J. HYM. RES. Vol. 13(2), 2004, pp. 250-261

Hooked Hairs and Not So Narrow Tubes: Two New Species of Latreille from Texas (: Apoidea: )

John L. Neff

Central Texas Melittological Institute, 7307 Running Rope, Austin, Texas 78731; email: [email protected]

Abstract. —Two new species, Colletes bumeliae Neff and Colletes inuncantipedis Neff, are de- scribed from central Texas, U.S.A. Both have arrays of specialized, recurved setae on their foretarsi which are used to extract pollen from flowers of lanuginosum (A. Michaux) Pennington (), their primary floral host. Nests and provisioning behavior are described for C. in- uncantipedis.

Colletes is a nearly cosmopolitan genus pers. com.). In addition, females of some of consisting of more than 440 species Mesoamerican Colletes (including C. isth- (Michener 2000, Kuhlmann 2003). Rough- micus Swenk and C. mexicanus Cresson) ly 100 species have been recorded from have a metasomal scopa comprised of se- North and Central America, making Col- tae with hooked apices (Griswold et al. letes one of the larger genera of American 1995). I here add to this short list by de- bees (Michener et al. 1994). While many scribing two distinctive new species from species of Colletes are believed to be oli- central Texas. Females of both species golectic (Stephen 1954, Moldenke 1979, have unusual arrays of hooked setae on Mader 1999, Muller and Kuhlmann 2003) their foretarsi which are employed in the very little has been reported on either collection of pollen from the partially hid- den anthers of flowers of Lin- their pollen collecting behavior or any Sideroxylon morphological features which may en- naeus (Sapotaceae). hance pollen harvesting by these bees. This is MATERIALS not particularly surprising since AND METHODS most North American Colletes lack obvi- Morphological nomenclature follows ous specializations for pollen harvesting Michener (2000) with the addition of the beyond the usual branched body hairs terms clypeal apicomedial and apicolater- and tarsal and tibial of brushes simple se- al fovea for the sensillae bearing fovea on tae (Thorp 1979). The few exceptions in- the apical margin of the clypeus. Defini- clude the sternal setal brushes of females tions for abbreviations and measurements of some members of the consors of group are as follows: UIOD—upper interocular used for Stephen (1954) pollen collection distance (minimal distance between com- during vibratile pollen from harvesting pound eyes on upper part of face); the flowers of Chamaesaracha, and — Quincula LIOD lower interocular distance (mini- (all Physalis Solanaceae) 1954, J. mal (Stephen distance between compound eyes on L. Neff pers. obs.), and of hooked arrays lower portion of face); MIOD—maximal setae on the and frons of an clypeus un- interocular distance (maximal distance be- described Mexican that aid in col- tween species compound eyes); OCD—ocellar di- from the nototribic anthers lecting pollen ameter; PW—puncture width; Tl, T2, T3 of flowers of the Lamiaceae (G. Dieringer, -Metasomal Tergum 1, Tergum 2, Ter- Volume 13, Number 2, 2004 251

.. — 3 . S3 ... Metasomal Ster- C. C. simulans gum ; SI, S2, scopiventer Swenk, Cresson,

. . . — num 1, Sternum 2, Sternum 3 ; BL C. skinneri Viereck, C. sleveni Cockerell, C. body length (length front of vertex to pos- sivenki Stephen, C. texanus Cresson, C. terior margin of T2); FW—forewing length thoracicus Smith, C. willistoni Robertson, C. (measured from base of the—radial cell to wihnattae Cockerell and C. wootoni Cock- apex of marginal cell); HW head width erell. (maximal width in frontal view). Acro- — Colletes bumeliae Neff, new species nyms used include: BFL Brackenridge — Field Laboratory of The University of Tex- Diagnosis. The female of Colletes bume- as, Austin, Texas; SEMC—Natural History liae is easily distinguished from all other Museum, University of Kansas, Lawrence, Nearctic Colletes, except C. inuncantipedis, Kansas; TAMU—Entomology Collection by the distinctive arrays of hooked hairs of Texas A & M University, College Sta- on its forebasitarsi. It fails at couplet 71 in tion, Texas and J.L.N.—J.L.Neff. the female key of Stephen (1954) due to The lengths of the foretarsi (and femora the absence of dark hairs on the meso- and tibiae) of female Colletes bumeliae and scutum and scutellum; mesepisternum C. inuncantipedis were compared with strongly punctate, not rugose; and absence those of females lacking the specialized of basal fascia on T2. The male runs to Col- hooked hairs. Lengths were adjusted for letes kansensis Stephen (couplet 71 in Ste- size by dividing the appropriate leg length phen's key) but is readily distinguished by by head width. Adjusted tarsal and other the shape of the 7th sternite, the puncta- leg length measures did not differ be- tion of Tl fine and sparse medially, and tween C. bumeliae and C. inuncantipedis so the tegulae translucent brown, not black. they were combined for further analyses. Colletes bumeliae is clearly closely related to For comparative purposes, I also used two C. inuncantipedis. The form of the genitalia measures of forewing length as the divi- and the metapleural prominence suggest sor: as defined above and, in unworn kansensis may be the closest relative to specimens, from the base of the radial cell these two species. Unfortunately, the fe- to the distal wing apex). Seventy-five in- male of kansensis is unknown. — = dividuals from 39 species (two or rarely Description. 9. Measurements: (N one female per species), were used for the 9) BL: 7.0-8.2 mm; FW: 4.5-5.2 mm; HW analysis (11 females were excluded due to 2.8-3.1 mm. Head: Face 1.24-1.29 X wider extensive wing wear for the second wing than long, greatest distance between eyes length analysis). Colletes species included exceeding eye length, inner orbits strongly in the tarsal length analysis are: C. aesti- convergent below and arched inward valis Patton, C. algarobiae Cockerell, C. az- above (UIOD 1.11-1.15 X LIOD, UIOD tekus Cresson, C. aridus Stephen, C. bea- 0.81-0.89 X MIOD). Malar space about Va merorum Stephen, C. birkmanni Swenk, C. as long as wide. Clypeus slightly pro- brevicomis Robertson, C. cercidii Timber- duced with depressed, flattened, impunc- lake, C. clypeonitens Swenk, C. compactus tate, apical margin; clypeal disk shining, Cresson, C. delicatus Metz, C. deserticola densely, striately punctate, often with Timberlake, C. eulophi Robertson, C. fulgi- small, shining, impunctate medioapical dus Swenk, C. gilensis Cockerell, C. gypsi- area; punctures smaller, denser and non- colens Cockerell, C. hyalinus Provancher, C. striate basally and laterally. Clypeal api- intennixtus Swenk, C. louisae Cockerell, C. comedial fovea large and round, apicola- latitarsis Robertson, C. mandibularis Smith, teral fovea smaller and much weaker. Disc C. mitclielli Stephen, C. nigrifrons Titus, C. of supraclypeal area raised, surface dull paniscus Viereck, C. perileucus Cockerell, C. and microstriate with sparse, moderate, prosopidis Cockerell, C. saritensis Stephen, punctures; punctation of lateral surfaces of Hymenoptera Research 252 Journal smaller and much denser. Median line punctures; interspaces shining; hypoepi- meral area Me- carinate from above supraclypeal area to shining, sparsely punctate. median ocellus. Disc of labrum rounded, tepisternum shining, quadrately pitted; median small, rugose, shining, impunctate, without metapleural prominence with short, curved, carinate rim groove. Frons with punctures strong, opaque, above small dense and nearly contiguous. Facial fovea shining declivity. Propodeum but with dorsal area pit- deeply impressed, broadest medially shining, quadrately surface of trian- restricted above, curving inward towards ted; posterior propodeal surface lateral ocellus; upper margin of fovea at gle shining; posterior propodeal summits of outside pos- or slightly above line between triangle coarsely roughened, of fovea nar- terolateral of weakly eyes, upper inner margin margins propodeum subrectan- rowed, within 1 OCD of lateral ocellus, fo- carinate. Fore basitarsis broad, width 0.37 X fore bas- vea tapering below, extending to level just gular (basal length, X fore hind bas- above upper margin of antennal fossae; itarsal length 0.58 tibia); as broad as surface dull, microstriate. Vertex shining, itarsis broad, roughly 3x long, sides Abdomen: shin- densely, minutely punctured except shin- subparallel. Terga with narrow ing impunctate area between lateral ocelli ing, impunctate margins; and facial fovea. Gena narrow, broadest punctation of Tl fine and dense laterally in below; genal width 0.5 X eye width at lev- but much sparser, nearly impunctate medial lA to Y discal of distal el of antennal insertion; punctures dense 5; punctation Puncta- above becoming striate below and on hy- terga uniformly fine and dense. postomal area. Scape slender, length 5 X tion of S 1-2(3) moderately dense, slightly first coarser than finer on dis- apical width. Minimal length flagellar terga, becoming of sterna with segment short, minimal length slightly tal sterna, apical margins translucent. of S6 less than apical breadth. Middle antennal narrowly Apex slightly surface with of segments slightly longer than broad. Tho- depressed, shining density Ves- rax: Pronotal spine short but sharp; broad punctures decreasing towards apex. titnre: Pile of face dense and basally, abruptly narrowing to acute tip. white, par- frons but Tegulae sparsely punctate, most punc- tially obscuring surface on x tures in apical h. Mesoscutum shining, sparse on vertex, clypeus and supracly- disc strongly punctate with punctures sep- peal area; hairs of clypeus, short, sparse, arated 1-1.5 pw on anterior %, punctures simple and semi-appressed; 1-2 long, much sparser on posterior Vz, punctures bent, flattened simple setae in apicomedial finer and denser on anterior and posterior fovea of clypeus, 3-4 shorter branched se- mesoscutal margins. Scutellum shining, tae in apicolateral fovea. Pile of vertex punctures strong, similar to discal meso- pale yellowish-white, hairs branched and scutal punctures on posterior %, punctures relatively dense in and behind ocellar tri- smaller and denser on posterior margin angle but sparse and simple in ocello-oc- but very sparse, nearly impunctate on an- ular space; hair of upper part of gena terior third. Metanotum rugose. Mesepis- dense with numerous short branched ternum anterior to quadrately pitted epi- hairs, appressed along upper posterior sternal groove densely punctate; punc- margin of eye, becoming much sparser tures slightly coarser than those of meso- and less branched below. Pile of meso- less than 1 scutum, punctures pw apart. scutum and scutellum pale, yellowish- Mesepisternum posterior to groove with white, hairs short with numerous long in punctures decreasing size and density branches; dense fringe of white, branched posteriorly (except punctures dense along hairs on posterior margin of pronotal lobe, with meso-metepisternal suture) posterior pile of mesepisterna, erect and sparse, not most punctures

carinate. Fore S2 weaker on dis- gins of propodeum weakly becoming progressively ex- tal sterna. Color: Black; antennae black to trochanter simple; fore femur slightly X as dark brown; mandibles with % panded mediodorsally, 3.6 as long apical reddish as in fe- broad; fore tibia 4.2 X as long as broad; translucent brown; wings dark brown with tarsi hind tibia not expanded, 5.0 X as long as male; legs ferrugi- 4.5 times as nous, tibial brown, broad; posterior basitarsi long spurs pale Tl shin- Material examined.— 9 : USA: as broad, sides parallel. Abdomen: Holotype fine Texas: Co: (30° 12.99' ing- lateral and posterior punctures Bastrop Sayersville L. Neff; on and dense but well separated on disc: N, 97° 20.99' W); ll-VI-1991; J. sometimes with median impunctate line, flowers of Sideroxylon lanuginosum (depos- ited 6: same data as ho- apical margin depressed, apex narrowly SEMC). Allotype Para 18 translucent. T2-6 with surface shining, lotype (deposited SEMC). types: 1 and 2 9 same data as 1 punctures fine and dense, roughly pw 66 9, holotype: 1 translucent. 17 6 and 1 9 , same data 9-VI-1986; apart, margins narrowly except dense- 9 same data 10-VI-1990 and taken strongly depressed, surface dull and , except with in 9 same data 3-VI-1992 ly punctate. Sterna weakly shining, nest; mm, except of fine reticulate shagreening, fine punctures and at flowers Sideroxylon lanuginosum; data 2 6 1-2 pw apart throughout except on nar- 3 6 6, same except 6-VI-1994; 6, 2 same row, impunctate translucent margins; gen- same data except 17-VI-1994; 9 9, 1 9 Swift italia, S7 and S8 as illustrated (Figs. 3a-d); data except 30-VI-1997; , Camp A. penis valves without dorsal wing; distal Military Training Area, 12-VI-2003, 2 Pedernales Falls processes of S7 short and membranous. Hook. Blanco Co.: 9 9, 25- Vestiture: Pile of face long, pale yellowish State Park (30°19.94' N, 98°15.37' W), L. in 2 same white, concealing facial surface except apex VI-1988, J. Neff, nest; 6 6, of clypeus exposed. Pile of vertex pale data except 2-VII-1988 and at flowers of data ochraceous, much sparser, fine hairs in Sideroxylon lanuginosum; 1 male same ocello-ocular areas with denser, conspicu- except 27-VI-1997. ously branched hairs in and behind ocellar Etymology. —From the sapotaceous ge- triangle. Pile of gena pale yellowish white nus Bumelia Swartz, a junior synonym of near vertex, becoming whiter and longer Sideroxylon (Pennington 1990), the appar- below and on hypostomal area. Pile of me- ent sole pollen host of the species. I find soscutum and scutellum pale ochraceous. bumeliae to be more mellifluous than Pile of sides of thorax sparse, pale yellow- names based on Sideroxylon. In addition, ish white with dense dorsolateral propo- recent molecular studies suggest the res- deal fringe pale ochraceous above. Hair of urrection of Bumelia for the American spe- Tl with Sid- legs yellowish-white. long pale cies of Sideroxylon may be justified as yellowish-white hair, dense laterally but eroxylon appears to be paraphyletic (An- very sparse medially, not extending to derberg and Swenson 2003). complete, white, apical fascia. T2-5 with Colletes Neff, complete, white, fascia, fascia inuncantipedis apical apical new absent on T6; T2-5 with hairs of disc very species short, black, with increasing mix of longer, Diagnosis. —The female of Colletes inun- hairs pale posteriorly, lateral portions of cantipedis is distinguished from all other T2-5 white. T6-7 with short, sparse, pale, American Colletes (except C. bumeliae) by hairs. SI with appressed hairs white, the distinctive arrays of hooked setae on S2-6 with sparse; pile white, that on discs its foretarsi. Females can be distinguished and semi-appressed; apices from C. bumeliae by their smaller size, tinct apical fascia of ap- presence of dark hair on the vertex and branched ->sed, hairs, fascia strongest on mesoscutum, and more complete discal Volume 13, Number 2, 2004 255 punctation of Tl. The males of C. inuncan- dark hairs, veins dark brown with pter- tipedis are very similar to males of C. bu- ostigma dark brown. = meliae but are slightly smaller and differ in 6 .—Measurements: (N 7), BL: 6.3-7.4 the shape of S7 and the well-formed pro- mm; forewing 4.3-5.3 mm, HW 2.6-2.9 podeal spine (rudimentary in C. bumeliae). mm. Head: Face 1.20-1.24 X wider than — = Description. 9. Measurements: (N long, greatest distance between eyes 10) BL: 6.6-7.8 mm; FW: 4.5-4.9 mm; HW slightly greater than eye width; inner mar- 2.8-3.0 mm. Head: Face 1.20-1.29 X wider gins strongly convergent below and weak- than long, greatest distance between eyes ly incurved above, UIOD distance 1.31— exceeding eye length, inner orbits conver- 1.41 X LIOD, UIOD 0.91-0.94 x MIOD. gent below and arched inward above Malar space 0.4 times as long as wide. (UIOD 1.11-1.21 LIOD, UIOD 0.80 X 0.86 Clypeus densely, finely, punctate, discal MIOD). Malar and clypeus as in C. bume- punctures slightly sparser apically, apical liae except clypeal striae irregularly con- margin narrowly depressed. Labrum with vergent in medioapical impunctate area. disc evenly rounded, impunctate and Disc of supraclypeal area raised, shining, shining. Supraclypeal area densely punc- microstriate with strong, well-separated tate, medial punctures roughly twice di- punctures; punctation of lateral surfaces ameter of peripheral punctures. Median smaller and much denser. Median line line carinate from apex of supraclypeal carinate from above supraclypeal area to area to just below median ocellus. Punc- preocellar triangle. Labrum, frons, vertex tures of frons coarse and dense. Facial fo- and facial fovea as in C. bumeliae. Gena as vea well defined, narrow, expanded sub- in C. bumeliae but slightly wider, width apically, upper portions of facial fovea approx. 0.6 X eye width at level of anten- strongly depressed along edge of eyes, nal insertion. Flagellum as in C. bumeliae. outer margins carinate, surface of fovea Thorax: Thorax and legs as in C. bumeliae dull. Punctures of vertex fine, dense with except forebasitarsis slightly longer and impunctate space laterad of ocelli. Gena narrower; basal width 0.33 X length; shining, finely punctate, distinctly de- length 0.60 X length of foretibia. Abdo- pressed along eye margin medially, nar- men: Terga as in C. bumeliae except discal row above but considerably wider below. punctation stronger, more uniform, with- Scape strongly punctate, short, approxi- out median impunctate area. Punctation mately 2x longer than apical width; first of sterna similar to C. bumeliae but coarser, flagellar segment short, length 0.9 X apical particularly on S6 where apical punctures width; middle flagellar segments 1.5-1.6 are strong and dense. Vestiture: Pile of X as long as wide. Thorax: Prothoracic face as in C. bumeliae except vertex with spines short and sharp. Mesoscutum shin- mixture of yellowish-white and darker ing, strongly punctate with punctures sep- brown to black hair. Pile thorax as in C. arated by 1-2 pw anteriorly and laterally, bumeliae except brown to black hair discal punctures 2-3 pw apart; punctures sparsely mixed among pale pile of scu- of scutellum similar, dense laterally and tum. Hair of legs C. bumeliae. Pile of terga posteriorly but very sparse, nearly im- as in C. bumeliae except T-5 without pale punctate anteriorly. Mesepisterum, ante- apical fascia. Pile of sterna darker and rior to quadrately punctate mesepisternal denser, particularly on distal Vi of S6. Col- groove, densely punctate, punctures or. Black, except apical half of mandibles, slightly coarser than those of scutum, tibial spurs, and tarsal claws dark reddish punctures less than 1 pw apart. Punctures brown; apices of T 1-4 and S 1-5 narrowly of mesepisterna posterior to groove simi- translucent brown.. Tegulae dark brown to lar to those of scutum, dense anteriorly black; wings hyaline, with abundant short but sparser posteriorly and ventrally to Hymenoptera Research 256 Journal of

usual of hooked setae on the 2-3 pw apart, interspaces shining. Hypoe- arrangement well foretarsi and is a combination of inuncantis pimeral area shining with strong sep- covered with hooks) and arated punctures. Metepisternum shining, (Latin—— pedis quadrately pitted; metapleural promi- (Latin leg), nence small, rugose, with short, curved carinate rim above small shining declivity. encountered Propodeum as in C. bumeliae. Foretrochan- We have occasionally of Colletes bumeliae at, and ter simple; forefemur slightly expanded males nectaring flowers of tex- mediodorsally, 3.6 X as long as broad; patrolling, the Eysenhardtia and one male bore fore tibia 4.2 X as long as broad; hind tibia ana Scheele (Fabaceae) on its mouth- not expanded, 5.0 X as long as broad; pos- pollinia of Asclepiadaceae C. bumeliae and C. in- terior basitarsi 4 X as long as broad, sides parts, but males of are observed parallel. Abdomen: Tl shining, lateral and uncantipedis normally only punctures fine and dense but about flowers of Sideroxylon lanuginosum posterior = well separated on disc, apical margin de- (A. Michaux) Pennington (Sapotaceae) ( of C. pressed and narrowly translucent. T2-6 Bumelia lanuginosa) (Fig. lb). Females with surface shining, punctures fine and bumeliae and C. inuncantipedis have only dense, roughly 1 pw apart, margins nar- been observed at flowers of S. lanuginos- rowly translucent. T7 strongly depressed, urn, with most females bearing scopal surface dull and densely punctate. Sterna loads or extensive loads of pollen on their weakly shining, with fine reticulate sha- foretarsi. All scopal pollen loads examined = greening, fine punctures 1-2 pw apart consisted solely of Sideroxylon pollen (n throughout except on narrow, impunctate 20). All evidence points to both species be- translucent margins; genitalia and S8 as in ing oligolectic on Sideroxylon. C. bumeliae (Figs. 3a-d), S7 as figured (Fig. Sideroxylon lanuginosum is a widespread 3e) with elongate, strong distal processes tree or shrub (2-15 m tall) of the Southeast (not short and membranous as in C. bu- and South Central U.S.A. and adjacent meliae). Vestiture: as in C. bumeliae. Color: Mexico, commonly known as gum bume- Black; antennae black to dark brown; lia, gum elastic, chittamwood or wooly- mandibles with apical % translucent red- bucket bumelia (Cheatham et al. 2000). dish brown; wings as in female; legs dark Sideroxylon lanuginosum normally flowers brown with tarsi ferruginous, tibial spurs in middle to late June in central Texas, brown. pale producing large numbers of small, pale, Material examined. —Holotype 9. USA: short-lived, nectiferous flowers. The tu- Texas: 0. mi. Bastrop Co., Bastrop, 5 N (30° bular, distally spreading corolla is 3-4 mm 08.11' 97° N, 19.24' W), J.L. Neff, 30-VI- long. Each flower has five fertile anthers, 1997. on flowers of Sideroxylon lanuginos- each of which is subtended by a petaloid urn (deposited SEMC). 6 : same data as ho- staminode. The anthers are not included lotype (deposited SEMC). Paratypes: 3 9 5 in the basal floral tube but rather are par- and 19 8 same data as 6, holotype. 13 9 9, tially hidden in the distal folds of the co- same data as holotype except 17-VI-1998. rolla (Fig. lb). I have observed female C. Travis Co.: 1 BFL 6, Austin, (30° 17.10' N, inuncantipedis first inserting their heads 97° 46.83' L. W), 6-V1-1986, J. Neff, on into the corolla, apparently to lap nectar, flowers of Eysenhardtia texana; 1 6, same and then inserting their forelegs into the data 12-VI-1987 and on flowers of except corolla, apparently to gather pollen (Fig. Sideroxylon lanuginosum; 2 8 6, same data la). The hooked foretarsal hairs of female 18-VI-l 1 same data except ?87; 8, except C. bumeliae and C. inuncantipedis often bear much Sideroxylon pollen and aid ' obviously The #!/.— name refers to the un- in extracting pollen from the partially hid- Volume 13, Number 2, 2004 257

Fig. 1. Flowers of Sideroxyhm lanuginosum. a. Pollen-collecting female of Colletes inuncantipedis inserting fore- legs into corolla, b. Sideroxi/lon inflorescence.

= = den anthers. The exact mechanics of how (n 18) vs. 0.511 ± .034, n 75, P < C. bumeliae and C. inuncantipedis extract .0001, unpaired t-test). pollen of S. lanuginosum has yet to be de- The foretarsal arrays of hooked setae of termined because most pollen collecting females of Colletes bumeliae and C. inuncan- visits are relatively rapid (< 5 sec), and tipedis are apparently unique among Ne- occur high in the canopy. The foretarsi, arctic and Neotropical Colletes species with their arrays of hooked hairs, appar- (Figs. 2a, b). The foretarsi of most New ently are pulled over the anthers which World Colletes I have examined only bear fe- are partly hidden by the lateral folds of simple hairs (Figs. 2e, f), although the the corolla and the subtending stami- males of few species, such as C. skinneri or nodes. This pollen collecting behavior ap- C. wootoni, have foretarsal combs with pears to be analogous to that of various dense arrays of apically hooked hairs Calliopsis (Verbenapis) spp. which employ (Figs. 2c, d). The function of the hooked hooked tarsal hairs to extract pollen from hairs for these bees is unclear as neither is anthers hidden in the narrow corollas of closely associated with tubular flowers. various Verbenaceae (Shinn 1967), with Colletes zvootoni apparently is polylectic the obvious difference that the corollas of while C. skinneri appears to be oligolectic L. Sideroxylon are not particularly narrow on papilionoid legumes (J. N. pers. and the anthers are much more exposed. obs.). While the adjusted combined length of The foretarsal combs of C. bumeliae and the femur and tibia of the forelegs does C. inuncantipedis are remarkably similar to not differ between C. bumeliae and C. in- the those found on the foretarsi of the fe- uncantipedis and a sample of North Amer- males of the west Palearctic species, C. na- = ican Colletes (0.871 ± .032 (n 19) vs. sutus Smith. Colletes nasutus is oligolectic = = 0.881 ± .044 (n 75), p .3408, unpaired on Anchusa (Boraginaceae) and uses its t-test), the adjusted foretarsal lengths of C. foretarsal arrays to extract pollen from the bumeliae and C. inuncantipedis are signifi- anthers included in the narrow corolla cantly shorter than average of the sample tubes (Muller 1995). Females of C. nasutus of North American Colletes, (0.490 ± .011, differ from C. bumeliae and C. inuncanti- Journal of Hymenoptera Research 258

foretarsal setae, b. Colletes d. Colletes Fig. 2. Foretarsi of female Colletes with closeup of a, inuncantipedis. c,

wootoni. e, f. Colletes birkmanni.

Nos- pedis in having foretarsi of normal length tae. These two species, C. anchusae are oli- but elongate forefemora and -tibiae, ap- kiewiscz and C. wolfi Kuhlmann, parently an adaptation for extracting pol- golectic on Cynoglottis (Boraginaceae) and len hidden in narrow corolla tubes (Muller use their tarsal setal arrays to extract pol- 1995). Shortened foretarsi (and modified len from the narrow floral tubes of Cynog- forefemora) are found in two closely re- lottis flowers (Muller and Kuhlmann lated European Colletes whose foretarsi 2003). bare array 1 of stout, flattened, curved se- Muller (1995) and Muller and Kuhl- Volume 13, Number 2, 2004 259

Fig. 3. Genitalia and associated sterna, a. dorsal view, genital capsule, Colletes bumeliae. b. ventral view, genital capsule, C. bumeliae. c. S8, dorsal view, C. bumeliae. d. S7, dorsal view, C. bumeliae e. S7, dorsal viev\ , C. inuncantipedis.

mann (2003) used forewing length as their wing length is expected to be an acceler- measure of body size, rather than head- ating, positive function of body size due width. Curiously, when either internal or to considerations of wing loading (Dan- total wing length was used in the denom- forth 1989). In addition, the use of internal inator when adjusting for body size in the markers of wing venation to avoid prob- comparison of American bees, no signifi- lems of wing wear in estimating wing cant difference was found between the tar- length may further complicate matters sal lengths of females with and without since wing venation extends distally with hooked hairs (0.306 ± 0.007 (n 19) vs. increasing body size (Danforth 1989). = = 0.302 ± 0.027 (n 75), p 0.5488 or 0.255 Head width was more strongly correlated = = ± 0.008 (n 18) vs 0.254 ± 0.022, (n with transtegular distance (another mea- = 65), P 0.8390). However, there are sev- sure of body size), than was wing length = = eral reasons why wing length may be a (R .945 vs. R .894) in the sample of less than ideal estimator of body size in American Colletes, suggesting it should be many groups. First, wing length is of- a superior estimator of body size, at least ten compromised by wing wear. Second, among groups like Colletes which lack ob- 260 Journal of Hymenoptera Research

Pollen in the vious cephalic allometry. Ultimately, these entrance. semi-liquid provi- estimators will need to be tested against sions was 100% Sideroxylon. between actual body dry weight. Most pollen foraging occurs Nests of C. inuncantipedis are unknown 0900 and 1400 CDST, with some foraging but nests of C. bumeliae were discovered in as late as 1630 hrs. Full daily provisioning deep sandy soils near Sayersville, Bastrop series for three females indicated they Co., Texas, as well as in sandy alluvial de- made 11-12 pollen trips per day. Mean the River at Ped- 6.7 posits along Pedernales pollen trip duration was 19.3 ± min (n = ernales Falls State Park, Blanco Co., Texas. 73, 9-38). Time in the nest between pol- near were = Nests Sayersville loosely len trips averaged 4.9 ± 2.5 min (n 72, grouped along a heavily shaded, unpaved 2-21). a woodland. At road through post oak Since C. bumeliae and C. inuncantipedis Pedernales nests were scattered Falls, are often locally abundant and contact a road cut the alluvial de- along through with the anthers and stigma of S. lanugi- with one of 5-7 nests clus- posits group nosum by both males and females is un- tered within the entrance of a aban- large, avoidable during foraging, these bees po- doned mammal (armadillo?) burrow. Two tentially are important pollinators of S. nests were excavated, one each at Peder- lanuginosum. However, as they are small nales Falls and one at Sayersville, both bees visiting a large tree, actual pollinator with similar structure. The excavated nest efficacy needs to be demonstrated directly, at Sayersville, and other nearby nests, or at least inferred from degree of pollen were on level ground and had a simple carryover and frequency of interplant fan like tumulus formed of soil pushed moves in order to have any confidence in away from the entrance. Entrances to the statements on its importance as a polli- nests at Pedernales Falls were in a near nator. The overall importance of these vertical bank and thus lacked tumuli. The bees in the reproductive biology of S. lan- first 15-20 cm of the main burrows de- uginosum is probably not great since they scended gradually to a depth of 5-10 cm appear to be much more restricted in dis- before descending almost vertically. Both tribution than the tree. Even in central nests were lost before reaching any cells, Texas, the most common visitors of the Sayersville nest at 40 cm and the Ped- gum bumelia are often various ernales Falls nest at 28 cm. In both cases, wasps, particu- the fast males of the problem was backfilled laterals. Bur- larly flying Myzinum spp. as well as males of various rows were unlined and had an interior di- (Tiphiidae), halictid and bee rather ameter of 5.0-5.5 mm. In both cases, the megachilid species, than C. bumeliae or C. burrows each had several soil septa, each inuncantipedis. Colletes 5-10 mm thick. No cells were recovered at bumeliae and C. inuncantipedis are known from five sites Sayersville but eight single cells were re- currently only covered from the Pedernales Falls exca- in three counties in central Texas. Al- vation at depths of 30-50 cm. All cells though their ranges overlap, they have not were horizontally oriented and from 10 to been found together at the same site. In all 15 cm from the estimated location of the cases, the sites had both S. lanuginosum main burrow. Individual cells were of the and some nearby areas of sandy soils. The classic of membranous sandwich bag type ranges these two species are likely to with a closure folding (Torchio et al. 1988). expand with further collecting since the One cell recovered intact was 11 mm long fauna of Sideroxylon flowers is very poorly with a maximal diameter of 7 mm. Each known. Despite the wide distribution of cell had a 5 collar, roughly mm in diam- the genus, there are no records of any hy- which extended 3 mm from the cell menopterous visitors to either Sideroxulon Volume 13, Number 2, 2004 261 or Bumelia in the Hymenoptera Catalog uni-muensterde/Landschaftsoekologie/Institut/ Mitarbeiter/Kuhlmann/Coll-check/Index.htm (Krombein et al. 1979). Mader, D. 1999. Geologische und biologische Entomobk- ACKNOWLEDGMENTS ologie der rezenten Seidenbiene Colletes: Band I. Lo- gabook; Koln, Germany, xliii + 807 pp. I thank Beryl Simpson for improving the manu- Michener, C. D. 2000. The Bees of the World. The Johns script and many other things; the Texas Parks and Hopkins University Press, Baltimore. Maryland; Wildlife Department for permission for studies at xiv + 913 pp. Falls State C. R. Pedernales Park; Larry Gilbert for permis- Michener, D., J. McGinley, and B. N. Danforth. sion for studies at BFL, Austin, Texas; Terry Griswold 1994. The Bee Genera of North and Central America for a useful reference and other correspondence; John (Hymenoptera: Apoidea). Smithsonian Institution Mendenhall for assistance with SEM; John Abbott Press, Washington, D.C. viii + 209 pp. and Al Hook for the loan of material from the Ento- Moldenke, A. R. 1979. Host- coevolution and the mology Collection at BFL; Rob Brooks for access to diversity of bees in relation to the flora of North specimens at SEMC; and Ed Riley for access to ma- America. Phytologia 43: 357^19. terial at TAMIL Michael Engel and an anonymous Miiller, A. 1995. Morphological specializations in reviewer found many glitches in the original manu- Central European bees for the uptake of pollen script. from flowers with anthers hidden in narrow co- rolla tubes (Hymenoptera: Apoidea). Entomologia LITERATURE CITED Generalis 20: 43-57. Miiller, A., and M. Kuhlmann. 2003. Narrow flower A. A., and U. Swenson. 2003. Evolution- Anderberg, specialization in two European bee species of the in A cladistic ary lineages Sapotaceae (): genus Colletes (Hymenoptera: Colletidae). Euro- based on ndhF data. Interna- analysis sequence pean Journal of Entomology 100: 631-635. tional Journal Plant Sciences 164: 763-773. of Pennington, T. D. 1990. Sapotaceae. Flora Neotropica Cheatham, S., M. C. Johnston, and L. Marshall. 2000. 52: 1-771. The Wild the Useful of Texas, Southeastern Shinn, A. F. 1967. A revision of the bee genus Calli- United States, the Southern Plains, and Northern opsis and the biology and ecology of C. andreni- Mexico. Volume 2. Useful Wild Aus- Plants, Inc.; formis (Hymenoptera, Andrenidae). University of tin, Texas; xxiii + 599 pp. Kansas Science Bulletin 46: 753-936. B. N. 1989. The evolution of Danforth, hymenopteran Stephen, W. P. 1954. A revision of the bee genus Col- The of size. wings: importance Journal of Zoology, letes in America north of Mexico (Hymenoptera: London 218: 247-276. Colletidae). University of Kansas Science Bulletin. Griswold, T., F. D. Parker and P. E. Hanson. 1995. 36: 149-527. The bees 650-691. In: P. E. (Apidae). Pp. Hanson Thorp, R. W. 1979. Structural, behavioral and physi- and I. D. eds. Gauld, The Hymenoptera of Costa ological adaptations of bees (Apoidea) for col- Rica. Oxford University Press, Oxford, England. lecting pollen. Annals of the Missouri Botanical

Krombein, K. V., P. D. Hurd, Jr. and D. R. Smith. Garden 66: 788-812. 1979. in Catalog of Hymenoptera America North of Torchio, P. F., G. E. Trostle and D. J. Burdick. 1988. Mexico. Smithsonian Institution Press. Washing- The nesting biology of Colletes kincaidii Cockerell ton, D.C. xxx + 2735 pp. (Hymenoptera: Colletidae) and development of Kuhlmann, M. 2003. Checklist of the Old World its immature forms. Annals of the Entomological Species of the Bee Genus Colletes Latr. www. Society of America 81: 605-625.