C. D. Dondale and J. H. Redner.
Revision of the milvina group of the wolf spider genus Pardosa (Araneae: Lycosidae).
Psyche 91:67-118, 1984.

Full text (searchable PDF, 6916K)
Durable link: http://psyche.entclub.org/91/91-067.html

The following unprocessed text is extracted from the PDF file, and is likely to be both incomplete and full of errors. Please consult the PDF file for the complete article.

REVISION OF THE MILVINA GROUP
OF THE WOLF SPIDER GENUS PARDOSA
(ARANEAE: LYC0SIDAE)l
BY C. D. DONDALE AND J. H. REDNER
Biosystematics Research Institute
Research Branch, Agriculture Canada,
Ottawa, Ontario, Canada KIA OC6
The genus Pardosa C. L. Koch is a large and widespread group of wolf spiders. Several species groups have been recognized in North America (Lowrie and Dondale 1981). The objective of the present paper is to define and revise the milvina group. The milvina group of the genus Pardosa consists of 18 American species of wolf spiders. The greatest concentration of species seems to be in the southeastern United States and Mexico. Only P. milvina (Hentz), P. saxatilis (Hentz), and P. littoralis Banks range northward as far as Canada, and only P. portoricensis Banks, P. hamifera F. Pickard-Cambridge, and P. littoralis have been found in the West Indies. P. fastosa (Keyserling) extends into South America as far as Ecuador. None are found on the U.S. Pacific coast or in the U.S. Rocky Mountains, and few specimens are recorded from the Great Plains.
Terminology for anatomical parts is defined by Dondale and Redner (1978) and by Figures 2-7 and 41-45 here. Measurements are given as the mean and standard deviation for 10 to 20 specimens or as the range for fewer.
The external genitalia, in our opinion, provide the best indicators of relationships within the milvina group, The inferred relationships among the species are shown in Figure 1. Females of all species in the group have the lateral margins of the median septum raised and thickened at the site of the copulatory openings (character 1, Fig. 1; Fig. 45). This modification appears to enlarge the openings and to reinforce the 1Monuscript received by the editor December 10, 67
margins, and we speculate
1983.
Pnchs 91:67-118 11084). htlpJ/psychc mtclubor@i/SI-Oe7 hlml



================================================================================

Psyche
[Vol. 91




================================================================================

19841 Dondale & Redner - Genus Pardosa 69 that each opening may accommodate not only the embolus, which is a slender shaft, but the enlarged, sclerotized tip of the conductor as well. The condition is not found elsewhere in Pardosa except in two species of the Old World wagleri group, which have a quite different kind of median septum, hood, and copulatory tubes. We infer the condition found in the milvina group to be apomorphic, and the absence of the condition to be plesiomorphic. The base of the median septum is more or less rectangular in females of certain species of the group (character 4, Fig. 1; Fig. 50). This condition is unique among the species of Pardosa, and is inferred to be apomorphic. The plesiomorphic state is a triangular base. Also, a strongly tapered anterior part of the septum (character 9, Fig. 1; Fig. 65), in combination with a single hood cavity, is inferred to be apomorphic and a broader anterior part to be plesiomorphic.
The epigynal hood may extend posteriad at the sides, thus defining a raised, tapered, median area (character 12, Fig. 1; Fig. 58). This condition, found only in specimens of the saxatilis complex, is inferred to be apomorphic. The simpler, lobed hood found in the other members of the milvina group is inferred to be plesiomorphic.
The median apophysis, which is believed to assist indirectly in aligning the embolus tip with the copulatory opening, may be unusually short and blunt (character 2, Fig. 1; Fig. 3), with its basal process located at midlength on the apophysis (character 5, Fig. 1; Fig. 8) and having its mesal margin thickened (character 7, Fig. 1; Fig. 8). These states are inferred to be apomorphic and the more slender, elongate apophysis, with the basal process located at the base and with the mesal margin not thickened, to be plesiomorphic. The conductor in the milvina group is a stout rod, lying transversely in the groove between the apical division of the genital bulb and the tegulum. In the unexpanded bulb, only its tip is visible (Fig. 3), but it becomes fully exposed by dissection (Fig. 6). The apomorphic states of this character are inferred to be "angulate or pointed on basal margin" (character 3, Fig. 1; Fig. 6), "fluted at tip" (character 6, Fig. 1; Fig. 1 I), "possession of dark, shiny knob near tip" (character 8, Fig. 1 ; Fig. 21), and "knob excavated" (character 10, Fig. 1; Figs. 38, 40, arrows). The plesiomorphic states are inferred to be "straight basal margin", "thick tip", "knob lacking", and "knob not excavated".




================================================================================

70 Psyche [vo~. 91
Certain segments of the male palpus may be covered dorsally with reflective white setae (characters 1 1, 1 1 a, Fig. 1 ; Figs. 16- 18).These setae probably serve as a visual cue to the female during male courtship, and their presence is believed to be apomorphic. In saxatilis, only the patella bears such a covering; in atlantica, both patella and tibia are covered, and inparvula, both segments plus the basal half of the cymbium are covered. Males of only the remotely related P. tesquorum (Odenwall) have a similar covering on the palpal patella. We infer that the presence of this covering in the milvina group is apomorphic, and that the plesiomorphic state of the character is the involvement of a single segment. A name associated in the past with the milvina group, namely, Lycosa canadensis Blackwall, 1 87 1, treated as a junior synonym of P. milvina by Charnberlin (1908), could not be identified by us, and is treated here as nomen dubium. The holotype of cana- densis was an unidentifiable juvenile and is no longer in exist- ence. Pardosa accurata Becker, 1886, described from "Mexico", has been identified as an unknown species of Lycosa, s.l., through examination of the holotype by us; the type is a juvenile speci- men deposited in the Institut Royal des Sciences Naturelles de Belgique.
Total length 3.00 to 7.50 mm. Carapace smoothly convex at sides, vertical anteriorly at sides, approximately uniform in height between dorsal groove and posterior row of eyes, covered sparsely with short setae; eye area black, often iridescent; anterior row of eyes somewhat procurved, distinctly shorter than middle row, with median eyes equal in size to lateral eyes or somewhat larger and lo- cated slightly closer to laterals than to each other. Legs moderately long and strong, with thin scopulae and sparse setae; femur I with 3 dorsal macrosetae, 2 prolaterals (near tip), 2 or 3 retrolaterals; tibia I with 2 slender bristlelike dorsals, 2 prolaterals, 2 retrolaterals, 3 pairs of ventrals; basitarsus I with 1 bristlelike dorsal, 3 prolaterals, 2 or 3 retrolaterals, 3 paired and 1 unpaired ventral; trochanters with deep notch distally at tip. Abdomen ovoid, covered with dense, short setae and with cluster of longer, erect setae at anterior end.



================================================================================

19841 Dondale & Redner - Genus Pardosa 71 Male palpus usually black and contrasting with pale areas on legs and carapace, often with fringes of black setae at sides of segments; palea (Fig. 6) projecting liplike at tip of genital bulb, giving rise prolaterally to slender, rather short, slightly curved embolus and giving rise basally to elongate, stout conductor that lies largely hidden between base of palea and tegulum; terminal apophysis toothlike or bladelike, located retrolaterodistally (Fig. 6); median apophysis short, straight or somewhat curved, with strong basal process (Fig. 3).
Epigynum (Fig. 41) rather long (often longer than twice maximum width of median septum), with median septum broad posteriorly and slender or evanescent anteriorly; hood small, distinct, with single cavity; copulatory openings with margins raised and thickened. Copulatory tubes (Fig. 44) short, angulate or somewhat curved, with swelling on ventral or lateral side; sperma- thecae small, bulbous.
KEY TO MALES OF THE MILVINA GROUP
Distal process of median apophysis short, broad, lacking hook at tip (Fig. 3) ............................ -2 Distal process of median apophysis elongate, usual1 y hooked at tip (as in Fig. 9, arrow) ............... -4 Terminal apophysis broad (ventral view, Figs. 2, arrow, 5); conductor smoothly curved on basal margin (Fig. 5) ............................. bellona Banks Terminal apophysis more slender (ventral view, Figs. 6, 7); conductor angular or pointed on basal margin (Figs. 6, 7, arrows) ...................................... 3 Conductor with hooked point on basal margin (Fig. 6); ..................... embolus bent near tip (Fig. 6) ..................... delicatula Gertsch and Wallace Conductor with angulate basal margin (Fig. 7); embolus gently curved at tip (Fig. 7) ....................... ................. hamifera F. Pickard-Cambridge Median apophysis with basal process small, located about midlength of apophysis (Fig. 8); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 11) ......- 5 Median apophysis with basal process located at base of apophysis or, if located at middle then large (Figs. 19,24);



================================================================================

Psyche [vo~. 91
conductor thickened at tip, bearing dark, shiny, knoblike process near tip (as in Figs. 21, 26, 38) ........... .8 Median apophysis with distal process straight at tip (Fig. 8, arrow) ................ sagei Gertsch and Wallace Median apophysis with distal process hooked at tip (as in Figs. 9, 10, 14) ................................. .6 Terminal apophysis separated from mesal swelling by acute angle (Fig. 12); median apophysis with distal process strongly hooked (Fig. 9, arrow) ............. ........................... fastosa (Keyserling) Terminal apophysis separated from mesal swelling by broad curve (Fig. 13, arrow); median apophysis with distal process weakly hooked (Fig. 10) ............ .7 Median apophysis with marginal swelling broad (Fig. 10, arrow) ............... desolatula Gertsch and Davis Median apophysis with marginal swelling narrow (Fig. 14, arrow) ........................... mayana spa. Patella (and sometimes additional segments) of palpus covered dorsally with reflective white setae (most notice- able in living or dried specimens) (Figs. 16-18); conductor sinuous along basal margin (Fig. 21, arrows) ...... .9 Patella of palpus covered dorsally with dark setae (though few reflective white setae may be present); conductor with single curve along basal margin (as in Figs. 25, 38). .. ............................................... 11 Tibia (as well as patella, and sometimes basal half of cymbium) covered dorsally with reflective white setae (Figs. 17, 18) ................................. .10 Tibia of palpus covered with dark setae (Fig. 16) .... ............................... saxatilis (Hentz) Basal half of cymbium (except narrow band at basal margin) covered dorsally with dark setae (Fig. 17). ... ............................ .atlantica Emerton Basal half of cymbium covered with reflective white setae (Fig. 18) ........................... .parvula Banks Terminal apophysis large, extending basad to or beyond tip of embolus (Figs. 22, arrow, 25) ................ .............................. littoralis Banks Terminal apophysis smaller, extending only short dis- tance basad (as in Figs. 24, 35, 42) ............. .12



================================================================================

19841 Dondale & Redner - Genus Pardosa 73 Terminal apophysis arched retrolaterally (Figs. 23, arrow, 26); median apophysis small, occupying about one-third length of genital bulb (Fig. 23) ....... .saltonia sp.n. Terminal apophysis arched mesally, or not arched (Figs. 27, 34, 40); median apophysis larger, occupying distinctly more than one-third of genital bulb (Figs. 24, 31, 42). ............................................ 13 Terminal apophysis with mesal swelling toothlike (Fig. 27, arrow); median apophysis thick throughout most of its length, with basal process swollen (Fig. 24) .......... ........................... pauxilla Montgomery Terminal apophysis with mesal swelling not toothlike, i.e., lacking sharp point (Fig. 34); median apophysis slender at middle (Figs. 28, 37) ........................... .14 Median apophysis with distal process expanded (ventral view, Fig. 28, arrow). Dorsum of abdomen with pale median band flanked by paired distinct, dark, longi- tudinal bands .................. portoricensis Banks Median apophysis with distal process more slender (Figs. 31, 37, 42). Dorsum of abdomen dark or mottled, or, if pale mesally then lacking paired dark longitudinal bands ........................................... 15 Terminal apophysis broad, blunt (Fig. 30, arrow). Species occurring only north of Tropic of Cancer (Map 5) ... ............................. .milvina (Hentz) Terminal apophysis more slender, pointed (Figs. 33, 36, 39). Species occurring only south of Tropic of Cancer (Map 6) ....................................... 16 Median apophysis abruptly angled on mesal margin (Fig. 35, arrow). Carapace width less than 1.35 mm. Dorsum of abdomen with median band of reflective white setae . . .......................... guadalajarana sp.n. Median apophysis curved on mesal margin (Figs. 37,42). Carapace width greater than 1.35 mm. Dorsum of abdomen lacking median band of reflective white setae ........................................ 17 Terminal apophysis rather long (Fig. 36). Tibia and basitarsus I with fringe of long, erect setae along prolateral and retrolateral surfaces. Carapace with black lateral margins ..... longivulva F. Pickard-Cambridge



================================================================================

Psyche [vo~. 91
Terminal apophysis shorter (Fig. 39). Tibia and basitarsus I lacking fringe. Carapace usually with pale lateral ..........................
margins marialuisae sp.n.
KEY TO FEMALES OF THE MILVINA GROUP
OCCURRING IN EASTERN CANADA AND UNITED STATES Median septum extending anteriad nearly to level of hood ...............................
(Figs. 46, 60, 65) .2
Median septum extending anteriad one-half length of ..................
epigynum or less (Figs. 58, 70) .4
Median septum broad anteriorly (Fig. 60, arrow) ..... ............................. littoralis Banks Median septum distinctly tapered anteriorly (Figs. 46, 65) ................................................ 3 Median septum with expanded posterior part concave at lateral margins (Fig. 65, arrow) pauxilla Montgomery Median septum with expanded posterior part convex at lateral margins (Fig. 46, upper arrow) .............. ................. delicatula Gertsch and Wallace Hood continuing posteriad at sides where it defines a raised, tapered median area (Fig. 58, arrows) ...... .5 Hood continuing posteriad at sides where it defines a .........
depressed, non-tapered area (Fig. 70, arrow) .................................. .milvina (Hentz) Species restricted to Great Lakes-St. Lawrence region and southward in Appalachian Mountains (Map 3) ....... ........................... saxatilis (Hentz) Species restricted to southeastern coastal plain (including Florida) and the Mississippi basin (Map 3) ....... .6 Species restricted to Florida and eastern Gulf Coast region (Map 3) ..................... .parvula Banks Species restricted to Atlantic coast States east of the Appalachians and in the Mississippi basin (Map 3) ... ................................ .atlantica Emerton KEY TO FEMALES OF THE MILVINA GROUP
OCCURRING IN SOUTHWESTERN UNITED STATES, MEXICO. AND CENTRAL AMERICA
Median septum with expanded posterior part approxi- mately rectangular (Figs. 50, 52, 54, 56) ............



================================================================================

Dondale & Redner - Genus Pardosa 75
........ desolatula Gertsch and Davis, mayana sp.n., sagei Gertsch and Wallace, and fastosa (Keyserling) Median septum with expanded posterior part more or less triangular (Figs. 48, 62, 68) ..................... .2 Median septum broad anteriorly, bordered by pair of ridges that extend posteriad along interior surface of septum (Figs. 41, arrow, 43, 44) ...... bellona Banks Median septum slender or evanescent anteriorly, lacking bordering ridges ................................ .3 Epigynum with curved ridges posteriorly (Figs. 46, lower arrow, 48, 72, 74, 76) ........................... .4 ........... Epigynum lacking curved ridges (Fig. 62). ................................. saltonia sp.n. Curved ridges concealing lateral angles of median septum (Fig. 46, lower arrow) ............................ .............. delicatula Gertsch and Wallace Curved ridges not concealing lateral angles of median ......................................... septum 5 Epigynum short (ratio of epigynal length to greatest ................. median septum width less than 2:l) ................. hamifera F. Pickard-Cambridge Epigynum longer (ratio of epigynal length to greatest .............
median septum width more than 2:l)
.6
Carapace width usually less than 1.6 mm. Retromargin of chelicera with 2 teeth ........... guadalajarana sp.n. Carapace width usually greater than 1.6 mm. Retro- ................. margin of chelicera with 3 teeth .7
Copulatory tubes with small swelling at base (Fig. 75). Carapace margins dark ........................... ................ longivulva F. Pickard-Cambridge Copulatory tubes with large swelling on lateral margins ...... (Fig. 77, arrow). Carapace margins usually pale ............................ marialuisae sp.n. KEY TO FEMALES OF THE MILVINA GROUP
OCCURRING IN THE WEST INDIES
Median septum rather broad anteriorly (Fig. 60, arrow) .............................. littoralis Banks Median septum distinctly tapered anteriorly (Figs. 48, 68) ................................................ 2



================================================================================

76 Psyche [vo~. 91
Copulatory tubes broad, and spermathecae widely sepa- rated (Fig. 49). Epigynum long (ratio of epigynal length to greatest median septum width more than 2:l) ........ ................... hamifera F. Pickard-Cambridge Copulatory tubes slender, and spermathecae narrowly separated (Fig. 69, arrows). Epigynum shorter (ratio of epigynal length to greatest median septum width less than 2: 1) ........................... portoricensis Banks Pardosa bellona Banks
Figures 2, 5, 41, 43, 44; Map 1
Pardosa bellona Banks, 1898:275, Fig. 21 (pi. 16). Two male and two female syntypes from Coral de Piedras, Baja California (Eisen and Vaslit), deposited in MCZ, examined. Syntypes from San Miguel de Horcasitas, Magdalena Island, and from San Jose del Cabo, deposited in California Academy of Sciences, presumed destroyed. Gertsch 1934:21. Gertsch and Wallace 1935:3, Fig. 14. Roewer 1954: 189. Bonnet 1958:3359.
Male.
Total length 4.50 k0.34 mm; carapace 2.28 k0.18 mm long and 1.75 k0.13 mm wide (20 specimens). Carapace with dark orange median and submarginal areas, and with pair of broad, dark longitudinal bands bordering pale median area; margins dark. Sternum yellow orange, sometimes with several small black spots. Chelicerae orange brown to black, pale mesally; retromargin with 3 teeth. Legs yellow orange; femur I often black basally; tibia and basitarsus I with sparse fringe of long lateral setae. Abdomen pale mesally, dark laterally; venter orange yellow. Terminal apophysis broad; median apophysis short, broad, with large basal process, lacking hook at tip of distal process (Fig. 2); conductor rounded on basal margin (Fig. 5).
Female.
Total length 4.85 k0.35 mm; carapace 2.34 k0.17 mm long and 1.86 k0.13 mm wide (18 specimens). General color and structure as in male but dark pigment on carapace margin may break into spots, femur I lacking dark pigment at base, and leg I lacking fringe of long lateral setae. Median septum rather broad posteriorly, little tapered anteriorly, flanked anteriorly by ridges that continue along internal surface (Fig. 41); copulatory tubes rather thick, with swelling on ventrolateral surface (Figs. 43, 44). Diagnosis. Specimens of P. bellona most resemble those of P. delicatula and P. hamifera, differing by the broad terminal



================================================================================

19841 Dondale & Redner - Genus Pardosa 77 apophysis, by the rounded basal margin of the conductor, and by the anteriorly broad median septum with paired ridges. Range. Utah south to Colima, Mexico. Included, but not mapped here, are the following: Utah: Monroe Canyon, 112' lO'W, 38'38'N. Nevada: Las Vegas. California: Seeley, 7 mi. west of El Centro; Indian Wells, Riverside Co.
Natural History. Males have been collected in December to February, and June to September, females in January, March, June to September, and November. Females with egg sacs were collected in January, March, and May to July.
Pardosa delicatula Gertsch and Wallace
Figures 3, 6, 45-47; Map 1
Pardosa pauxilla: Montgomery 1904:268 (part, not lectotype). Pardosa delicatula Gertsch and Wallace, 19354, Figs. 13, 17. Male holotype and female allotype from Edinburg, Hidalgo Co., Texas (Stanley Mulaik), deposited in AMNH, examined. Gertsch and Davis, 1940: 5, Roewer 1954: 189. Bonnet l958:3365.
Male. Total length 4.78 k0.32 mm; carapace 2.59 k0.15 mm long and 1-90 k0.14 mm wide (20 specimens). Carapace with dark orange median and submarginal areas, and with pair of dark brown longitudinal bands flanking pale median area; margins dark, pale, or marked with series of 3 or 4 dark spots. Sternum orange yellow, sometimes suffused with black or with faint, V-shaped dark mark. Chelicerae variable in color; retromargin with 3 teeth. Legs yellow orange; femur I dark prolaterally at base. Abdomen dull yellow mesally, darker laterally, sometimes black throughout; venter pale. Terminal apophysis rather slender (ventral view); median apophysis short, broad, with large basal process, lacking hook at tip of distal process (Fig. 3); conductor with sharp, hooked point on ventral surface; embolus bent near tip (Fig. 6). Female. Total length 5.73 k0.79; carapace 2.63 k0.20 mm long and 2-02 k0.23 mm wide (20 specimens). General color and structure as in male but carapace paler, usually having pale margins. Median septum strongly tapered anteriorly, lacking lateral ridges, convex along lateral margins of posterior part, partly concealed laterally by curved ridges (Fig. 46); copulatory tubes rather slender (Fig. 47).




================================================================================

78 Psyche [VOL 91
Map 1.
Collection localities for Pardosa bellona (squares), P. delicatula (stars), P. hamfera (half-circles), P. littoralis (circles). Diagnosis. Specimens of P. delicatula most resemble those of P. bellona and P. hamifera but differ by the sharp, hooked point on the conductor, by the bent embolus, and by the concealment of the lateral angles of the median septum by curved ridges. Range. Oklahoma and Mississippi south to Chihuahua and Nuevo Leon, Mexico.
Natural History.
Males and females have been collected in every month except January, and females with egg sacs were taken from April to October. Common habitats are pastures and other grasslands, where they are taken in numbers by pitfall trap; one large collection was taken by sweep net in Louisiana sweet potato fields at night.




================================================================================