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) Author Buck, Matthias Author Cobb, Tyler P. Author Stahlhut, Julie K. Author Hanner, Robert H. text Zootaxa 2012 2012-10-01 3502 1 48 journal article 10.11646/zootaxa.3502.1.1 6ba8f5c9-f02f-4529-9b47-6fb31fc4a585 1175-5326 210194 6126D769-A131-49DD-B07F-0386E62FF5B9 Polistes ( Fuscopolistes ) hirsuticornis Buck , sp. nov. ( Figs 20 , 28 , 30, 31 , 37 , 45 , 53 , 61 , 65 ) (IAV Figs B 10.14 , B 10.16 , B 10.18 , C 80.1 – 7 ) Polistes sp. A: Buck et al . ( 2008 ) : http://www.biology.ualberta.ca/bsc/ejournal/bmc_05/ 80 p_spa.html Type material . Holotype 3 (debu00289048, GB# EU 649650 ): U.S.A. , MARYLAND, Anne Arundel Co., Milt's Pit, 4 mi SW Davidsonville, 38 ° 52 ' 40 "N , 76 ° 40 ' 10 "W , 20 m , 25 Aug 2007 , M. Buck ( DEBU ). Paratypes : 31 Ƥ, 23 33 . U.S.A. DELAWARE: New Castle Co., 1 Ƥ (debu00288935), Chesapeake & Delaware Canal at Delaware Bay, 39 ° 33 ' 29 "N , 75 ° 34 '02"W , 4 m , 26 Aug 2007 , sandy area, M. Buck ( DEBU ). FLORIDA: Alachua Co., 1 Ƥ, Gainesville, DPI [= Division of Plant Industry], 26 Apr 1994 , woods–mixed forest, L. Stange ( FSCA ); 1 Ƥ, Gainesville, Agricultural Experiment Station, " 5195 ", no date (probably collected between 1911 and 1946 ), J.R.W. [= J.R. Watson] ( FSCA ); 1 Ƥ (with eclosed male Strepsiptera pupa), same as previous except "Ed 1200 " ( FSCA ); 1 Ƥ (with> 20 deutonymphs of Sphexicozela ), Gainesville, “ 55 d 19 ”, “K.W. Cooper Collection 1960–1965 ” ( USNM ). ILLINOIS: Champaign Co., 1 Ƥ, Champaign, 25 Sep 1956 , J.F. McAlpine ( CNCI ). MARYLAND: 1 Ƥ (debu00289053, GB# EU 649646 ), 7 33 (debu00289045–... 47 , ... 49 –... 52 , GB# EU 649651 ), same as holotype ; 2 Ƥ, 4 33 (debu00289289–... 92 , GB# EU 649652 ), same as holotype except 2 Sep 2007 ( DEBU , Ƥ: PMAE ); Anne Arundel Co., 1 3 (debu00289496), Patuxent Wildlife Res. Ref., North Tract, 39 °02' 36 "N , 76 ° 45 ' 37 "W , 30 m , 1 Sep 2007 , M. Buck ( DEBU ); 1 Ƥ (USGS-DRO-084749), [Jug Bay Wetlands Sanctuary], 38.7847 °N , 76.7003 °W , 15 Sep 2007 , S.W. Droege ( PMAE ); Prince George's Co., 5 33 (debu00288728–... 32 , GB# EU 649648 ), Greenbelt, Springfield Rd., Beltsville Agr. Res. Center, 39 °01' 50 "N , 76 ° 50 ' 20 "W , 40 m , 24 Aug 2007 , M. Buck ( DEBU ); 1 Ƥ (USGS-DRO- 052349 ), [same locality as previous], 39.02935 °N , 76.8401 °W , 18–19 Apr 2005 , S.W. Droege, ( DEBU ); 1 Ƥ (debu00288818), 2 33 (debu00288819–... 20 , GB# EU 649649 ), Patuxent Wildlife Res. Ref., Visitor Center, 39 °01' 38 "N , 76 ° 47 ' 50 "W , 40 m , 24 Aug 2007 , M. Buck ( DEBU ); 1 3 (debu00288881), Patuxent Wildlife Res. Ref., Duvall Pond, 39 °02' 43 "N , 76 ° 47 ' 28 "W , 27 m , 24 Aug 2007 , M. Buck ( DEBU ); Largo, Hwy 214 & Church Rd., 38 ° 54 '02"N , 76 ° 45 ' 32 "W , 40 m , 1 Ƥ (debu00289018), 25 Aug 2007 , 1 Ƥ (debu00289449), 1 3 (debu00289448), 1 Sep 2007 , M. Buck ( DEBU ); 1 Ƥ (USGS-DRO- 173782 ), [Bowie, Governor's Bridge], 38.9431 °N , 76.69661 °W , 19 Sep 2010 , S.W. Droege ( PMAE ); 1 Ƥ (debu01009146, GB# EU 649645 ), [Bowie], 38.9623 °N , 76.7501 °W , 17 Sep 2006 , K. Hutson, ( DEBU ); Dorchester Co., 1 3 (USGS-DRO- 137577 ), 38.3491 °N , 75.9366 °W , 5 Oct 2009 , S.W. Droege ( PMAE ). MISSOURI: Boone Co., Columbia , 1 Ƥ, 20 Jun 1967 , 7 am– 4 pm, 1 Ƥ, 17–30 Mar 1968 , malaise trap, F.D. Parker ( USNM ); 2 Ƥ, Columbia , “MO 49 E 14 ”, “K.W. Cooper Collection 1960–1965 ” ( USNM ); Clark Co., 1 Ƥ, Rose Pond Conservation Area, T 64 N R 6 W S. 14 (sel/ 4 ), 6 Sep 1998 , 1500 hrs, Helianthus tuberosa flowers, M. Arduser ( PMAE ). NEW YORK : Rockland Co., 1 3 , Nyack, 1885 [ sic !, circular label, J.L. Zabriskie collection] ( AMNH ); Suffolk Co., 1 Ƥ, Wading River, 24 Jun 1989 , R. Foster ( AMNH ). NORTH CAROLINA : Pasquotank Co., 1 Ƥ, Elizabeth City, 5 Apr 1929 , no collector ( USNM ; specimen coated by thick, greasy deposit); Dare Co., 1 Ƥ, Kill Devil Hills, 24 May 1952 , K.V. Krombein ( USNM ). SOUTH CAROLINA : Pickens Co., 6 Ƥ (one without metasoma), Clemson, 31 Mar 1951 , W.R.M. Mason ( CNCI ). VIRGINIA: 1 Ƥ, Nelson Co., 25 Jun 1926 , W. Robinson ( USNM ). TABLE 1. Development of pronotal fovea in Polistes hirsuticornis sp. nov.

absent	minute, shallow	moderately developed	fairly large and deep	Total
Females	16 ( 55 %)	12 ( 41 %)	1 ( 3 %)	-	29
Males	4 ( 17 %)	11 ( 46 %)	5 ( 21 %)	4 ( 17 %)	24

TABLE 2. Morphometric parameters: females of Polistes hirsuticornis sp. nov. , P. parametricus sp. nov. and similar species. Explanations: Avg—arithmetic mean, Range—observed minimum and maximum values (ranges marked by an asterisk include measurements that were not part of the sample used for the statistical analyses), SD—standard deviation, n —number of measured specimens, THSD—Tukey's HSD (pairwise comparisons): species with identical letters have mean values that are not statistically different ( p > 0.05), F—F statistic for ANOVA within each morphometric parameter. For each ANOVA, p value was <0.001 except for CW/CL ( p = 0.294; no statistically significant differences).

Morphometric parameter—Ƥ Species	Avg	Range	SD n	THSD	F
Clypeal width / length metricus (CW/CL) bellicosus	0.98 0.99	0.91-1.04 0.95-1.04	0.029 32 0.023 25
fuscatus	0.99	0.94-1.03	0.024 24	1.25
parametricus	0.99	0.94-1.03*	0.025 25
hirsuticornis	1.01	0.96-1.06	0.025 25
Clypeus-eye contact length / Antennal hirsuticornis socket vertical diameter parametricus (CEC/AS) bellicosus	0.61 0.74 0.85	0.38-0.85 0.54-0.97 0.65-1.08	0.09 25 0.11 25 0.09 25	a b c	75.19
metricus	1.02	0.86-1.21	0.11 32	d
fuscatus	0.99	0.76-1.23*	0.11 25	d
Flagellomere 1 length / bellicosus Scape length fuscatus (FIL/SL) parametricus	0.74 0.75 0.76	0.69-0.77 0.68-0.79* 0.71-0.83	0.021 25 0.019 25 0.024 25	a b b	15.30
hirsuticornis	0.77	0.74-0.79	0.014 25	bc
metricus	0.78	0.73-0.82	0.022 32	c
Genal width / metricus Eye width bellicosus (GW/EW) fuscatus	0.73 0.76 0.78	0.48-0.85 0.67-0.90 0.67-0.92*	0.067 32 0.055 25 0.060 25	a ab bc	56.14
hirsuticornis	0.81	0.69-0.90	0.054 25	c
parametricus	0.99	0.80-1.21	0.095 25	d
Postocellar line / parametricus Ocellocular line metricus (POL/OOL) fuscatus	0.47 0.49 0.51	0.38-0.58* 0.43-0.56 0.44-0.65*	0.031 25 0.035 32 0.032 25	a ab bc	48.59
bellicosus	0.53	0.48-0.63	0.037 25	c
hirsuticornis	0.60	0.51-0.69	0.046 25	d
Scape length / width parametricus (SL/SW) hirsuticornis	3.28 3.28	3.15-3.43 2.94-3.64	0.08 25 0.15 25	a a
bellicosus	3.51	3.26-3.72	0.10 25	b	69.36
fuscatus	3.53	3.35-3.69	0.09 25	b
metricus	3.64	3.43-3.79	0.09 32	c
Upper interocular distance / metricus Head width fuscatus (UID/HW) bellicosus	0.42 0.43 0.43	0.40-0.44 0.41-0.44 0.42-0.45	0.009 32 0.008 25 0.008 25	a b b	90.18
parametricus	0.44	0.42-0.47	0.012 25	c
hirsuticornis	0.46	0.45-0.48	0.009 25	d

...... continued on the next page TABLE 2. (Continued)

Morphometric parameter—Ƥ	Species	Avg	Range	SD	n	THSD	F
Upper interocular distance /	metricus	0.91	0.87-0.96	0.024	32	a
Lower interocular distance	bellicosus	0.93	0.89-0.97	0.021	25	ab
(UID/LID)	fuscatus	0.93	0.88-0.97 *	0.025	25	b	52.29
parametricus	0.94	0.89-0.99 *	0.026	25	b
hirsuticornis	1.00	0.95-1.05	0.024	25	c
Upper interocular distance /	parametricus	3.17	2.83-3.43	0.18	25	a
Malar space (UID/MS)	hirsuticornis metricus	3.47 3.60	3.13-3.74 3.22-4.06	0.17 0.20	25 32	b b	73.37
bellicosus	3.86	3.47-4.22	0.20	25	c
fuscatus	4.03	3.62-4.68	0.24	25	d
Upper interocular distance /	metricus	1.01	0.94-1.07	0.031	32	a
Scape length (UID/SL)	bellicosus fuscatus	1.03 1.04	0.98-1.09 0.98-1.10 *	0.031 0.029	25 25	b b	83.20
parametricus	1.07	1.01-1.14 *	0.024	25	c
hirsuticornis	1.14	1.08-1.20	0.027	25	d
Number of hamuli	hirsuticornis	17.0	14-20	1.5	25	a
fuscatus	18.5	15-27	2.4	25	b
parametricus	19.3	15-24 *	2.3	25	bc	24.00
bellicosus	20.4	15-25 *	2.1	25	c
metricus	21.9	18-26	1.9	32	d

Diagnosis . Structurally distinct from all Nearctic members of Fuscopolistes based on the suite of characters mentioned in couplet 1 of the key below as well as the following ones (character states of related species in parentheses): female tergum 1 strongly and evenly convex (strongly convex anteriorly behind reception of propodeal muscle, weakly convex medially, but similar to P. hirsuticornis in some P. c a ro l in a ); female stinger slightly decurved in apical half (straight). Colouration as in dark or ferruginous forms of P . fuscatus with poorly developed yellow markings (except in midwestern specimens which have well developed yellow markings); some females (Virginia, Florida) nearly completely ferruginous and hence similar to P. c a ro l i n a . Description. Structural characters. FEMALE (IAV Figs C 80.1 , C 80.3 ). Fore wing length 13.5 –16.0 mm. For morphometric measurements see Table 2 ; differences from other species include: the relatively large upper interocular distance (UID/LID, UID/ HW), long postocellar line (POL/OOL), short malar space (UID/MS), wide clypeus (CW/CL), short clypeus-eye contact (CEC/ AS ), short scape (UID/SL, SL/SW), and small number of hamuli. Head (IAV Fig. C 80.5 ). Clypeus with apex minutely truncate, length of truncation ca. 0.75 × width of median ocellus. Clypeal disk convex, finely micropunctate except V-shaped apical area which is finely shagreened; diameter of micropunctures ca. 8 μm, much smaller than in other species ( P. f u s c a t u s : 12–16 μm); punctures near centre of micropunctate area separated by at least one puncture diameter (separated mostly by less than puncture diameter in P. fuscatus ), clypeus therefore more shining; macropunctures scattered and of variable sizes, small and inconspicuous in basal half of clypeus, becoming increasingly larger towards apex, where they are large and conspicuous (macropunctures smaller on average than in P. f u s c a t u s ); 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 bearing erect hairs whose tips are bent. 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 with widely scattered minute hairs ( 15 μm, visible at 20 × magnification). Mandible of the usual shape with three apical teeth and one receded tooth on dorsal margin; inner surface with the usual longitudinal ridges between first/second and second/third teeth; oblique, receded ridge present on undersurface of fourth tooth; outer surface bare except for scattered macropunctures of moderate size, each bearing small bristle. Antenna (IAV Fig. B 10.18 ). Flagellomeres 1–8 or 9 with relatively long, bristly hairs posteriorly and dorsally (length ca. 60 μm, with longest ones up to 90 μm; hairs becoming sparser and shorter on distal flagellomeres), these hairs absent to short (maximum length ca. 30 μm) in most other species (except sometimes in P. rubiginosus ). TABLE 3. Morphometric parameters: males of Polistes hirsuticornis sp. nov. , P. parametricus sp. nov. and similar species. Explanations: Avg—arithmetic mean, Range—observed minimum and maximum values (ranges marked by an asterisk include measurements that were not part of the sample used for the statistical analyses), SD—standard deviation, n —number of measured specimens, THSD—Tukey's HSD (pairwise comparisons): species with identical letters have mean values that are not statistically different ( p > 0.05), F—F statistic for ANOVA within each morphometric parameter. For each ANOVA, p value was <0.001 except for CW/CL ( p = 0.051; no statistically significant differences).

Morphometric parameter— 3	Species	Avg	Range	SD	n	THSD	F
Clypeal width / length	parametricus	0.98	0.92-1.04 *	0.028	22
(CW/CL)	bellicosus	0.99	0.94-1.07	0.033	25
metricus	1.02	0.93-1.10 *	0.036	25	2.44
fuscatus	1.02	0.93-1.12	0.046	25
hirsuticornis	1.03	0.99-1.11	0.033	24
Flagellomere 1 length /	hirsuticornis	0.89	0.83-0.94	0.029	24	a
Scape length (FIL/SL)	bellicosus parametricus	0.92 0.94	0.87-0.96 0.89-0.99	0.022 0.026	25 22	b bc	44.21
fuscatus	0.96	0.90-1.02	0.029	25	c
metricus	0.99	0.93-1.06	0.032	25	d
Flagellomere 10 width /	parametricus	0.78	0.71-0.82	0.027	21	a
Flagellomere 1 width (FXW/FIW)	hirsuticornis metricus	0.79 0.81	0.75-0.83 0.73-0.93 *	0.021 0.035	24 25	a ab	12.66
bellicosus	0.88	0.82-0.94	0.033	25	b
fuscatus	0.91	0.85-0.97	0.035	25	b
Genal width /	hirsuticornis	0.80	0.70-0.88	0.043	23	a
Eye width (GW/EW)	fuscatus bellicosus	0.83 0.83	0.75-0.96 0.72-1.01	0.054 0.078	25 25	ab ab	3.87
parametricus	0.87	0.75-1.03 *	0.070	22	b
metricus	0.87	0.75-0.97	0.063	25	b
Postocellar line /	parametricus	0.48	0.39-0.61	0.048	22	a
Ocellocular line (POL/OOL)	metricus bellicosus	0.51 0.54	0.44-0.66 * 0.40-0.63	0.044 0.056	25 25	ab ab	22.42
fuscatus	0.54	0.44-0.68 *	0.056	25	b
hirsuticornis	0.61	0.55-0.71	0.039	24	c
Scape length / width	hirsuticornis	3.07	2.92-3.25	0.08	24	a
(SL/SW)	parametricus	3.20	3.00- 3.56	0.12	22	b
fuscatus	3.33	3.02-3.53	0.11	25	c	52.04
bellicosus	3.42	3.11-3.61	0.12	25	d
metricus	3.45	3.17-3.76 *	0.11	25	d

...... continued on the next page TABLE 3. (continued)

Morphometric parameter—3 Species	Avg	Range SD n	THSD	F
Upper interocular distance / Flagellomere metricus 1 length fuscatus (UID/FIL) bellicosus	0.92 0.97 1.01	0.82-0.99* 0.044 25 0.89-1.06 0.042 25 0.96-1.10 0.039 25	a b c	117.94
parametricus	1.02	0.91-1.10 0.046 22	c
hirsuticornis	1.18	1.11-1.29* 0.046 24	d
Upper interocular distance / fuscatus Head width metricus (UID/HW) bellicosus	0.41 0.41 0.41	0.39-0.43 0.010 25 0.39-0.43 0.011 25 0.39-0.43 0.011 25	a a a	35.28
parametricus	0.41	0.39-0.43 0.011 22	a
hirsuticornis	0.44	0.42-0.46 0.010 24	b
Upper interocular distance / fuscatus Lower interocular distance bellicosus (UID/LID) metricus	0.93 0.94 0.95	0.89-1.01* 0.024 25 0.87-1.03 0.044 25 0.87-1.01 0.037 25	a a a	30.80
parametricus	0.97	0.92-1.04 0.028 22	a
hirsuticornis	1.03	0.96-1.07 0.026 24	b
Upper interocular distance / parametricus Malar space bellicosus (UID/LID) metricus	2.58 2.59 2.63	2.32-2.82 0.13 22 2.25-2.83 0.16 25 2.28-2.87 0.16 25	a a a	25.97
fuscatus	2.80	2.49-3.05 0.16 25	b
hirsuticornis	2.96	2.57-3.19 0.16 24	c
Upper interocular distance / metricus Scape length fuscatus (UID/SL) bellicosus	0.91 0.93 0.93	0.85-0.97 0.032 25 0.87-1.00 0.036 25 0.89-0.99 0.026 25	a a a	69.10
parametricus	0.96	0.90-1.01 0.029 22	a
hirsuticornis	1.05	1.00-1.13 0.036 24	b
Number of hamuli hirsuticornis	16.5	13-22 2.3 24	a
fuscatus	18.4	10-24* 2.6 25	ab
parametricus	18.6	12-24 2.6 22	b	11.21
bellicosus	19.3	15-26 2.6 25	b
metricus	21.7	16-29 3.4 25	c

Mesosoma (IAV Fig. C 80.7 ). Pronotum with fovea absent (IAV Fig. B 10.14 ) or represented by a small, superficial depression, exceptionally (one female from South Carolina ) with a small but fairly deep fovea (not penetrating cuticle as usually in P. fuscatus ) ( Table 1 ). Pronotal carina somewhat shortened laterally, becoming evanescent shortly below level of pronotal lobe; median portion of carina (behind ocellar triangle) more strongly arched than paramedian sections which are almost straight before curving ventrally at humeral angle (carina more or less evenly curved dorsally between humeral angles in P. fuscatus ); ventral third of pronotum almost completely flat (with a very shallow groove in continuation of pronotal carina in P. f us c a tu s ). Mesosoma more shining than in other species, punctures smaller and interspaces larger. Diameter of mesopleural micropunctures ca. 8 μm, separated by about their own width, interspaces shining (in P. fuscatus micropuncture size ca. 14–16 μm, separated by about half their width, interspaces less shining); macropunctures twice the size of micropunctures, scattered and inconspicuous, each giving rise to a longer hair with bent apex. Dorsal groove of mesopleuron on average weaker than in P. f u s c a t u s , median third very faintly impressed or not impressed at all (but often clearly marked by a dark line). Metapleuron below metapleural pit similar in sculpture to mesopleuron, hardly more shining than latter (in P. fuscatus with much smaller micropunctures and more shining than mesopleuron). Scutum with scattered and inconspicuous longer hairs between regular pubescence, apices of these hairs bent at apex; macropunctures very inconspicuous. Scutellum with the usual faint median longitudinal keel, 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 punctate laterally (weakly microstriate in P. fuscatus ), median half becoming increasingly ridged posteriorly, with microstriae and a few delicate ridges, (ridges much weaker than in coarsely ridged species such as P. rubiginosus ), depressed median groove highly shining; 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 (IAV Fig. B 10.16 ) of unusual shape, wide ventrally, dorsal third narrowed fairly abruptly to about one third of ventral width, sides of dorsal section subparallel with narrowly rounded apex (in other Fuscopolistes narrowed gradually with upper width at least half of lower width, usually more). Legs and wings without unusual features, morphology essentially as in P. fuscatus , except fore coxae slightly more shining. Metasoma (IAV Fig. C 80.1 ) with tergum 1 1.06–1.19 × as wide as long (length measured from attachment of petiole muscle to apex of tergum); anterior half of tergum moderately and evenly curved in lateral view with degree of convexity gradually decreasing posteriorly (in P. fuscatus typically rising steeply for a short distance behind attachment of propodeal muscle, then fairly abruptly levelling off onto weakly curved posterior surface; IAV Fig. C 77.4 ). 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 distinct 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 ), finely and fairly densely micropunctate (similar to tergum), macropunctures more numerous than on tergum, with macrosetae longer and fairly conspicuous. Punctation of following sterna becoming slightly sparser and slightly coarser (except sternum 6 which is more finely and densely punctate than the previous sternum); macropunctures fairly numerous, in some specimens quite distinct and obvious; macrosetae becoming longer towards apex of metasoma. Sternum 6 with the usual median tuft of hairs at extreme base (usually concealed by previous sternum). Stinger with apical half slightly to moderately decurved (straight or nearly straight in other species). Each stinger lancet with ca. five fine barbs near apex, as in other species. MALE (IAV Figs C 80.2 , C 80.4 ) characters as in female except as follows. Fore wing length 12.5–15.5 mm . For morphometric measurements see Table 3 ; differences from other species include (similar as in female): the relatively large upper interocular distance (UID/LID, UID/HW), long postocellar line (POL/OOL), short malar space (UID/MS), wide clypeus (CW/CL), short scape (UID/SL, SL/SW), short FI (FI/SL), and small number of hamuli. Head (IAV Fig. C 80.6 ). Clypeus usually at least narrowly separated from eye (at most by ca. 0.4 MOD ), in two specimens in contact with eye for a distance of up to 0.6 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. fuscatus very slightly concave, rarely flat); surface entirely and densely micropunctate with weakly shining interspaces, macropunctures scattered, those in dorsal third of clypeus bearing pale, longer macrosetae with bent apices, those in lower 2 / 3 inconspicuous, with short and straight macrosetae; diameter of micropunctures about the same as in female but punctures closer together and clypeus therefore less shining (only slightly more shining than in males of P. f u s c a t u s ); 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 P. fuscatus ; subantennal longitudinal grooves usually less impressed than in P. f u s c a t u s , with outer margin more rounded (usually more carinate in P. fuscatus ). 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. fuscatus ), macropunctures smaller than in female and only bearing weak hairs; inner surface of mandible without ridges. Labrum narrow and parallelsided (width ca. 0.5 MOD ), with several small bristles near the narrowly rounded apex. Antenna differs from P. fuscatus as follows ( Table 3 ): scape almost always stouter, flagellomere 1 almost always shorter, apical flagellomeres more slender, flagellomere 10 narrower, and flagellomere 11 usually less tapered than in P. fuscatus . Male flagellum longer than in female, apically curved backwards as in other Fuscopolistes . Flagellomere 1 with outstanding hairs very short and inconspicuous (length ca. 35 μm, similar to P. f u s c a t u s ); following flagellomeres with hairs even shorter or absent. Tyloids similar to P. f u s c a t u s , almost linear on flagellomere 1 , becoming increasingly wider towards flagellomere 7 where tyloid covers entire ventral surface (as in flagellomeres 8–11 ); micropubescence of tyloids becoming sparser distally, apical four flagellomeres with large, shining, bare patches. Mesosoma. Pronotum with fovea quite variable, on average better developed than in female ( Table 1 ): usually small and shallow, rarely absent, sometimes moderately developed, in four specimens fairly large and deep but not penetrating cuticle as usually in P. fu s ca t us . Ventral extension of pronotal carina also slightly better developed than in female, extending close to ventral corner of pronotum as a weak rounded ridge. Metasoma with tergum 1 1.00– 1.12 × as wide as long. Sternum 2 less convex than in female (similar to P. fuscatus ), 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 ). 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. f u s c a t u s ). 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 ). Genitalia (for morphometric parameters see Table 4 ). Paramere and volsella of the usual shape ( Fig. 20 ). Paramere with base of spine haired medially and laterally. Parameral spine with the usual gland openings: one at base dorsolaterally, and one near middle laterally which extends as a groove towards apex. Digitus ( Fig. 28 ) with apical process almost straight to slightly curved, inner surface of process sometimes with 1–5 hairs; inner surface of main body with numerous long hairs ( Fig. 37 ), outer surface with the usual black, scale-like bristles; preapical ventral curvature very weak (unlike P. fuscatus and P. metricus ); anteroventral lobe usually rounded ( Fig. 30 ), in one specimen slightly pointed ( Fig. 31 ). Cuspis ( Fig. 20 ) as in other species, with long, retrorse hairs at apex and along dorsal margin of distal half, a cluster of black, scale-like bristles at dorsal angle, moderately long hairs on basal prominence, and short hairs along ventral margin and dorsal margin of basal half. Aedeagus ( Fig. 45 ) relatively slender, lateral projections of median expansion variable, quite tapered in one specimen, more robust in others ( Fig. 61 ). Row of teeth on ventral edge of penis valve extending anteriorly about halfway onto median expansion ( Figs 53 , 65 ); teeth in one well aligned row. Teeth inclined anteriorly, of moderate size, unequal, usually one large tooth alternating with one slightly smaller tooth (in one specimen with only a few smaller teeth). Colouration . See detailed description in Buck et al . ( 2008 : "sp. A"). Remarks . Based on the presence of several unique morphological features (absent or poorly developed pronotal fovea, fine mesopleural punctation, shape of propodeal orifice, haired female flagellum, sparsely haired eyes, curved stinger) and unusual body proportions (expressed by extreme values for most morphometric parameters, see Tables 2 , 3 ) P. hirsuticornis is one of the most distinctive species in the subgenus Fuscopolistes . However, this morphological distinctiveness is masked by a highly variable colour pattern that mimics that of several other sympatric species. Since Polistes taxonomy has traditionally relied mostly on colour patterns P. hirsuticornis has been confused with no less than four species: P. fuscatus , P. metricus , P. bellicosus and P. carolina . Most females of P. hirsuticornis resemble sympatric populations of P. fuscatus , which in turn has various colour forms that closely resemble the other three mentioned species. Females that are similar to P. bellicosus occur mostly in the Midwest where P. bellicosus is absent. Females with patterns like P. m e tr i c us (or the newly described P. p a r a m e t r i c u s , see below) occur in the northeastern and east-central part of the range. Specimens that look like P. carolina or extreme ferruginous forms of P. fuscatus have been found in Virginia and Florida. The situation is different in males. Since the male of P. fuscatus has all-dark apical flagellomeres (see key below) it is immediately differentiated from all other species including P. hirsuticornis . Furthermore, P. hirsuticornis has less extensive yellow markings on head, meso- and metasoma than P. f u s c a t u s . Males of P. hirsuticornis look therefore most like P. m e t r i c u s (which is usually larger in size) and particularly P. parametricus (which usually has darker brown apical flagellomeres, see description below). Unfortunately, no males from the southern and western part of the range are available. They are probably patterned like dark forms of P. carolina and P. bellicosus , respectively. TABLE 4. Male genitalia: morphometric parameters of aedeagus, digitus and paramere of eastern Nearctic Fuscopolistes . Explanations: Avg—arithmetic mean, Range—observed minimum and maximum values, SD—standard deviation, n —number of measured specimens. P . bellicosus WA is a form with wide aedeagus that might be a different species than the typical form. " Polistes carolina " possesses two different forms that correspond to separate clusters in the genetic analysis (see discussion of molecular study). Morphometric parameter Species Avg Range SD n State/Province Etymology . This species is named for the unusually long bristly hairs of the basal female flagellomeres.

Aedeagus: bellicosus WA Length from base of median fuscatus expansion to apex / Width of median expansion (AL/MEW) carolina A metricus	1.61 1.68 1.80 1.82	1.54–1.67 0.09 1.56–1.82 0.09 1.69–1.88 0.10 1.69–1.94 0.12	2 10 3 5	AR, FL AR, FL, MD, TX, ON AR, SC FL, MD, MO, TX
bellicosus	2.00	1.88–2.11 0.10	6	FL, MD, TX
rubiginosus	2.14	2.00–2.22 0.08	5	MO, TX
parametricus	2.17	2.14–2.22 0.05	3	MD
hirsuticornis	2.24	2.08–2.50 0.22	3	MD
carolina X/TX	2.24	2.11–2.40 0.11	5	FL, TX
dorsalis	2.63	2.38–3.00 0.23	5	FL, MD, TX
Aedeagus: bellicosus WA Length from base of median metricus expansion to apex / carolina A Minimum width near apex (AL/MPW) fuscatus bellicosus	5.1 5.3 5.5 5.5 7.1	4.8–5.5 0.46 5.0–6.0 0.38 5.0–6.0 0.50 4.4–6.3 0.59 6.1–8.3 0.78	2 5 3 10 6	as above
hirsuticornis	7.6	6.3–10.0 1.65	3
rubiginosus	8.3	7.1–10.0 1.18	5
parametricus	8.9	8.0–10.0 1.03	3
carolina X/TX	9.6	8.3–11.5 1.27	5
dorsalis	11.5	10.0–15.4 2.24	5
Aedeagus maximum tooth length (μm) dorsalis rubiginosus hirsuticornis carolina A	35.0 36.8 38.5 42.8	32–40 3.3 32–40 3.4 32–40 4.7 40–47 3.4	5 5 3 4
bellicosus WA	43.5	42–45 2.1	2	as above
parametricus	44.0	40–48 4.0	3
carolina X/TX	44.6	39–48 4.0	5
bellicosus	45.5	42–48 2.5	6
metricus	48.4	44–50 3.1	5
fuscatus	71.0	64–88 10.5	10
Digitus: fuscatus Apical process length / Total length metricus bellicosus	0.17 0.18 0.21	0.10–0.21 0.032 0.15–0.20 0.021 0.18–0.24 0.026	10 5 6
carolina A	0.22	0.21–0.24 0.013	4
bellicosus WA	0.23	0.21–0.24 0.021	2	as above
parametricus	0.23	0.21–0.27 0.038	3
carolina X/TX	0.23	0.19–0.28 0.032	5
hirsuticornis	0.24	0.21–0.26 0.021	3
rubiginosus dorsalis	0.26 0.26	0.23–0.29 0.024 0.24–0.28 0.018	5 5

...... continued on the next page TABLE 4. (continued)

Morphometric parameter Species	Avg	Range SD	n	State/Province
Paramere length (excl. spine) / height carolina A	2.25	2.24–2.26 0.01	2
fuscatus	2.39	2.20–2.61 0.13	10
bellicosus	2.42	2.31–2.54 0.10	6
bellicosus WA	2.3 5	1
parametricus	2.42	2.38–2.45 0.04	3	as above
hirsuticornis	2.43	2.29–2.58 0.12	3
metricus	2.49	2.32–2.82 0.19	5
dorsalis	2.50	2.28–3.00 0.29	5
carolina X/TX	2.51	2.22–2.76 0.22	5
rubiginosus	2.57	2.32–3.00 0.28	5

Distribution ( Fig. 1 ). The distribution of this species is insufficiently documented, especially in interior areas of the eastern United States . It appears likely that the northernmost and southernmost currently known localities (coastal areas of New York and northern Florida) are in fact close to the northern and southern distribution limit of this species. We have examined large numbers of Polistes from central and south Florida but did not find any P. hirsuticornis . The westernmost currently known specimens are from central Missouri. We suspect that the species ranges from the Atlantic all the way to easternmost parts of Texas, Oklahoma and perhaps Kansas (likely absent from most of the Appalachians area). FIGURE 1 . Locality records for Polistes hirsuticornis sp. nov. (triangles) and P . parametricus sp. nov. (circles). Phenology ( Fig. 5 ). Females have been collected from the second half of March to the end of September except for the month of July. Male specimens occur from the end of August to the beginning of October. The unusual phenology of females is discussed below in the section "Suspected social parasitism". Parasites and symbionts . One female from Florida (see material examined) carries an eclosed male Strepsiptera pupa between terga 3 and 4 . We assume it belongs to a species of the genus Xenos ( Stylopidae ), which are known to parasitize several other sympatric species of Polistes . Another female from Florida (see material examined) carried> 20 deutonymphs of Sphexicozela (Acari: Winterschmidtiidae ) (det. D.E. Walter, 4 specimens mounted on slides). This genus currently includes only one named species from Europe ( Mahunka, 1970 ), and our specimens probably belong to an undescribed species. Sphexicozela mites have only been found in association with Polistes , but their biology is poorly known ( O'Connor, 1994 , and in litt .). Deutonymphs are usually found in the genital chamber between the last tergum and sternum (as in our case), and are therefore easily overlooked. Various stages of the mite (larvae, protonymphs, deutonymphs and adults) occur in nests and on larvae or pupae of Polistes ( Mahunka, 1970 ; Rusina & Orlova, 2011 ), but their feeding behaviour has not been studied. It is not clear whether they feed on the prey of the wasps or on immature stages of the wasps themselves (O'Connor, in litt .).