Allocosa senex ( Mello-Leitão, 1945 ), Mello-Leitao, 1945

Simó, Miguel, Lise, Arno A., Pompozzi, Gabriel & Laborda, Álvaro, 2017, On the taxonomy of southern South American species of the wolf spider genus Allocosa (Araneae: Lycosidae: Allocosinae), Zootaxa 4216 (3), pp. 261-278 : 267-270

Allocosa senex ( Mello-Leitão, 1945) revalidated

Figs 3 , 5 C–D, 6C–D, 7A–E, 8B, 9

Glieschiella senex Mello-Leitão, 1945: 254 (male holotype from Colón , Entre Ríos, Argentina (32°12´S, 58°8´W), M. Birabén leg., XI/1941, MLP 16470, specimen very damaged, examined).

Allocosa brasiliensis: Capocasale 1990: 133 , figs 1–5 (misidentification); Capocasale, 2001: 272, figs 8–10 (misidentification).

Remarks. Capocasale (2001) considered A. senex as a junior synonym of A. brasiliensis based on the similar morphology of the terminal and median apophyses of the male bulb. After examination of the type specimens, we concluded they are not coespecific due to clear differences in the body size, colouration, the shape of the male cymbium and internal genital morphology of the female.

Other material examined. Argentina. Buenos Aires. 10 km NE Quequén (38°34'S, 58°40'W), 2.IX.1972 , 1 female ( MACN Ar 36342); General Madariaga (37°1'S, 57°7'W), 13.IV.1951, A . Bachmann leg., 1 male, 2 immatures (MACN Ar 3 234); XII.1952, Orfila, De Carlo & Di Amico leg., 1 female, 1 immature ( MACN Ar 3738); Entre Ríos. Parque Nacional El Palmar (31°52'S, 58°12'W) , 18.IV.1981, Zanetic & P. Golobbof leg, 1male, 1 female (MACN Ar 34581); 22–23.XI.2003, C. Grismado, A. Ojanguren & F. Labarque leg., 2 females ( MACN Ar 34582). Córdoba. Copina (31°34'S, 64°39'W) , 26.XI.2010, 1 male (FCE Ar 1057); 1 male (FCE Ar 1058); 1 male (FCE Ar 1059); 1 male (FCE Ar 1060); 1 male (FCE Ar 1061); 1 male (FCE Ar 1062); 1 male (FCE Ar 1063); 1 male (FCE Ar 1064); 1 female (FCE Ar 1065); 1 female (FCE Ar 1066); 1 female (FCE Ar 1067); 1 female (FCE Ar 1068); 1 female (FCE Ar 1069); 1 female (FCE Ar 1070); 1 female (FCE Ar 1072), all collected by G. De Simone; Nono (31°47'S, 65°00'W) , 08.I.2011, 1 female (FCE Ar 1073); 1 male (FCE Ar 2905); 1 female (FCE Ar 2906); 1 female (FCE Ar 2907); 1 female (FCE Ar 2908); 1 female (FCE Ar 2909); 1 female (FCE Ar 2910); 1 female (FCE Ar 2911); 1 female (FCE Ar 2912); 1 female (FCE Ar 2913), all collected by G. De Simone; Santa Fé, Laguna Setúbal (31°35'S, 60°36'W), 10.III.1950, G. Martínez leg , 1 male (MACN Ar 36343). Brazil. Río Grande do Sul. Barra do Ribeiro, Parque Municipal (30°17'S, 51°18'W), VII.2006, L.E. Costa-Schmidt leg. , 2 females (FCE Ar 2392). Pelotas , Colônia Z3, camping (31°41'S, 52°09'W), 11.IX.2006, L.E. Costa-Schmidt leg. , 2 males, 1 female, 2 immatures (FCE Ar 2393); Pelotas, Laranjal , paleodunas (31°46'S, 52°15'W), 11.IX.2006, L.E. Costa-Schmidt leg. , 2 males, 8 females, 2 immatures (FCE Ar 2394); Tapes, Santuário (30°41'S, 51°23'W), IX.2006, L.E. Costa-Schmidt leg. , 2 females (FCE Ar 2396); Tapes, Portal Tapes Hotel , (30°39'S, 51°23'W), IX.2006, L.E. Costa-Schmidt leg. , 2 females, 1 immature, (FCE Ar 2397); Rio Grande, Praia do Cassino , (32°12'S, 52°11'W), 12.IX.2006, L.E. Costa-Schmidt leg. , 2 males, 12 females (FCE Ar 2398); Cidreira (30°10'S, 50°12'W), 8.II.2003 , 1 female (MCTP 16514); 23.II.2003, 1 female (MCTP 16512); 4.III.2003, 2 males (MCTP 16 517); 3.III.2003, 1 male ( MCTP 16516 ), all collected by J. A. M. Pinto; Viamão, Itapuã (30°17'S, 51° 1'W) , 22.IX.2003, 1 female (MCTP 16511); 10.II.2004, 1 male (MCTP 16507); 13.I.2004, 1 female (MCTP 16504); 26.I.2004, 1 female (MCTP 16505); 9.III.2004, 1 female (MCTP 16 508); 10.XI.2004, 1 male (MCTP 16506); 13.I.2004, 1 male ( MCTP 16510 ), all collected by A.C.K. Ferreira; Pinhal–Cidreira (30°13'S, 50°13'W) , 23.X.1992, R. M. Alberto leg., 1 female, 2 immatures ( MCTP 2692 ). Uruguay. Canelones. El Pinar (34°47'S, 55°52'W) , 27.I.2008, R. Postiglioni leg., 1 male (FCE 2348). Lagomar (34°50'S, 55°58'W) , 28.XI.2004, 1 male (FCE Ar 2691); 21.I.2005, 1 female (FCE Ar 2692); 28.X.2004, 4 females (FCE Ar 2693); 17.IV.2005, 5 females (FCE Ar 7310); 21.X.2005, 1 female (FCE Ar 2694); 26.VI.2004, 1 male (FCE Ar 2696), 1 female (FCE Ar 2697); 29.VIII.2004, 3 males, 1 female (FCE Ar 2699); 21.I.2005, 1 male (FCE Ar 2700); XII.2004, 1 female (FCE Ar 2703); 1.VIII.2004, 4 females (FCE Ar 2718); 17.IV.2005, 1 male (FCE Ar 7307); IV.2009, 1 female (FCE Ar 6636); 6.VI.2009, 1 female (FCE 6637); V.2009, 1 male (FCE Ar 6638); 8.I.2010, 10 females, 16 immatures (FCE Ar 6640), all collected by M. Simó; Marindia (34°46'S, 55°49'W), I.1992, F. Costa leg., 1 male (FCE Ar 992), 1.X.2007, 1 male (FCE Ar 2345); 1 female (FCE Ar 2350); 8.XII.2007, 1 male (FCE Ar 2342); 20.XI.2007, 1 male (FCE Ar 2343); 20.XII.2007, 1 female (FCE Ar 2344); 20.XI.2007, 1 male (FCE Ar 2346); 1 male (FCE 2347); 28.XII.2007, 1 male (FCE Ar 2349); 1 female (FCE Ar 2351); 20.XI.2007, 1 male (FCE Ar 2353); 18.X.1996, 1 male (FCE Ar 2721); 1 male (FCE Ar 2723); 1 female (FCE Ar 2730); 26.V.1998, 1 male, 1 female (FCE Ar 2724), all collected by A. Aisenberg; Salinas (34°47'S, 55°49'W), 16.V.2005, M . Simó leg., 2 females (FCE Ar 2695); M. Simó leg., 13.II.2005, 1 male (FCE Ar 2701); San José de Carrasco (34°51'S, 55°58'W), 19.XI.2008, A. Aisenberg leg. , 1 male (FCE Ar 2341); Santa Ana (34°47'S, 55°27'W), 8.I.2005, M . Simó leg., 2 males (FCE Ar 2702), 2 females (FCE Ar 2707). Colonia . Colonia city (34°27'S, 57°51'W) 10.XI.1919. A.G. Fren leg., 1 male , 1 female, 5 immatures ( MACN Ar 36 344). Florida. Río Yi (33°26'S, 56°16'W), 9.X.2010, D . Gómez leg., 1 female, 1 immature (FCE Ar 4566). Lavalleja. Río Santa Lucía, Paso del Molino (34°17'S, 55°13'W), 22.II.2007 , 1 male, 4 females (FCE Ar 2726). Montevideo. Playa Mulata (34°53'S, 56°04'W), 17/II/2005, M . Simó leg., 1 female (FCE Ar 7308); 23.X.2009, C. Jorge leg., 1 male (FCE Ar 4565). Playa Carrasco (34°53'S, 56°03'W), 5.I.2005, M . Simó leg., 1 male (FCE Ar 7268); 5.I.2005, M. Simó leg., 1 female (FCE Ar 7270); 16.X.2009, 1 female (FCE Ar 6639); Playa Verde (34°53'S, 56°04'W) , 17.II.2005, M. Simó leg., 4 females (FCE Ar 7305); 30.XII.2009, M. Simó leg., 2 males, 3 females, 8 immatures (FCE Ar 6641); Río Negro. Las Cañas (33°09'S, 58°21'W), 20.XII:2002, V. Quirici leg. , 2 females (FCE Ar 2725); Club Biguá (33°14'S, 58°01'W), 7–8.III.2011, R . Postiglioni & V. Telesca leg., 1 male (FCE Ar 4906); 1female (FCE Ar 4907); 1 female (FCE Ar 6081); 1 male (FCE Ar 6082); Rocha. La Paloma (34°39'S, 54°10'W), 14.I.2008, L. Barufaldi leg. , 1 male (FCE Ar 2352); Laguna de Rocha (34°39'S, 54°15'W), 19.XII.2002 , 1 male (FCE Ar 2720); Santa Teresa (34°00'S, 53°31'W) , 21.XI.2009, 1 male (FCE Ar 2992); 22.XI.2012, 1 male, 1 female (FCE Ar 4587); 24.XI.2013, 1 male (FCE Ar 6554), all collected by A. Laborda. Barra de Chuy-La Coronilla (33°49'S, 53°27'W), 25.XI.2011, 2 immatures (FCE Ar 6608); 2 immatures (FCE Ar 6609); 1 immature (FCE Ar 6611) ; 1 female (FCE Ar 6613); 1 female (FCE Ar 6614); 1 immature (FCE Ar 6 615); 1 immature (FCE Ar 6616); 1 immature (FCE Ar 6617); 1 immature (FCE Ar 6618); 1 immature (FCE Ar 6620); 24.I.2012, 3 females (FCE Ar 6621); 1 immature (FCE Ar 6 622); 1 female (FCE Ar 6623); 2 immatures (FCE Ar 6626); 2 females; 1 immature (FCE Ar 6627); 1 immature (FCE Ar 6629); 1 female, 1 immature (FCE 6630), all collected by V. Mourglia. Tacuarembó. Bosque ribereño, sobre Río Tacuarembó (31°37'S, 55°39'W) , 28.IV.2011, M. Simó leg., (FCE Ar 6127); II.2012, M. Simó leg., 1 male (FCE Ar 6258).

Diagnosis. Males of A. senex resembles A. alticeps and A. marindia sp. nov. in the elongated cymbium of the male palp, but differ by the body size notoriously much larger (commonly 1.5–2 times), and by the sclerotized palea and pointed terminal apophysis of the male bulb ( Figs 3 D, 5C). Females can be recognized by the vulval chambers not projected laterally ( Figs 3 F, 6D).

Description. Male (FCE Ar 4906). The body is densely covered with setae. Carapace yellowish beige, with cephalic region dark brown. Thoracic groove narrow and shallow. Chelicerae, labium and endites dark brown. Clypeus dark brown, with four median setae. Sternum dark brown, with dark edges, acuminate posteriorly. Legs yellowish beige. Opisthosoma yellowish, with light brown spots. Venter yellow. Spinnerets yellow, densely covered with dark setae, anterior laterals the longest. Total length 16.60. Carapace 9.40 long, 7.20 wide. Clypeus 0.50 high. Eye diameters and interdistances AME 0.30, ALE 0.20, PME 0.55, PLE 0.50, AME–AME 0.30, AME– ALE 0.10, PME–PME 0.70, PME–PLE 0.80, PLE–PLE 1.50. Chelicerae 5.70 long, with three promarginal teeth, the median the largest, and three retromarginal teeth, increasing in length from basal to apical. Opisthosoma 8.0 long, 5.6 wide. Leg formula IV/I/III/II; segment length: I femur 7.80/patella 3.30/tibia 6.70/ metatarsus 6.1/ tarsus 3.2/total 27.10. II 7.40/3.20/6.20/6.0/3.20/26.0. III 6.90/3.40/5.80/6.50/3.60/26.2. IV 8.1/4.0/7.60/7.90/4.1/32.0. Leg spination: I, femur p 1ap, d1-1-1, r1-1; patella p1; tibia p1-1, v2-2 -2, r1-1; metatarsus p1-1-2, r1-1-2, v2-2 - 1. II, femur p1-1, d1-1-1, r1-1; patella p1; tibia p1-1, r 1-1, v1-1 -2; metatarsus p1-1-2, r1-1-2, v2-2 - 1. III, femur p1-1, d1-1-1, r1-1; patella p1, r1; tibia p1-1, d1, r1-1, v2-1 -2; metatarsus p1-1-2, r1-1-2, v2-2 - 1. IV, femur p1-1, d1-1-1, r1ap; patella p1, r1; tibia p1-1, d1, r1-1, v2-2 -2; metatarsus p1-1-2, r1-1-2, v3-2 -1. Palpus: cymbium yellowish, with two apical macrosetae. Palea conspicuous and membranous with pointed terminal apophysis. Median apophysis bifurcated ( Figs 3 A–D, 5C).

Female (FCE Ar 4907). As in the male except for the following: sternum yellowish beige, with dark edges, acuminate posteriorly. Legs yellowish. Total length 13.0. Carapace 6.20 long, 4.80 wide. Clypeus 0.30 high. Eye diameters and interdistances AME 0.30, ALE 0.20, PME 0.45, PLE 0.40, AME–AME 0.15, AME–ALE 0.05, PME–PME 0.5, PME–PLE 0.55, PLE–PLE 1.05. Chelicerae 3.50 long, with three promarginal teeth, the median the largest, and three retromarginal teeth, increasing in length from basal to apical. Opisthosoma 7.50 long, 4.6 wide. Leg formula IV/I/II/III; segment length: I femur 5.30/patella 2.60/tibia 4.50/ metatarsus 4. 0/ tarsus 2.0/total 18.40. II 5.10/2.50/4.20/3.90/2.30/18.0. III 4.60/2.30/3.80/4.40/2.40/17.5. IV 6.0/3.10/5.50/5.60/3.0/23.20. Leg spination as in the male. Palpus with conspicuous prolateral macrosetae in patella, tibia and tarsus. Tarsal claw pectinated and well developed. Epigynum rather flat, with a setaceous plate. Copulatory openings small, situated laterally on the posterior margin of the epigynal plate ( Figs 3 E, 6C). Head of spermathecae with porose area situated in an apical projection. Vulval chambers projected laterally ( Figs 3 F, 6D).

Variation: Measurements, males (n=5): carapace length 9.4–6.5, width 7.0–5.1; femur I length 7.7–5.5. Females (n=5): carapace length 7.2–6.3, width 5.4–4.6; femur I length 6.0–5.0.

Natural history. Many behavioural and ecological studies were developed on this species during the last two decades, when it was considered a junior synonym of A. brasiliensis . Allocosa senex ( Fig. 8 B) is a very common wolf spider in fluvial, estuarine and oceanic coastlines from Southern South America, and have been chosen as a model organism for many ecological and behavioural studies. The species constructs burrows in the sand ( Capocasale 1990; Aisenberg et al. 2011b; Aisenberg & Peretti 2011a, 2011b; Albín et al. 2015; De Simone et al. 2015) and shows sex roles reversal, with males bigger than females. Females are the mobile sex, which initiates courtship ( Aisenberg et al. 2007; Aisenberg 2014), males produce volatile pheromones ( Aisenberg et al 2010a) and females can be cannibalized by the male ( Aisenberg et al. 2009, 2011a). Copulation takes place inside male burrows ( Aisenberg et al. 2007; Aisenberg & Costa 2008; García Díaz et al. 2015) during warmer seasons ( Costa et al. 2006; Costa 2014; Costa & Simó 2014; Carballo et al. 2015). Furthermore, it presents sexual dimorphism represented by morphological, behavioural and physiological features strongly associated to inhabiting the dune lines of natural sandy coasts ( Aisenberg et al. 2010b; Aisenberg & Peretti 2011a). Another study demonstrated the parasitoidism caused by the pompilid wasp Anoplius bicinctus (Dahlbom) , mainly on juveniles and females ( Stanley et al. 2013). The species has been catalogued as a good bio-indicator for coastal environments, since it is highly associated with psammophile vegetation ( Costa 1995; Aisenberg et al. 2011b, 2011c; Ghione et al. 2013; Jorge et al. 2015).

Distribution. Northeastern Argentina, southern Brazil and Uruguay ( Fig. 9 ).

Figures

FIGURE 3. Allocosa senex (Mello-Leitão). A – C, male left palp. A, ventral view. B, prolateral view. C, retrolateral view. D, close up of the bulb, ventral view. E – F, epigynum. E, ventral view. F, dorsal view. Scales. A, B, C: 1 mm; D, E: 0.5 mm. Abbreviations: CO: copulatory opening; FD: fertilization duct; HS: head of the spermatheca; MA: median apophysis; PR: palea region; TA: terminal apophysis; VC: vulval chamber.

FIGURE 5. Structures of male palpus. Allocosa marindia sp. nov.: A, bulb, ventral view; B, apical macrosetae. Allocosa senex (Mello-Leitão): C, bulb, ventral view; D, apical macrosetae. Allocosa alticeps (Mello-Leitão): E, bulb, ventral view; F, apical macrosetae. Abbreviations: MA: median apophysis; PR: palea region; ST: subtegulum; TA: terminal apophysis.

FIGURE 8. Living specimens. A, Allocosa marindia sp. nov.; B, Allocosa senex (Mello-Leitão) (A and B from Playa Pascual, San José, Uruguay); C – D, Allocosa alticeps (Mello-Leitão) from Pehuen-có, Buenos Aires, Argentina; C, female carrying spiderlings. D, specimen at the burrow entrance.

FIGURE 9. Distribution records of Allocosa marindia sp. nov., Allocosa brasiliensis (Petrunkevitch), Allocosa senex (Mello- Leitão) and Allocosa alticeps (Mello-Leitão).