Colletes ultravalidus Hall & Ascher, 2016

Hall, H. Glenn, Almquist, David T. & Ascher, John S., 2016, A new species of Colletes (Hymenoptera: Apoidea: Colletidae) from northern Florida and Georgia, with notes on the Colletes of those states, Journal of Melittology 2016 (58), pp. 1-20 : 3-11

Colletes ultravalidus Hall & Ascher , new species

ZooBank: urn:lsid:zoobank.org:act:AC1D4A58-9479-4557-BA0C-0F30437DE68F

( Figs. 1 , 3 , 5 , 7 , 9 , 11 , 13 , 15 )

DIAGNOSIS: Both female and male of C. ultravalidus are distinguished from those of all other eastern North American species of the genus by their exceptionally long malar area, especially the male. This is a large bee with dense fulvous mesosomal pubescence, dark metasomal terga with apical fasciae absent or weak, and strongly infuscated wings. Color varies, and some specimens have paler pubescence. Colletes ultravalidus is most similar to, but somewhat larger than, C. validus Cresson ( Figs. 2, 3 ). The malar area exceeds in length that of C. validus , previously considered the Nearctic species with the longest malar area within the genus. The proportionately longer malar area comes at the expense of a proportionately shorter compound eye ( Figs. 4–11 ; Table 1 ). In contrast to C. ultravalidus , C. validus has pronounced apical fasciae on the metasomal terga, ochraceous rather than fulvous pubescence, most notable on the mesosoma, and lighter wings. Colletes ultravalidus tends to have more fuscous hairs intermixed with light hairs, for example on the legs of females. Colletes ultravalidus resembles C. thoracicus Smith , as both have abundant mesosomal fulvous pubescence, strongly infuscated wings, and lack metasomal fascia in females. However, both sexes of C. thoracicus are easily distinguished from C. ultravalidus by their much shorter malar area (less than half the width of the mandible base in females, subequal in males) and shorter clypeus. Females of C. thoracicus are slightly smaller and more slender, males more so. Three other species of Colletes in the eastern United States have notably long malar areas, C. longifacies Stephen , C. productus Robertson , and C. compactus Cresson. All have conspicuous tergal fascia, an average smaller body size than C. ultravalidus , and malar area not exceeding the width of the mandible at its base, thus much shorter than the malar area of either C. ultravalidus or C. validus . The S7 of C. ultravalidus and C. validus males have a nearly identical shape which separates them from other species of Colletes ( Figs. 12, 13 ). Genital armatures of the two species differ subtly in the position of the dorsal wings ( Figs. 14, 15 ).

DESCRIPTION: ♀: Structure. Body length 14.27± 0.98 mm, inter-tegular distance 3.62± 0.05 mm (n=7) [13.37± 0.55 mm and 3.17± 0.17 mm (n=6), respectively, for C. validus ]. Head 3.78± 0.18 mm long (n=7), length about 0.9x head width. Mandibles strongly crossed in ventral view, one extending nearly one-fifth its length beyond side of other. Clypeus, 0.44x head length, extends below mandible articulation by 0.5x length of malar area, strongly protuberant, with moderately deep median sulcus; apical margin with paired, variably shaped, depressions. Supraclypeal area strongly protuberant. Malar area about 0.31x length of head, 0.49x length of compound eye, almost 1.9x width of mandible base [0.25x, 0.37x, and 1.66x respectively for C. validus ]. Vertex excavated lateral to ocelli. Antenna F2 about 0.8x length of F1, F2–F9 subequal in length, length subequal to width, F10 about 1.1x length of F1. Compound eye about 0.57x length of head [0.63x for C. validus ], inner margin subparallel ( UOD /LOD ratio = 0.99). Lateral ocelli slightly closer to each other than to compound eyes. Facial fovea short, wide, and shallow, lower margin about one-third distance from antenna to lateral ocellus, upper margin nearly level with lower margin of lateral ocellus. Facial fovea curves from inner orbital margin toward lateral ocellus, ending about half distance between these two. Genal area about 1.2x width of eye. Pronotal ridge with short, lateral angle, with wide base. Metapleural protuberance acarinate. Fore coxal spine lacking. First recurrent vein of forewing meets center of second submarginal cell. Posterior basitarsus 3.9x as long as broad. T 1 slightly impressed narrow apical rim; T 2– T 5 apical impressed area widens from about one fourth to one third of exposed surface of segment; S1 with narrow, central, rounded, elevated protuberance; S5 with wide, shallow, semicircular emargination.

Pubescence and Color. Integument nearly all black/piceous. Labrum and mandible with long, thick, erect, simple, fuscous hairs along ventral edge of mandible and apex of labrum, may be intermixed with few light hairs [pubescence all light in C. validus ]. Lower paraocular area with sparse, short, ochraceous, plumose hairs intermixed with few, longer, fuscous, simple or somewhat branched hairs. Upper paraocular area and frons with dense, long, fulvous-brownish, plumose hairs intermixed with smaller number of fuscous hairs between and lateral to antennae, relatively bare between antennal base and vertex. Vertex with long, plumose fuscous hairs intermixed with shorter light hairs; more dense row of long light pubescence extending from upper occiput along upper margin of vertex. Genal area with sparse, long, fuscous, simple or somewhat branched hairs below and adjacent to compound eyes, mixed with long, fulvous, plumose pubescence above, more dense toward occipital ridge. Mesosoma with dense, light, fulvous hairs on pronotal lobe, as row along omaulus, on lateral mesoscutum anterior to wings, on metanotum, and on boundary between lateral and posterior surfaces of propodeum. Sparser, light, fulvous pubescence, intermixed with varying amounts of fuscous pubescence, on disc of mesoscutum [ Stephen (1954) describes presence of abundant black pile on median discal areas of mesoscutum and mesoscutellum of C. validus , whereas these areas largely worn bare in both C. ultravalidus and C. validus specimens investigated here, with fulvous pubescence mainly on mesoscutum anteriorly and mostly long, erect, fuscous pubescence along posterior margin of mesoscutellum]. Mesepisternum with long, light pubescence, sparse below wings, more dense ventrally [distribution of hairs on mesepisternum of C. validus similar, but pubescence ochraceous to white versus light fulvous in C. ultravalidus ]. Leg pubescence generally dark brown on basal segments, becoming dark brown or fuscous on distal segments [variable, three FSCA specimens with light leg pubescence; in C. validus light brown pubescence on basal segments, becoming ochraceous or white on distal segments]. Wings deeply infuscated [lightly infuscated in C. validus ], veins and stigma piceous. T 1 with fulvous hairs long and sparse on anterior declivitous surface, denser laterally; in some specimens fringe of pale ochraceous hairs along outermost lateral margin [hair distribution similar to that described by Stephen (1954) for C. validus , pubescence light ochraceous and less dense laterally]. Apical tergal fascia entirely or mostly absent, but better developed in some specimens with appressed, isolated, individual hairs or weak, fragmented, patches of tomentum along apical rim, sometimes more substantial laterally and across T 4 and T 5 rims [moderately wide and dense, apical fascia present in C. validus , interrupted medially, broadly on T 1 narrowly on T 2, entire on T 3– T 5; Stephen (1954) described sparse, long, erect, black pile on tergal discs of C. validus , whereas pile very short and light on discs of specimens examined]. Unworn specimens of C. ultravalidus have very sparse, short fuscous and light pubescence on tergal discs. T 2– T 5 pubescence erect, simple, light, short laterally, becoming sparser, darker, and longer from anterior to posterior; apical rims black or faint amber [impunctate tergal rims of C. validus noticeably amber where fascia has been lost]. T 6 with dense, appressed, fuscous pubescence covering disc; erect, long, fuscous pubescence laterally. S1 protuberance with long fulvous hairs. S2–S5 with sparse fascia arising along basal margin of narrow, impressed, amber, apical rim; pubescence becoming denser, longer, lighter laterally; disc with appressed, short, sparse, light and sparse, erect, fuscous pubescence; light hairs of disc become increasingly longer, denser laterally. S6 with long, thick, appressed, fuscous hairs intermixed with fewer light hairs on disc.

Surface Sculpture. Clypeus with large and small deep punctures, widely and unevenly separated along entire length; punctures much denser, somewhat elongate and separated by one to two puncture diameters in median sulcus and along margin adjacent to lower paraocular area; interspaces smooth and shiny over nearly entire length; extreme base with elongate, dense punctures and/or rough texture. Supraclypeal area densely and coarsely punctured except center and lower margin; interspaces faintly imbricate except strongly imbricate along upper and upper-lateral margins and lineolate laterally and on lower margin. Malar area with few (~18) widely spaced, deep punctures; interspaces smooth and shiny, faintly lineolate just above mandible base and just below compound eye. Lower paraocular area with narrow rugulose-punctate strip of punctures, irregular in size and shape with some elongate, adjacent to and delineated from smooth malar area; punctures becoming large, deep, dense interiorly, spaced by less than one puncture diameter, more elongate lateral to antenna; interspaces shiny. Upper paraocular area rugulose-punctate with smaller, nearly confluent, elongate, obscure punctures; interspaces rough. Frons with large, elongate, confluent punctures. Vertex with smooth patch above compound eye and fovea, with widely spaced large punctures and fine, small punctures inbetween; obscure punctures between ocelli and along upper margin of vertex; surface rough between ocelli, imbricate along upper margin. Genal area densely punctured along orbit and between eye and preoccipital ridge, lineolate along hypostomal cavity. Pronotum dorsal surface and preomaular area with widely spaced punctures, evenly imbricate. Mesoscutum with deep, dense punctures, nearly contiguous on anterior half, laterally, and along posterior margin; punctures larger, less dense on disc. Mesoscutellum with dense, deep punctures, somewhat elongate, laterally and posteriorly, widely spaced anteriorly with shiny interspaces; imbricate along anterior-most margin and posteriorly [in C. validus punctures more poorly defined and somewhat denser on anterior of mesoscutum and posterior of mesoscutellum]. Tegula with fine punctures separated by two to three puncture diameters. Mesepisternum with deep, dense punctures, well defined on hypoepimeral area, separated by no more than one puncture diameter; surface somewhat rough, although shiny; punctures more obscure, surface increasingly rough and lineolate ventrally. Metapleural protuberance with shallow obscure punctures; surface very rough and somewhat rugulose. Metepisternum rugulose ventrally with slanted rugae dorsally. Metanotum finely rugulose. Metapostnotum with vertical (longitudinal), evenly spaced rugae forming a transverse row of shiny pits. Propodeum, lateral surface with few indistinct, very widely spaced punctures; granulate; posterior surface rugulose. T 1 with declivitous anterior surface nearly impunctate; deep large punctures on disc, separated by one to three puncture diameters, becoming smaller and denser along slightly impressed apical rim; interspaces shiny; apical-most narrow margin impunctate. T 2– T 5 with punctures somewhat smaller than those on T 1, denser basally and along narrow, median area; finer punctures along impressed apical rim; interspaces shiny. Metasomal sterna with widely-spaced punctures separated by three to six puncture diameters, finer and denser on posterior segments; interspaces imbricate.

♂: Structure. Body length 12.62± 0.36 mm, inter-tegular distance about 2.9± 0.14 mm (n=6) [somewhat larger than C. validus , 11.76± 0.25 mm and 2.71± 0.27 mm (n=7), respectively]. Form more slender than female. Head about 3.66± 0.11 mm long, length about 0.97x head width. Clypeus about 0.44x length of head. Malar area about 0.31x length of head, 0.53x length of compound eye, about 2.3x width of mandible base [0.26x, 0.41x, and 1.92x, respectively for C. validus ]. Antennae, F2 about 1.5x length of F1, F2–F9 subequal in length, and length about 1.5x their width, F10 about 1.5x length of F1. Compound eyes about 0.54x length of head [0.59x for C. validus ], inner margins very slightly convergent below ( UOD /LOD ratio = 1.1). Facial fovea shallow, shorter than in female, upper margin nearly level with lower margin of middle ocellus. Genal area width about 0.9x width of eye. Mandibles, clypeus, ocellar spacing as in female. Posterior basitarsi 4.8x as long as broad. Mesosoma and metasoma structure as in female. S5 lacks median emargination present in female. Genital armature with dorsal wing of penis valve (terminology from Stephen, 1954) slightly more distal than that of C. validus ( Figs. 14, 15 ).

Pubescence and Color. Generally greater percentage of fuscous hairs relative to fulvous hairs than in female. Integumental color as in female. Clypeus with dense, long, fulvous, plumose pubescence on dorsal third drape over middle third. Lower paraocular area with long, fulvous hairs and a few scattered, fuscous, simple or somewhat branched hairs laterally. Upper paraocular area and frons with dense, mixed brown and fuscous pubescence around and just above antennal base, relatively bare between just above antennal base and vertex. Antenna with brown, short black hairs on scape longer and more apparent than on female [light, short, and less apparent in C. validus males and females]. Vertex with dense, mostly fuscous pubescence mixed with fulvous hairs between ocelli and along upper margin. Genal area with long, fulvous, plumose pubescence below; hairs shorter above, and denser toward occipital ridge, intermixed with a few, fuscous, simple or somewhat branched hairs, mainly along orbit. Mesosomal hair distribution and color similar to female, darker and somewhat denser on mesoscutum but not obscuring surface. Legs with dense, fulvous, plumose pubescence on posterior of femur, somewhat less dense on hind femur; hairs simple on tibia and tarsomeres and slightly darker on some specimens. Wings deeply infuscated, somewhat lighter than females [clear in C. validus males]; veins and stigma piceous. T 1 anterior and lateral hair distribution similar to female but lacking lateral fringe of light pubescence; erect, short, light and dark hair mixture on disc. T 1– T 5 with sparse, appressed, short, light tomentum forming weak, fragmented, apical fascia, more substantial than on female [fascia on C. validus males more dense and complete]. T 2 and T 3 discs with abundant short, erect, fuscous, simple hairs, intermixed with sparse longer erect fuscous hairs on T 4 and T 5. T 6 and T 7 with decumbent long hairs over disc, dark on T 6, mixed with fulvous on T 7. Metasomal sternal hair distribution similar to female but longer and much denser overall. S2–S5 with long plumose fulvous hairs on discs becoming progressively shorter and confined to apical center on posterior segments, flanked by short erect fuscous pubescence on remainder of discs. S4 and S5 with simple, long, erect, light pubescence laterally.

Surface Sculpture. Clypeus surface largely hidden by pubescence; apical threequarters more densely punctate, especially basally and laterally, than in female; patches of elongate, contiguous punctures adjacent to rugulose-punctate areas. Supraclypeal area more densely and evenly punctate than in female; punctures large, deep and contiguous. Malar area shiny and sparsely punctate as in female, more distinctly lineolate on lower fourth above mandibles and just below compound eyes. Lower paraocular area more evenly punctate than in female, punctures separated by no more than one puncture diameter adjacent to malar area; interspaces shiny, surface rugulose and rough; punctures become contiguous toward antennal bases. Upper paraocular area rugulose-punctate with large, deep, somewhat elongate punctures; surface rough. Frons with large, deep, round, confluent punctures. Vertex as in female but smooth patch above compound eyes not well defined. Genal area similar to female but punctures more obscure and surface rougher. Mesoscutum and mesoscutellum punctures as in female but spaced somewhat more evenly. Mesepisternum and metepisternum as in female. Terga similar to female; punctures slightly more dense and more evenly separated; T 4– T 7 interspaces with faint imbrication becoming more pronounced posteriorly. Metasomal sterna similar to female.

HOLOTYPE: ♀, USA [ United States of America]: FL [Florida]: Franklin County: Apalachicola , 29.755084N 85.100426W, 27 January 2013. D. T. & L.D. Almquist (deposited in AMNH).

ALLOTYPE: ♂, USA: FL: Wakulla County, St. Marks NWR [National Wildlife Refuge], Medart, 30.057731N 84.386362W, 29 December 2013. D. T. & L.D. Almquist (deposited in AMNH).

PARATYPES: 1♀, USA: FL: Levy County: Goethe State Forest , 10 mi Road, 29.12139N 82.589167W, 2 April 2011, H.G. Hall (deposited in HGH collection). All the following caught at same location by same collectors as allotype : 2♀♀, 10 February 2013 (1 each deposited in SNMNH and FSCA) ; 1♀, 24 February 2013 (deposited in AMNH) ; 2♀♀, 11 March 2013 (1 each deposited in FSCA and HGH collection) ; 2♂♂, 29 December 2013 (1 each deposited in AMNH and SNMNH) ; 2♂♂, 12 January 2014, C. Siervogel additional collector (1 each deposited in FSCA and HGH collection) ; 8♀♀, 22 March 2014, HGH additional collector (2 deposited in AMNH, 2 in FSCA, 1 in HGH collection, 3 in SNMNH) .

ADDITIONAL MATERIAL EXAMINED: 1♀, Okaloosa County, Eglin AFB, 13 February 1993, P. Skelley, M. Thomas, R. Turnbow ( FSCA); 1♀, Taylor County: Steinhatchee , 27 February 1993, Lloyd Davis ( FSCA); 1♀, USA: GA [Georgia]: Brantley County : Hoboken, 16 February 1997, P. Skelley, ( FSCA); 1♀, USA: FL: Franklin County: near Apalachicola, first specimens discovered could not be relocated at time of description (Cornell University Insect Collection) .

DISTRIBUTION: Along the Gulf Coastal areas of northern Florida, extending to southeastern Georgia, in sandhill communities adjacent to wetlands ( Fig. 16 ).

ETYMOLOGY: The specific epithet validus, Latin for “powerful”, refers to the similar species C. validus , with the prefix ultra from Latin, “beyond”, referencing the longer malar area of the new species.

COMPARATIVE MATERIAL EXAMINED fOR COLLETES VALIDUS : USA: NORTH CAROLINA: Sampson County: 1♀, Ivanhoe, 15 March 1949, T. B. Mitchell ( FSCA); USA : NEW JERSEY: Burlington County: 3♂♂, Lebanon State Forest , 27 April 1974, C. Porter ( FSCA); Ocean County: 1♀, 2♂♂, 1 June 1973, J.G. & K.C. Rozen, R. McGinley, C. Thompson, ( AMNH _ BEE 18759 , 18761,18764 ); USA: MASSACHUSETTS: Dukes County, Martha’s Vineyard : 1♂, Manuel F. Correllus State Forest, 7 May 1994, P.Z. Goldstein ( AMNH _ BEE 131834 ); 1♀, 2♂♂, Aquinnah, 41 19.122 N 70 48.100 W, 30 April 2011 – 27 May 2011, P.Z. Goldstein ( AMNH _ BEE 170009 , 170008,170006 ); 1♂, Edgartown, 1–3 May 2011, P.Z. Goldstein ( AMNH _ BEE 166683 ) .

Figures

Figure 1. Photograph of female of Colletes ultravalidus, new species.

Figure 16. Map of collection sites (green circles) and nesting sites (red circles).

Figures 2–3. Dorsal habitus of females. 2. Colletes validus Cresson. 3. C. ultravalidus, new spe- cies. Note difference in size, hair color on body and legs, infuscated wings, and metasomal tergal fascia. Scale bars = 3 mm.

Figures 4–5. Face of females. 4. Colletes validus Cresson. 5. C. ultravalidus, new species. Scale bars = 1 mm.

Figures 6–7. Lateral view of female heads. 6. Colletes validus Cresson. 7. C. ultravalidus, new species. Scale bars = 1 mm.

Figures 8–9. Face of males. 8. Colletes validus Cresson. 9. C. ultravalidus, new species. Scale bars = 1 mm.

Figures 10–11. Lateral view of male heads. 10. Colletes validus Cresson. 11. C. ultravalidus, new species. Scale bars = 1 mm.

Figures 12–13. Metasomal sternum seven. 12. Colletes validus Cresson, dorsal view. 13. C. ultravalidus, new species, ventral view. Scale bars = 0.5 mm.

Figures 14–15. Genital armature. 14. Colletes validus Cresson. 15. C. ultravalidus, new species. Arrows point to dorsal wings of penis valves, slightly more distal in C. ultravalidus. Scale bars = 1 mm.