgarda atricolor most re-sembles W. mexicana in its size, blunt scape, andeye proportions. It can be distinguished by thecopulatory pores, which are visible in ventral

view, and the shape of the scape (Figure 219).DESCRIPTION.—Female: Holotype. Total

length 2.6 mm (estimate). Carapace 1.15 mmlong, 0.98 mm wide, 0.77 mm high. Sternum.0.58 mm long, 0.58 mm wide. Cephalothoraxlight red-tan, legs similar. Abdomen 1.45 mmlong, 1.55 mm wide, 1.89 mm high; uniform dullbrown, smoothly ovoid, dorsal muscle scars asfour distinct pits. AME % PME diameter, sepa-ration V-j their diameter. PME separation 0.4their diameter. ALE, PLE subequal, slightlylarger than AME, separated from AME, PME by% their diameter. Clypeus height slightly morethan 2 times AME diameter. Epigynum a smoothplate with lateral pits, central transverse pit, pos-terior rim incised laterally revealing copulatorypores, also medially; short, broad scape notcheddistally (Figure 219). Dorsal view, Figure 220.

NUMBER 422 91

Leg lengths of holotype female (±0.03 mm). Male: Unknown.j JJ jjj j V V A R I A T I O N . — U n k n o w n .

Femur 1.29 1.24 0.86 1.07 NATURAL HISTORY.—Unknown.Patella 0.46 0.43 0.36 0.38 RANGE.—Islands of Principe and Sao Tome,Tibia 0.84 0.77 0.53 0.65 off African coast.Metatarsus 0.81 0.83 0.62 0.71Tarsus 0.46 0.48 0.41 0.40

Total 3.86 3.75 2.78 3.21

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