Polistes (Fuscopolistes) parametricus Buck

Buck, Matthias, Cobb, Tyler P., Stahlhut, Julie K. & Hanner, Robert H., 2012, Unravelling cryptic species diversity in eastern Nearctic paper wasps, Polistes (Fuscopolistes), using male genitalia, morphometrics and DNA barcoding, with descriptions of two new species (Hymenoptera: Vespidae), Zootaxa 3502, pp. 1-48 : 14-19

Polistes (Fuscopolistes) parametricus Buck , sp. nov.

( Figs 2–4 , 21 , 29 , 34 , 38 , 46 , 54 , 62 , 66 )

(IAV Figs B10.35, B10.40, B10.43, C81.1–6)

Polistes sp. B: Buck et al. (2008): http://www.biology.ualberta.ca/bsc/ejournal/bmc_05/81p_spb.html

Type material. Holotype 3 (debu00289459, IAV Fig. C81.2): U.S.A., MARYLAND, Prince George's Co., Largo, Hwy 214 & Church Rd., 38°54'02"N, 76°45'32"W, 40 m, 1 Sep 2007, M. Buck ( DEBU). Paratypes: 57 Ƥ, 24 33. U.S.A. DISTRICT OF COLUMBIA: Washington, 1 Ƥ, [date illegible] 1945, 1 3 (with 7 Sphexicozela mites), " 8.10.1943 ", M. Vogel ( USNM); 1 Ƥ, Washington, Rock Creek Park, Park Road meadow, 20 Sep 1988, R.A. Boettcher ( USNM). ILLINOIS: Hancock Co., 1 Ƥ, A.L. Kibbe Field Sta., 7 Jul 1978, stud[ent] coll. ( AMNH). MARYLAND: 1 3, Linwood, 9 Sep [19]08, "87", P.R. Myers ( USNM); Anne Arundel Co., 1 3 (debu00289286, GB# EU649665 ), Milt's Pit, 4 mi SW Davidsonville, 38°52'40"N, 76°40'10"W, 20 m, 2 Sep 2007, M. Buck ( DEBU); 1 Ƥ (debu00289498, GB# EU649658 ), Patuxent Wildlife Res. Ref., North Tract, 39°02'36"N, 76°45'37"W, 30 m, 1 Sep 2007, M. Buck ( DEBU); Baltimore Co., 1 3, 10 Sep 1967, C.F. Reed ( USNM); Montgomery Co., 1 Ƥ, Gaithersburg, 3 Nov 1967, W.F. Gimpel ( USNM); Plummers I., 1 Ƥ, 25 Apr 1961, K.V. Krombein, 1 Ƥ, 11 Apr 1971, P.D. Hurd, 1 Ƥ, Apr [19]08 ( USNM); Glen Echo, 1 Ƥ, 23 Apr 1980, 1 Ƥ, 2 Jul 1981, 1 Ƥ, 15 Apr, 1 Ƥ (partially eaten by dermestids), 20 Apr, 2 Ƥ, 24 Jul, 1 Ƥ, 13 Aug 1982, malaise trap, E.M. Barrows ( USNM); 1 Ƥ, 4 mi SW Ashton, 6 Jul 1986, malaise trap, G.F. & J.F. Hevel ( USNM); 1 3, Chevy Chase, Woodend, 26 Sep 1974, R.A. Boettcher ( USNM); 3 Ƥ (USGS-DRO-225928–...30), [Wheaton], 39.0406°N, 77.0155°W, 2 Aug 2011, [nesting in siding of house], S.W. Droege ( PMAE); Prince George's Co., 2 Ƥ (debu00288743–...4, GB# EU649656 ), Greenbelt, Springfield Rd., Beltsville Agr. Res. Center, 39°01'50"N, 76°50'20"W, 40 m, 24 Aug 2007, M. Buck ( DEBU); 1 3 (USGS-DRO-173684), [Greenbelt, Beltsville Agr. Res. Center], 39.0164°N, 76.8511°W, 21 Sep 2010, S.W. Droege ( PMAE); 3 33 (debu00288790–...2, GB# EU649662 ; 1 spm. with 2 Sphexicozela mites), Patuxent Wildlife Res. Ref. 1 mi E Montpelier, 39°02'57"N, 76°49'04"W, 50 m, 24 Aug 2007, power line, M. Buck ( DEBU); 1 Ƥ (debu00288834, GB# EU649657 ), 3 33 (debu00288847–...9, GB# EU649663 ) Patuxent Wildlife Res. Ref., Visitor Center, 39°01'38"N, 76°47'50"W, 40 m, 24 Aug 2007, M. Buck ( DEBU); 1 Ƥ, Patuxent Wildlife Refuge nr. entrance, 24 Apr 1966, D.R. Smith ( USNM); 1 Ƥ, " PWRC " [= Patuxent Wildlife Research Centre], 25 Jul 1977, W.N. Beyer ( USNM); 1 3 (with eclosed male Strepsiptera pupa), College Park, 18 Sep 1979, S. Gross ( USNM); College Park, Paint Branch [Stream Valley] Park, 1 Ƥ, 19 Apr, 1 Ƥ, 11 May 1986, E. Eaton ( PMAE); 1 3 (debu00289022, GB# EU649664 ), same as holotype except 25 Aug 2007, M. Buck ( DEBU); 1 3, Beltsville, 12 Sep 1915, J. Silver ( USNM); 1 3 (USGS-DRO-173786), [Bowie, Governor's Bridge], 38.9431°N, 76.69661°W, 19 Sep 2010, S.W. Droege ( PMAE); Washington Co., 1 3 (USGS-DRO- 140260), [Dargan], 39.3785°N, 77.727°W, 4 Sep 2009, S.W. Droege ( PMAE). MISSOURI: Shannon Co., 1 3 (GB# JN276781 ; with 6 Sphexicozela mites), ONSR [= Ozark National Scenic Riverways], Devil's Well [Hollow], 28 Sep 2005, old field, at Solidago, M. Arduser ( PMAE); St. Francois Co., 1 3, St. Joseph [= St. Joe] St. Pk., 27 Sep 2001, bike trail, M. Arduser ( PMAE); St. Louis Co., 1 3, Pelican I. Nat. Area, 27 Sep 1995, Eupatorium altissimum flowers, M. Arduser ( PMAE). NORTH CAROLINA : Macon Co., 1 Ƥ, Highlands, 3,800 ft, 4 May 1957, J.R. Vockeroth ( CNCI). OHIO: 1 Ƥ (USGS-DRO-213162), [rest stop along I-70, probably nr. Zanesville], 28 May 2010, S.W. Droege ( PMAE); Franklin Co., 1 Ƥ, Columbus [no date or collector] ( USNM); 2 Ƥ (one missing both antennae), as previous but "Mar", 1 3, "Sept", Bridwell ( USNM); Vinton Co., [Vinton Furnace Experimental Forest], 39.1921°N, 82.4048°W, 1 Ƥ (USGS-DRO-166527), 25 Jun–8 Jul 2010, 1 Ƥ (USGS-DRO-184801), 20 Aug–3 Sep 2010, 1 3 (USGS-DRO-183582), 1-14 Oct 2010, [cup traps], D. Hosack ( PMAE). PENNSYLVANIA: 1 Ƥ, "1572" [locality not given], collection C.F. Baker ( USNM); Cumberland Co., 1 Ƥ, N[ew] Cumberland, 11 Sep [19]09, "242", P.R. Myers ( USNM); Dauphin Co., 2 Ƥ, 1 3, Linglestown, 19 Sep [19]09, "244a", Kirk & Champlain, collection P.R. Myers ( USNM); Perry Co., Duncannon, 40°22'35"N, 77°6'50"W, 1 Ƥ, 9 Mar 2010, 1 Ƥ, 10 Apr 2010, 1 Ƥ, 21 Jun 2012, 1 Ƥ, 9 Aug 2012, S.M. Nacko ( PMAE). VIRGINIA: Arlington Co., 1 Ƥ, East Falls Church, 14 Jul [19]14, V. Roberts ( USNM); Fairfax Co., 1 3 (apical flagellomeres missing on both antennae), Dunn Loring, 30 Oct 1987, N.E. Adams ( USNM); Giles Co., 1 Ƥ (debu01009147, GB# EU649653 ), 10 km NW Blacksburg, 18 May 1997, sphagnum bog/fen, malaise, W. Scanlon ( DEBU); 1 Ƥ, "Montgomery Co." [probably in error], 8 km NW Blacksburg, 1000 m, 1–17 Aug 1987, malaise, BRC Hym. Team ( PMAE); 1 Ƥ (debu01009115, GB# EU649654 ), Mountain Lake Biol. Stn., 37°22’31”N, 80°31’18”W, 13–26 May 2001, S. Goodfellow ( DEBU); 1 Ƥ (debu01009148, GB# EU649659 ), "Jefferson Co." [= Jefferson Natl. For.], Cascades Recr. Area USFS, 17–28 May 1999, by bathroom, C. Rothfels ( DEBU); 1 Ƥ (with malformed tergum 4), Cascades Recr. Area, 37°21’0”N, 80°36’30”W, 11–25 May 2008, netted nr. underside of building roof, N. Moore ( DEBU); Montgomery Co., 1 Ƥ (debu00250120, GB# EU649655 ), Pandapas Pond, 37°16’30”N, 80°28’0”W, 19 May 2005, S.M. Paiero ( DEBU); 1 Ƥ (debu01009149, GB# EU649660 ), Jefferson N.F., Pandapas Pond Recr. Area, 17–28 May 1999, L. Soth ( DEBU). WEST VIRGINIA: Doddridge Co., 1 Ƥ, [Toll gate on old US-50], 39.2775°N, 80.908°W, 24 Sep 2009, J. Whitaker ( PMAE); Hampshire Co., 1 Ƥ, [Timber Ridge Camp], 39.2299°N, 78.4629°W, 20 Aug 2010, S.W. Droege ( PMAE); Hardy Co., 1 Ƥ (debu01009150, GB# EU649661 ), [3 mi NE Mathias], 38.9167°N, 78.8167°W, spring 2006, D. Smith ( DEBU); 4 Ƥ (somewhat mouldy but otherwise in good condition), as previous except 6–25 Apr 2006 ( PMAE); 1 Ƥ (somewhat mouldy but otherwise in good condition), as previous except 9–22 Jun 2006 ( PMAE). Other material examined. KENTUCKY: Wolfe Co., 1 Ƥ, Red River Gorge, Hwy 715 bridge at Red River, 13 Jul 1980, L. Guidry ( USNM). GW/EW only 0.80, but otherwise like typical P. parametricus .

Diagnosis. This species is intermediate between P. f u s c a t u s and P. metricus both of which occur in sympatry with P. p a r a m e t r i c u s. Females of P. parametricus look like dark varieties of P. m e t r i c u s, or like dark P. f u s c a t u s with reduced yellow markings although they never show black markings on the clypeal disc which are usually present in similarly patterned P. f u s c a t u s. The female of P. parametricus differs from both P. metricus and P. f u s c a t u s by the stouter scape and wider gena (see key below, Table 2 ). Furthermore, it has a less bulging sternum 2 than P. metricus , and more coarsely punctate sterna 3–5 as well as a longer malar space than P. f u s c a t u s (see key below, Table 2 ). Males can be distinguished from P. f u s c a t u s by the narrower flagellomere 10 ( Table 3 ), by the different distribution and smaller size of aedeagal teeth, and darker colouration of anterior surface of basal flagellomeres (see key below). Males with pale antennae (an uncommon colour variety) are very similar to P. metricus but can usually be separated from the latter by the stouter scape and shorter flagellomere 1 (see key, Table 3 ). Several other characters (smaller size, flat instead of concave clypeal disc, darker colouration of anterior surface of flagellum, presence of pale apical fascia on tergum 1) are also helpful in distinguishing P. parametricus males from P. metricus but do also occur in a small percentage of the latter.

Description. Structural characters.

FEMALE ( Fig. 2 ; IAV Figs C81.1, C81.3). Fore wing length 14.0– 18.5 mm (12.0 mm in one dwarfed female). For morphometric measurements see Table 2 . The most diagnostic parameters are: a wide gena (GW/EW), long malar space (UID/MS), and stout scape (UID/SL, SL/SW).

Head (IAV Fig. C81.5). Clypeus with apex minutely truncate, length of truncation ca. 0.75–1.0 × width of median ocellus. Clypeal disk convex, micropunctate except V-shaped apical area which is finely shagreened; diameter of micropunctures 12–16 μm (similar to P. f u s c a t u s and P. metricus ), punctures near centre of micropunctate area separated mostly by less than puncture diameter; macropunctures scattered and of variable sizes, smaller in basal half of clypeus, becoming increasingly larger towards apex, where they are large and conspicuous; macropunctures bearing stiff, more or less erect bristles (size and thickness correlated to puncture size), micropunctures bearing small appressed setulae. Interantennal tubercle moderately developed, similar to P. fuscatus . Frons densely micropunctate, less shining than clypeus, area around and in front of ocelli with scattered macropunctures that bear erect hairs whose tips are bent or straight. Upper 2/3 of gena micropunctate, lower third including malar space devoid of micropunctures (as in most Fuscopolistes, unlike P. rubiginosus ); macropunctures distinctly larger on lower third of gena than on upper 2/3. Occipital carina well developed on upper 2/3 of head capsule, absent on lower third, ending quite abruptly ventrally. Head behind lateral section of occipital carina with sparse fringe of longer hairs (length slightly longer than MOD), their tips bent anteriorly; fringe comparable to other Fuscopolistes, not as long and as dense as in the P. exclamans -group of Aphanilopterus . Eye bare. Mandible of the usual shape (see under previous species); outer surface bare except for scattered macropunctures of moderate size, each bearing small bristle. Basal flagellomeres (1–2 or 4) of antenna with very short, bristly hairs posteriorly and dorsally (length ca. 30 μm, rarely up to 50 μm; becoming sparser and shorter distally).

Mesosoma. Pronotum with fovea normally developed, usually deep (rarely small and shallow). Pronotal carina similar to P. fuscatus , laterally ending about half-way between pronotal fovea and ventral margin of pronotum, carinate to a point about half antennal socket diameter above pronotal fovea (below that rounded into a smooth welt); carina more or less evenly curved dorsally between humeral angles (median portion not elevated as in P. hirsuticornis ). Punctation of mesosoma similar to P. fuscatus and P. metricus : diameter of mesopleural micropunctures ca. 14–16 μm, separated by about half their width; macropunctures 2–3 × the size of micropunctures, scattered and fairly conspicuous to quite inconspicuous (best developed ventrally), each giving rise to a longer hair with bent apex. Dorsal groove of mesopleuron as in P. f u s c a t u s, with median third clearly impressed. Metapleuron with more shallow micropunctures and therefore more shining than mesopleuron (as in P. fuscatus and P. metricus ). Scutum with scattered and inconspicuous longer hairs between regular pubescence, apices of these hairs bent at apex; macropunctures very inconspicuous. Scutellum with the usual weak median longitudinal keel in anterior half (sometimes very faint), its macrosetae longer and more conspicuous than on scutum, macropunctures distinct but not very conspicuous. Metanotum with macrosetae longer than on scutellum but macropunctures hardly evident. Propodeum weakly microstriate laterally (as in P. fuscatus and P. metricus ), median half becoming increasing ridged posteriorly, with microstriae and a few ridges, the latter weak to fairly strong (but less so than in coarsely ridged species like P. rubiginosus ); pubescence of propodeum becoming longer posteriorly (ventrally), macrosetae more or less restricted to area just beside median groove, their length also increasing posteriorly (ventrally). Propodeal orifice variable, wide and usually gradually narrowing dorsally with upper width (below apical emargination) about half lower width (i.e., as in P. f u s c a t u s and P. m e t r i c u s), but sometimes dorsal third narrowed fairly abruptly to about one third of ventral width (as in P. hirsuticornis ).

Legs and wings without unusual features, morphology essentially as in P. fuscatus .

Metasoma (IAV Figs C81.1) with tergum 1 1.08–1.20 × as wide as long (length measured from attachment of petiole muscle to apex of tergum); anterior half of tergum typically moderately and evenly curved in lateral view with degree of convexity gradually decreasing posteriorly (in P. f u s c a t u s and P. m e t r i c u s typically rising steeply for a short distance behind attachment of propodeal muscle, then fairly abruptly levelling off onto weakly curved posterior surface; IAV Fig. C77.4). Sternum 1 transversely microstriate, postpetiolar part usually with several stronger ridges, near petiole always with one or more stronger, more or less clearly defined ridges. Terga finely micropunctate, punctures and associated setulae becoming larger posteriorly; macropunctures hardly visible and very sparse but becoming more numerous on posterior terga and towards base of tergum 1, associated macrosetae fairly long near base of tergum 1, very short and hardly noticeable on tergum 2, increasing in length towards apex of metasoma. Sternum 2 weakly to moderately convex in lateral view (similar to P. fuscatus , unlike P. metricus ), finely and fairly densely micropunctate (similar to tergum), macropunctures more numerous than on tergum, with macrosetae longer and fairly conspicuous. Punctation of sterna 3–5 (IAV Fig. B10.40) coarser and slightly to distinctly more spaced than in P. fuscatus (except posterior and lateral margins), punctures often more or less irregularly distributed, leaving small bare areas of two or more puncture diameters; in areas where punctures are sparse macropunctures are more numerous than micropunctures. Macrosetae increasing in length towards apex of metasoma. Sternum 6 more finely and densely punctate than the previous sternum, with the usual median tuft of hairs at extreme base (usually concealed by previous sternum). Stinger with apical half straight; each stinger lancet with ca. five fine barbs near apex.

MALE ( Fig. 3 ; IAV Fig. C81.2, C81.4). Characters as in female except as follows. Fore wing length 12.0–16.0 mm. For morphometric measurements see Table 3 .

Head ( Fig. 4 ; IAV Fig. C81.6). Clypeus usually at least narrowly separated from eye (at most by ca. 0.4 MOD), in about one third of specimens in contact with eye for a distance of up to 0.8 MOD. Clypeal apex narrowly rounded, less produced than in female. Clypeal disk usually flat in lower 2/3 (excluding area near lateral margin), rarely slightly convex (in P. f u s c a t u s very slightly concave, rarely flat, in P. m e t r i c u s often slightly concave); surface entirely and densely micropunctate with weakly shining interspaces, macropunctures scattered, bearing straight, pale hair-like macrosetae (except in dorsolateral corners where some setae have bent tips), which are longer in dorsal third; diameter of micropunctures smaller than in female and punctures closer together, clypeus therefore less shining; clypeal margin with a moderately dense fringe of silvery hairs (more dense than in P. f u s c a t u s). Interantennal tubercle moderately developed, similar to related species; subantennal longitudinal grooves usually less impressed than in P. f u s c a t u s, with outer margin usually more rounded (usually more carinate in P. f u s c a t u s). Gena and malar space micropunctate throughout; macropunctures of more or less uniform size. Mandible showing the usual sexual dimorphism; basal 3/5 of outer surface densely micropunctate and pubescent, with pubescence becoming denser and longer near upper articulation (but not forming distinct brush, its development similar to P. f u s c a t u s and P. metricus ), macropunctures smaller than in female and only bearing weak hairs; inner surface of mandible lacking ridges. Labrum narrow and parallel-sided (width ca. 0.5 MOD), with several small bristles near the narrowly rounded apex. Scape usually stouter than in P. metricus ; flagellum (IAV Fig. B10.35) longer than in female, apically recurved as in other Fuscopolistes; flagellomere 10 narrower than in P. f u s c a t u s ( Table 3 : FXW/FIW). Flagellomere 1 with outstanding hairs very short and inconspicuous (length ca. 25 μm); following flagellomeres with hairs even shorter to absent. Tyloids similar to related species, almost linear on flagellomere 1, becoming increasingly wider towards flagellomere 7 where tyloid covers entire ventral surface (as in flagellomeres 8–11); pubescence of tyloids becoming sparser distally, apical four flagellomeres with large, shining, bare patches.

Mesosoma. Pronotal fovea similar to female, shallow to virtually absent in two specimens.

Metasoma with tergum 1 1.01–1.17 × as wide as long. Sternum 2 less convex than in female (similar to P. fuscatus , usually less convex than in P. metricus ), posterior half of sternum 3 and sterna 4–6 flattened ventrally, sternum 7 slightly concave behind anteromedian tubercle, the latter small and not visible in profile (as in P. fuscatus and P. m e t r i c u s). Punctation of flattened area of sterna 3–6 very fine, macropunctures small as well, interspaces large and shining, pubescence extremely short; posterolateral corners of sterna with patches of fairly long setulae, increasing in length from sternum 3 to 6 (length on latter roughly 1 MOD; similar to P. fu s c at us and P. metricus ). Sternum 7 with scattered bristly hairs in posterior depression, forming a sparse fringe near apical margin (length of bristles increasing laterally), pubescence of disc otherwise slightly longer than on previous sternum, laterally with longer pubescence which is much shorter than posterolateral patch of hairs of previous sternum (similar to P. fuscatus and P. m e t r i c u s).

Genitalia ( Figs 21 , 29 , 34 , 38 , 46 , 54 , 62 , 66 ; Table 4 ) very similar to P. hirsuticornis , different as follows: preapical ventral curvature of digitus variable, weak to pronounced, in one specimen angulate (right digitus only, aberrant?); anteroventral lobe of digitus more angulate and pointed (as in most Fuscopolistes). Aedeagal teeth slightly longer.

Colouration very similar to dark varieties of P. metricus , far less variable than in P. hirsuticornis . See detailed description in Buck et al. (2008: "sp. B").

Etymology. The name parametricus has a twofold meaning. On one hand, it alludes to the great overall similarity with P. m e t r i c u s, and on the other hand it points to the importance of morphometric parameters in recognizing this species.

Distribution ( Fig. 1 ). The distribution of P. parametricus is poorly known, especially in the western and southern part of its range. Compared to P. hirsuticornis , the range appears to be more restricted, especially in the south. Our southernmost record is from North Carolina , close to the South Carolina and Georgia state lines. The species is apparently absent from Florida, from where we have studied plenty of material. The range along the northern Atlantic coast also appears to be more restricted than in P. hirsuticornis , with a notable absence of records from New Jersey and New York (areas well documented in collections examined by us). By contrast, the species is well represented in certain areas along the Appalachians (Giles Co., Virginia; Perry Co., Pennsylvania) where P. hirsuticornis is seemingly absent. In western Virginia and North Carolina the species reaches altitudes of more than 1,100 m. This is in sharp contrast to P. hirsuticornis , which is only known from low-lying areas below 300 m.

Phenology ( Fig. 6 ). Earliest and latest collection records for females are March 9 and November 3, respectively. Our data show a distinct spring maximum in April and a less pronounced summer maximum in August. Similar to P. hirsuticornis there is a sharp decline in early summer, with only two females collected during the month of June (discussion see below under "Suspected social parasitism"). Earliest and latest records for males are August 24 and October 30, respectively.

Parasites and symbionts. One male from College Park, Maryland (see under material examined) carries an eclosed male Strepsiptera pupa between terga 4 and 5. It probably belongs to a species of the genus Xenos (Stylopidae) , which parasitize several other sympatric species of Polistes . Two other males from Maryland and Washington, D.C. (see under material examined) carried deutonymphs of Sphexicozela (Acari: Winterschmidtiidae ) (det. D.E. Walter, mounted on slides). The mites were either found on the male genitalia which were pulled out of the genital chamber for examination (male from Maryland, 2 mites), or below the apical margins of sterna 5, 6 (6 mites) and on the exposed genitalia (1 mite, male from D.C.). For additional information on Sphexicozela see also the "Parasites and symbionts" paragraph under P. hirsuticornis .

Figures

FIGURE 1. Locality records for Polistes hirsuticornis sp. nov. (triangles) and P. parametricus sp. nov. (circles).

FIGURE 2. Female of Polistes parametricus sp. nov. on goldenrod; Smithburg, Doddridge Co., West Virginia, October 4, 2006 (Photo: Donna Race).

FIGURES 3 – 4. Male of Polistes parametricus sp. nov.; Duncannon, Pennsylvania, September 12, 2011 (Photos: Scott Nacko).

FIGURES 18 – 21. Paramere and volsella, inner view: 18. Polistes metricus (MD, Anne Arundel Co.), 19. P. rubiginosus (MO, Callaway Co.), 20. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 21. P. parametricus sp. nov. (MD, Prince George's Co.).

FIGURES 22 – 29. Digitus, outer surface: 22. Polistes bellicosus (MD, Anne Arundel Co.), 23. P. carolina (TX, Bastrop Co.), 24. P. dorsalis (MD, Anne Arundel Co.), 25. P. f u s c a t u s (ON, Wellington Co.), 26. P. metricus (MD, Anne Arundel Co.), 27. P. rubiginosus (TX, Bastrop Co.), 28. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 29. P. parametricus sp. nov. (MD, Prince George's Co.). ap: apical process; avl: anteroventral lobe; ba: basal articulation; sb: scale-like bristles.

FIGURES 30 – 34. Anteroventral lobe of digitus, outer surface (30, 33, 34), and inner surface (31, 32): 30, 31. Polistes hirsuticornis sp. nov. (MD, Anne Arundel Co.), 32. P. bellicosus (MD, Worcester Co.), 33. ditto (MD, Anne Arundel Co.), 34. P. parametricus sp. nov. (MD, Prince George's Co.).

FIGURES 35 – 38. Digitus, dorsal view (35), and ventral view (36 – 38): 35, 36. Polistes dorsalis (MD, Anne Arundel Co.), 37. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 38. P. parametricus sp. nov. (MD, Prince George's Co.). ap: apical process; avl: anteroventral lobe; ba: basal articulation; gr: groove; is: inner surface; os: outer surface.

FIGURES 39 – 46. Aedeagus, left lateral view: 39. Polistes bellicosus (MD, Worcester Co.), 40. P. carolina (TX, Bastrop Co.), 41. P. dorsalis (MD, Anne Arundel Co.), 42. P. f u s c a t u s (MD, Anne Arundel Co.), 43. P. metricus (MD, Anne Arundel Co.), 44. P. rubiginosus (MO, Callaway Co.), 45. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 46. P. parametricus sp. nov. (MD, Prince George's Co.). ap pt: apical portion; apd: apodeme (paired).

FIGURES 47 – 54. Apex of aedeagus, left lateral view: 47. Polistes bellicosus (MD, Worcester Co.), 48. P. carolina (TX, Bastrop Co.), 49. P. dorsalis (MD, Anne Arundel Co.), 50. P. f u s c a t u s (MD, Anne Arundel Co.), 51. P. metricus (MD, Anne Arundel Co.), 52. P. rubiginosus (MO, Callaway Co.), 53. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 54. P. parametricus sp. nov. (MD, Prince George's Co.).

FIGURES 55 – 62. Apex of aedeagus, ventral view: 55. Polistes bellicosus (MD, Anne Arundel Co.), 56. P. carolina (TX, Bastrop Co.), 57. P. dorsalis (MD, Anne Arundel Co.), 58. P. f u s c a t u s (MD, Anne Arundel Co.), 59. P. metricus (FL, Highlands Co.), 60. P. rubiginosus (MO, Callaway Co.), 61. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 62. P. parametricus sp. nov. (MD, Prince George's Co.). med ex: median expansion; pe val: penis valve. Arrows indicate the end of the row of aedeagal teeth.

FIGURES 63 – 66. Median expansion of aedeagus, ventrolateral view: 63. Polistes fuscatus (MD, Anne Arundel Co.), 64. P. metricus (MD, Anne Arundel Co.), 65. P. hirsuticornis sp. nov. (MD, Anne Arundel Co.), 66. P. parametricus sp. nov. (MD, Prince George's Co.).

FIGURES 5 – 7. Phenology of Polistes species: 5. P. hirsuticornis sp. nov., 6. P. parametricus sp. nov., 7. P. fuscatus. Data for P. hirsuticornis and P. parametricus based on all available material (see material examined under both species). Data for P. fuscatus based on random sample of specimens collected by E. Eaton in Cincinnati, Ohio, over several years.