Journal of Arachnology

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Journal of Arachnology 35(2):334-395. 2007
doi: 10.1636/SH-06-36.1

MORPHOLOGY AND EVOLUTION OF COBWEB SPIDER MALEGENITALIA (ARANEAE, THERIDIIDAE)

Ingi Agnarsson, Jonathan A. Coddington, Barbara Knoflach

aSystematic Biology-Entomology, Smithsonian Institution, NHB-105, PO Box 37012, Washington, DC 20013-7012, USA

bThe University of British Columbia, Departments of Botany and Zoology, 3549-6270 University Blvd., Vancouver, B.C. V6T 1Z4, Canada.

cUniversity of Innsbruck, Institute of Ecology, Division of Terrestrial Ecology and Taxonomy, Technikerstrasse 25, A-6020 Innsbruck, Austria

Abstract

This study elucidates the homology of elements of the male palps in the spider family Theridiidae. We survey and illustrate 60 species from 29 out of the 86 currently recognized genera representing all subfamilies. The study is buttressed by a phylogenetic framework, and uses a new method to evaluate critically competing homology hypotheses based on various criteria. Among the classic criteria for homology, topology performed better than special similarity, and much better than function. Guided by those results, we propose names for and correspondences among the broad diversity of theridiid palpal tegular sclerites. We discuss the phylogenetic utility and distribution of key palpal characteristics, and evaluate existing evolutionary hypotheses of the theridiid palp and its components.

Received: 14 June 2006; Revised: 22 May 2007

Keywords: Character homology, congruence, phylogeny, tests of homology, primary homology



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Appendix A – Abbreviations


BC bulb-cymbium lock mechanism

BH basal haematodocha

C conductor

CA cymbial apophysis

Cb conductor base

CHd theridiid cymbial hood

CHk theridiid cymbial hook

Cy cymbium

E embolus

EA embolic apophysis

Eb embolic division b

ES embolic sclerite

ETS extra tegular sclerite

MA median apophysis

MH median haematodocha

PC paracymbium

SC subconductor

ST subtegulum

T tegulum

Tp tegular pit

TTA theridiid tegular apophysis



Appendix B – Material Examined (deposited in the CTh Collection Thaler & Knoflach)


For additional material examined, see Agnarsson (2004).

Achaearanea lunata (Clerck 1757). Austria, Northern Tyrol, Innsbruck, Hötting, 15 May 1992, leg. Knoflach.

Achaearanea riparia (Blackwall 1834). Italy, Treviso, Quartier del Piave, Palu, pitfall trap, 1989/ 1990, leg. Targa.

Crustulina guttata (Wider 1834). Austria, Northern Tyrol, Ötztal, Längenfeld, 14 April 1992, leg. Knoflach.

Dipoena melanogaster (C.L. Koch 1837). Austria, Northern Tyrol, Innsbruck, Hötting, 15 May 1992, leg. Knoflach.

Enoplognatha latimana Hippa and Oksala 1982. Austria, Burgenland, Parndorf, 1988, leg. Thaler, Meyer, Steinberger.

Enoplognatha ovata (Clerck 1757). Austria, Northern Tyrol, Innsbruck, Martinswand, 3 August 1991, leg. Knoflach. Telfs, Zimmerberg, 17 July 1991, leg. Bertrandi. Ötztal Bahnhof, Forchet, 16 May 1992, leg. Knoflach. Kufstein, Langkampfen, tree eclector, 28 June–23 July 1988, leg. Thaler, Meyer, Steinberger.

Enoplognatha thoracica (Hahn 1833). Italy, Veneto, Treviso, pitfall trap, 1990–1991, leg. Schiroto, Paletti.

Episinus angulatus (Blackwall 1836). Austria, Northern Tyrol, Innsbruck, Kranebitten, 12 July 1991, leg. Knoflach.

Episinus theridioides Simon 1873. France, Corsica, Col de Vizzavona, 1100–1400 m, 1 October 1974, leg. Thaler.

Episinus truncatus Latreille 1809. Austria, Northern Tyrol, Innsbruck, Kranebitten, 20 July 1991, leg. Knoflach.

Euryopis flavomaculata (C.L. Koch 1836). Austria, Vienna, Lobau, 19 May–2 June 1972, leg. Steiner.

Keijia tincta (Walckenaer 1802). Austria, Northern Tyrol, Innsbruck, Kranebitten, 10 May 1991, leg. Knoflach.

Kochiura aulica (C.L. Koch 1838). Croatia, Rovinj, 29 July–26 August 1965, leg. Thaler.

Lasaeola tristis (Hahn 1833). Austria, Northern Tyrol, Ötztal, Sautener Forchet, 26 June 1992, leg. Bertrandi.

Neottiura bimaculata (Linné 1767). Austria, Northern Tyrol, Innsbruck, Kranebitten, 23 June 1991, leg. Knoflach.

Pholcomma gibbum (Westring 1851). Austria, Northern Tyrol, Innsbruck surroundings, Halltal, 13 June 1992, leg. Thaler.

Robertus neglectus (O. Pickard-Cambridge 1871). Italy, Veneto, Treviso, Riese, pitfall trap 1990– 1991, leg. Schiroto, Paoletti. Germany, near Immendingen, Zimmern, leg. Wunderlich 1973.

Robertus scoticus Jackson 1914. Austria, Carinthia, Großglockner 1700 m, pitfall trap, 1978, leg. Thaler.

Robertus ungulatus Vogelsanger 1944. Austria, Northern Tyrol, Innsbruck surroundings, Lanser Moor, fen, pitfall trap, 14 May–18 September 1963, leg. Thaler.

Rugathodes bellicosus (Simon 1873). Austria, Northern Tyrol, Obergurgl, 2600 m, 26 June 1992, leg. Thaler.

Simitidion simile (C.L. Koch 1836). Italy, Trentino, Civezzano, 30 April 1990, leg. Foddai.

Steatoda bipunctata (Linné 1758). Austria, Northern Tyrol, Innsbruck, Hötting, November 1992, leg. Knoflach.

Steatoda triangulosa (Walckenaer 1802). Italy, Toscana, Grosseto, Castiglione, 8 June 1987, leg. Thaler.

Theonoe minutissima (O. Pickard-Cambridge 1879). Germany, Kempten, Schorenmoos, 15 December 1974–17 May 1975, leg. Mendl.

Theridion conigerum Simon 1914. Germany, Oberharz, Ilsenhütte, June 1972, Staatliches Museum für Tierkunde Dresden, leg. Heimer.

Theridion nigrovariegatum Simon 1873. Switzerland, Unterengadin, Ramosch, 12 July 1987, leg. Thaler.

Theridion ohlerti (Thorell 1870). Austria, Northern Tyrol, Kühtai, 2200 m, 18 June 1992, leg. Bertrandi.

Theridion petraeum L. Koch 1872. Austria, Northern Tyrol, Innsbruck surroundings, Patscher-kofel, 2200 m, 7 July 1991, leg. Knoflach.

Theridion pictum (Walckenaer 1802). Austria, Northern Tyrol, Innsbruck West, university surroundings, 14 May 1992, leg. Knoflach.

Theridion pinastri L. Koch 1872. Austria, Northern Tyrol, Ötztal, Sautener Forchet, 26 June 1992, leg. Bertrandi.

Theridion sisyphium (Clerck 1757). Austria, Northern Tyrol, Innsbruck surroundings, Gnadenwald, 11 June 1991, leg. Bertrandi.

Figure 1. Two taxa each have three sclerites, but their homologies are ambiguous. They perform three different functions, indicated as F1–F3; occur in various relative positions (topology), and differ in color and shape (black or white, round or hexagon; special similarity). Depending on which criterion is primary, different primary homology hypotheses result. Under topology, r1 = a1, r2 = a2, and r3 = a3; under function, r1 = a1, r2 = a3, and r3 = a2; under special similarity, r1 = a3, r2 = a2, and r3 = a1. See text and Agnarsson and Coddington (in press) for explanation

Figure 2. Cladogram of Theridiidae (reproduced from Agnarsson 2004) labeled with subfamily and informal clade names, some of which refer to characters of the male palp. This cladogram forms the basis for the taxon choice in this study and is used to evaluate evolutionary hypotheses of palpal elements

Figure 3. Landmarks and descriptions of the major features and sclerites of the theridiid palp

Figures 4–7.—Dipoena melanogaster. 4, male palp mesal; 5, bulb removed from cymbium, ectal; 6, apical; 7, dorsal; note loops of sperm duct within T, MA and E

Figures 8–11.—Euryopis flavomaculata. 8, bulb removed from cymbium, mesal-dorsal; 9, ectal; 10, apical-dorsal; 11, ventral; conductor absent, note loop of sperm duct within MA

Figures 12–16.—12, Latrodectus geometricus C.L. Koch 1841; 13, Selkirkiella sp. note the bent and sharp tipped cymbial hook, a synapomorphic condition for Pholcommatinae, the tight juxtaposition of C and TTA is a synapomorphy of Selkirkiella; 14, Enoplognatha ovata; 15, Episinus maculipes Cavanna 1876, the huge and complexly folded C is a synapomorphy of Spintharinae; 16, Styposis selis Levi 1964, the ectal E with tip inside a TTA groove suggests affinities with Pholcommatinae. Scale bars: 12, 14, 15 = 100 μm; 13, 16 = 50 μm

Figures 17–23.—17–19, Steatoda americana (Emerton 1882). 17, male palp ventral expanded; 18, bulb dorsal, removed from cymbium; 19, tibia dorsal view; 20, 21, Steatoda albomaculata (De Geer 1778), 20, palp ventral; 21, bulb removed from cymbium, dorsal view (redrawn from Knoflach 1996a); 22, Episinus angulatus, bulb removed from cymbium, ventral (redrawn from Knoflach 1993b), 23, mesal. 22, 23 reproduced from Agnarsson (2004) with permission from Blackwell Publishing

Figures 24–27.—Episinus truncatus. 24, bulb removed from cymbium, mesal; 25, ventral; 26, ectal; 27, dorsal; a third tegular sclerite ETS is present; note convoluted sperm duct within E and loop within MA; ventral tegulum conducts a part of the distal E

Figures 28–32.—Crustulina guttata. 28, distal bulb, ectal; 29, palp expanded, ventral; 30, bulb removed from cymbium, apical; 31, cymbium, ventral; note large, mesal cymbial process; 32, bulb removed from cymbium, mesal-dorsal; embolus bears numerous processes

Figures 33–38.—Steatoda bipunctata. 33, bulb slightly expanded and removed from cymbium, ectal; 34, dorsal; 35, ventral; sperm duct narrows when leaving T, then widens within MA and again becomes constricted when passing to E; 36, tip of embolus; 37, tip of cymbium, ventral; 38, tip of cymbial hook

Figures 39–42.—Steatoda phalerata (Panzer 1801). 39, bulb removed from cymbium, ectal; 40, apical-ventral; 41, mesal; 42, ventral

Figures 43–47.—Steatoda triangulosa. 43, bulb removed from cymbium, mesal; 44, ectal-dorsal; 45, apical; 46, ventral; 47, tip of cymbium, ventral; note tegular pit in 44 close to conductor, into which an embolar process articulates

Figures 48–58.—48, Enoplognatha gemina Bosmans & van Keer 1999, palp ventral (redrawn from Levy 1998; sub E. mandibularis (Lucas 1846)); 49, Phoroncidia americana (Emerton 1882), palp loosened from cymbium, ectal (redrawn from Levi & Levi 1962); 50, Carniella schwendingeri Knoflach 1996, palp ectal (redrawn from Knoflach 1996b); 51, Enoplognatha sp. expanded; 52, 53, Argyrodes argyrodes (Walckenaer 1842) (redrawn from Saaristo 1978). 52, ventral; 53, schematic of bulb removed from cymbium and E removed from T; G-I, Anelosimus vittatus (C.L. Koch 1836). 54, palp ectal; 55, sperm duct trajectory, see Agnarsson (2004) for nomenclature and discussion; 56, schematic look at duct loops; 57, 58, Anelosimus sp. 57, palp ventral; 58, sperm duct trajectory. 50, 57, 58 reproduced from Agnarsson (2004) with permission from Blackwell Publishing

Figures 59–65.—59, 60, 64 Argyrodes elevatus Taczanowski 1873 male palp. 59, apical; 60, dorsal; 61, Neosphintharus trigonum (Hentz 1850), palp ventral; 62, 63, Anelosimus eximius. 62, apical view of mesal side, note strong cymbial incision, an Anelosimus synapomorphy; 63, apical view of ventral side, showing clearly the C coming out of the base of the SC; 64, hooked bulb to cymbium lock system; 65, Anelosimus sp. hooded BC-lock system. Rather than representing independent lines of evolution the hooded system is derived from the hooked one (see Fig. 201). Scale bars: 59–63 = 100 μm; 64, 65 = 50 μm

Figures 66–69.—66, 68, Enoplognatha ovata. 66, ventral; 68, ectal; 67, 69, E. latimana. 67, ventral; 69 ectal; note TTA conducting E in the unexpanded palp

Figures 70–74.—70, 71, 73, 74, Enoplognatha ovata. 70, 73, 74, bulb slightly expanded and removed from cymbium; 70, ventral; 71, cymbium and tibia, dorsal; 72, Enoplognatha latimana, cymbium and tibia dorsal; 73, “naturally” expanded, ventral-mesal, note embolus shifted into furrow of conductor; 74, apical, note third tegular sclerite

Figures 75–78.—Enoplognatha thoracica. 75, 76, 78, male palp slightly expanded, ventral, ectal, mesal. 77, distal palpal sclerites, ectal

Figures 79–82.—Robertus neglectus. 79, 80, male palp expanded, ectal-apical, ectal; 81, distal sclerites, ectal; only one tegular apophysis present; 82, embolus

Figures 83–86.—Robertus scoticus. 83–85, male palp, ventral, ectal, mesal; 86, distal palpal sclerites

Figures 87–90.—Robertus ungulatus. 87, 88, male palp, mesal, ventral; 89, median apophysis; 90, distal palpal sclerites; two tegular apophyses present

Figures 91–95.—Pholcomma gibbum. 91, bulb slightly expanded and removed from cymbium, ectal; 92, ventral; conductor hyaline, slender and apparently without guiding function, whereas MA and TTA show a broad groove, which presumably supports the embolus; 93–95, distal bulb, removed from cymbium, 93, mesal-dorsal; 94, apical-dorsal; 95, caudal-ventral; note loop of sperm duct within MA

Figures 96–99.—Theonoe minutissima. 96, 97 male palp, ventral-mesal, ectal; 98, bulb, ventral; note distal process of cymbium and distinct constriction of sperm duct within T; embolus and median apophysis probably fused; 99, Kochiura aulica, cymbium ectal, largely excavated

Figures 100–103.—Kochiura aulica. 100, male palp, ectal; 101, 102, bulb expanded, dorsal, apical; modified tibial and cymbial setae support the embolus, whereas conductor is comparatively inconspicuous; 103, embolus separated from palp, coiling up in spirals; its remarkable length measures three times the male's total body length

Figures 104–107.—Male palps ventral; 104, Theridion varians Hahn 1833, note furcated MA, typical of Theridion and relatives; 105, Theridion frondeum Hentz 1850, note also tegular pit (arrow) involved in a lock mechanism with E via a embolic apophysis (see also Avilés et al. (2006, fig. 4) for SEM photographs of the closely related T. nigroannulatum); 106, Ameridion sp. like all theridiines with a hooded BC-lock system (arrow); 107, Thymoites nr. prolatus (Levi 1959), the grossly enlarged tibial rim is shared with some other Thymoites. All scale bars = 100 μm

Figures 108–112.—108, Ameridion sp., arrow indicates tip of MA; 109, Achaearanea tepidariorum, like most other Achaearanea the TTA has been lost. Note the seam in the embolus (or possibly the fusing point between the E and MA, see discussion); 110, Theridula opulenta (Walckenaer 1842), among the simplest palps of theridiids, the TTA has been lost (independently from the loss in Achaearanea, see Fig. 201), the C is also absent, while a membranous connection exists between the T distally and cymbium. This membrane is possibly a homolog of the MA; 111, Ameridion sp., note elongated palpal femur; 112, Thymoites nr. prolatus, here not only the femur, but also the palpal patella is grossly elongated. Scale bars: 108, 109, 111, 112 =100 μm; 110 = 10 μm

Figures 113–120.—113–115, Theridion cochise Levi 1963 dissected palp. 113, cymbium; 114, bulb ventral, absence of TTA and flat based E are shared with some other Theridion, e.g.,T. grallator Simon 1900; 115, bulb dorsal; 116, Coleosoma floridanum Banks 1900, schematic drawing of a dissected palp, MA is present, but not shown; 117, Theridion frondeum, palp ventral; 118, Achaearanea trapezoidalis (Taczanowski 1873), the type species of Achaearanea, uniquely among theridiines, has a hooked BC-lock system (arrow); 119, Achaearanea tabulata Levi 1980 (redrawn from Knoflach 1991), like most Achaearanea lacks TTA; the MA is either lost as well or fused with the embolus; 120, Theridula emertoni Berland 1920 (redrawn from Levi & Levi 1962), lacks a conductor, TTA is also absent. The tegulum is distally attached to the cymbium via a membranous sclerite, most likely the MA

Figures 121–124.—Achaearanea lunata. 121–123, bulb slightly expanded and removed from cymbium, ventral, ectal, mesal; the TTA has been lost; it is uncertain whether the MA has been fused with the embolus, or lost, in which case the embolus base interacts with the BC-lock system; 124, distal ectal margin of cymbium in ventral view with cymbial hood

Figures 125–129.—Achaearanea riparia. 125–127, bulb slightly expanded, ventral, mesal, ectal-dorsal; 128, apical; conductor with scaly surface as present in many Achaearanea species. 129, distal cymbium in ventral view, with protruding tip

Figures 130–133.—Keijia tincta. 130, 132, 133, bulb slightly expanded, ventral, ectal, dorsal; 131, male palp, ventral; both tegular apophyses present; note tegular pit and corresponding embolar process (arrow)

Figures 134–138.—Neottiura bimaculata. 134–137, bulb removed from cymbium and slightly expanded, apical-ectal, ectal, apical-dorsal, ventral; three tegular apophyses present, which are complexly folded and connected by a large membrane; 138, tegulum, dorsal; note convoluted course of sperm duct within T

Figures 139–144.—Rugathodes bellicosus. 139, 141, bulb removed from cymbium and slightly expanded, dorsal, ectal; both tegular apophyses present; 140, male palp in ventral view; 142, distal cymbium in ventral view with cymbial hood; 143 embolus removed from bulb. 144, median apophysis, mesal. Embolus submerged deeply into tegulum, only its tip being free, accompanied by the conductor; its articulation into the tegular pit also inside but visible through tegulum (arrow)

Figures 145–148.—Simitidion simile. 145, 147, 148, bulb removed from cymbium and slightly expanded, mesal, dorsal, apical-ventral; sperm duct forms numerous coils within tegulum; 146, male palp, ectal; both tegular apophyses present, connected by a membrane

Figures 149–152.—Theridion pictum. 149, 151, 152, bulb removed from cymbium and slightly expanded, ventral, dorsal, mesal; both tegular apophyses present, both without sperm duct; conductor with broad, short channel supporting the embolus; scales on TTA indicate contact to the female epigynum; 150, male palp, ventral; arrow points to tegular pit

Figures 153–156.—Theridion conigerum. 153, embolus, ventral; distal part corrugated; 154, 156, bulb removed from cymbium and slightly expanded, ventral, apical-mesal; 155, distal bulb, dorsal; conductor lobe-like, forming a fold

Figures 157–160.—Theridion ohlerti. 157, male palp, ventral; embolus hidden by cymbium; 158–160, bulb removed from cymbium and slightly expanded, ventral, dorsal, apical-mesal; TTA small and submerged into tegulum; distal embolus corrugated; tegulum with distinct lobe close to conductor

Figures 161–164.—Theridion petraeum. 161, 162, male palp, mesal, ectal; 163, 164, bulb removed from cymbium and slightly expanded, dorsal, apical-mesal; conductor bifid, containing a groove and channel for embolus; ventral end of MA typically sickle-shaped

Figures 165–168.—Theridion pinastri. 165, 167, 168, bulb removed from cymbium and slightly expanded, apical-ventral, mesal, dorsal; the TTA is relatively small; 166 male palp, ventral; note articulation between embolus and tegulum

Figures 169–172.—Theridion sisyphium. 169, 171, 172, bulb removed from cymbium and slightly expanded, ventral, apical-mesal, dorsal; TTA small and bifid; 170, male palp, ventral

Figures 173–188.—Distal cymbium with cymbial hook, 173–182, and hood, 183–188. 173, Lasaeola tristis; 174, 175, Steatoda phalerata; 176, Dipoena melanogaster; 177, Euryopis flavomaculata; 178, 179, Pholcomma gibbum; 180, Episinus truncatus; 181, E. theridioides; 182, Robertus neglectus; 183, 184, Neottiura bimaculata; 185, Theridion nigrovariegatum; 186, Keijia tincta; 187, Simitidion simile; 188, Theridion sisyphium

Figures 189–194.—189, 190 Synotaxus monoceros (Caporiacco 1947) (Synotaxidae). 189, ventral, note patellar spur (arrow), a rather uniform tibia, and a large excavate TTA; 190, ectal, the E is a direct outgrowth of the tegulum, note also a distinct, cup shaped PC; 191, S. waiwai paracymbium; 192, Nesticus silvestrii Fage 1929, huge and rigid PC; 193, 194, Euryopis gertschi Levi 1951. 193, ventral, conductor absent; 194, ectal, note small membrane between T and E. Scale bars: 189, 190, 192–194 = 100 μm; 191 = 20 μm

Figures 195–200.—195, Pimoa rupicola (Simon 1884) (PEP = pimoid embolic process); 196, Linyphia triangularis (EM = embolic membrane, LC = lamella characteristica, R = radix, SPT = suprategulum, TA = terminal apophysis); 197, Synotaxus monoceros; 198, Nesticus cellulanus (Clerck 1757); 199, Eidmanella pallida, part of tegulum showing MA and E; 200, Euryopis flavomaculata (redrawn from Levi & Levi 1962). 195–199 reproduced from Agnarsson (2004) with permission from Blackwell Publishing

Figure 201. Evolutionary changes in the unique theridiid BC-lock system. The cladogram is taken from Agnarsson (2004), see Figure 2 for clade names. Theridiidae is indicated with bold lines. Black horizontal bars each indicate one instance of homoplasy: Spintharus independently evolved a hooded lock system and lacks the hood on the MA; Phoroncidia lacks the MA hood; the MA of Pholcomma hirsutum Emerton 1882 does not contain a loop of the sperm duct (but in P. gibbum does, see Figs. 91–95); Kochiura rosea (Nicolet 1849) lacks the MA hood, Tidarren has a hook BC lock system (uniquely among Theridiinae); the lock system of Theridula is unique (see text)

table

Table 1. Results of the method outlined in Fig. 1 and the text, as applied to three problematic theridiid palpal sclerites: median apophysis (MA), theridioid tegular apophysis (TTA), and conductor (C) with topology as the primary criterion. Similar tables were compiled for function and special similarity and are summarized under the SS and FNC columns in Table 2. Scores are given for topology (TOP), special similarity (SS) and function (FNC) as secondary criteria. Dashes are inapplicables; question marks are unknowns. Special similarity includes three points of comparison: flexible or fused tegular connection (Cxn), sperm duct presence or absence (Dct), and membranous or sclerotized texture (Tex), which three scores are averaged under SS for each sclerite under the equal weights point of view (see text). The strict gain/loss point of view (see text) is tabulated in the G/L column. As the primary criterion, topology naturally does not conflict with itself as a secondary criterion (all scores = 1, or agreement) but it conflicts with function for the TTA and C, and with special similarity for MA and C. Subtotals by taxon (averages for TOP, SS, FNC, and G/L) and counts of conflict for parsimony (PAR) appear at right; grand totals are counts or averages of raw scores under each sclerite and are carried forward to Table 2

table

Table 1. Extended.

table

Table 2. Results of the logic of Table 1 as applied to four prior analyses of theridiid palpal homologies (A04 = Agnarsson 2004; C90 = Coddington 1990; L62 = Levi 1953–1973; S78 = Saaristo 1978) and for all three primary criteria (TOP, FCN, SS). The column TOP carries forward the grand totals of Table 1. Either as mean performance under all criteria and accounting for gain/loss hypotheses (Grand mean) or simple step counting (parsimony) topology (TOP) as applied by Agnarsson (2004) outperforms other criteria and previous homology hypotheses

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