Boletes of Eastern North America

30 · Boletes of Eastern North America 3a. Cap at first reddish orange, rose red, rose pink, pinkish red, or pinkish purple → 4 (see also Boletus paluster [p. 132]) 3b. Cap at first pinkish cinnamon, dull brick red, brown, olive brown, reddish brown, yellow, or yellow brown → 8 4a. Stalk reticulation prominent, typically covering upper half or more of the stalk → 5 4b. Stalk reticulation or pseudoreticulation delicate to obscure or present only at the stalk apex → 13 5a. Flesh staining quickly and decisively blue when exposed → 6 5b. Flesh unchanging or changing only slightly and slowly blue when exposed → 7 6a. Flesh taste bitter to astringent, sour or acidic (like lemon) → Butyriboletus roseopurpureus (p. 184), cap pinkish purple when fresh, becoming darker purplish pink to dark purplish red, at times mottled in these colors, marginal areas becoming grayish with age (compare with Butyriboletus peckii [p. 182]) Alessioporus rubriflavus (p. 56), cap dark wine red at first, soon predominantly a yellow ground color with streaks and splashes of wine red, red brown, or ocher, developing olive tones on the disc in age 6b. Flesh taste not distinctive; cap bright rose red to rose pink, becoming pinkish brown to olive brown with age → Boletus speciosus (p. 155) 7a. Tubes and pores staining bluish green when cut or bruised; cap dull red at first, becoming pinkish red to orange red and fading to reddish orange to dull golden orange in age; distribution in the southeastern United States → Boletus carminiporus (p. 97) 7b. Tubes and pores staining blue and then brownish when cut or bruised; cap deep red to dull red or rose red, fading to brownish rose or pale tan to buffy brown from the disc outward; known distribution eastern Canada south to Georgia, west to Tennessee and Indiana → Butyriboletus peckii (p. 182) 8a. Cap and stalk bright yellow when young, developing reddish-brown tones with age; all parts instantly staining blue when handled or bruised → Neoboletus pseudosulphureus (p. 279) 8b. Cap pinkish cinnamon, dull brick red, brown, olive brown, reddish brown, grayish brown, or yellow brown → 9 9a. Stalk reticulation or pseudoreticulation prominent, extending well below the stalk apex → 10 9b. Stalk reticulation weak to uncertain or limited to the stalk apex only → 14 10a. Stalk reticulation brownish, especially at the central portion of the stalk, possibly yellowish at the apex; cap yellow brown to amber brown, reddish brown, or streaked dull brown over a yellow ground color, becoming ochraceous tawny to hazel with age → Boletus luridellus (p. 114)

Field Keys to the Boletes · 31 (compare with Buchwaldoboletus lignicola [p. 177], Boletus ferrugineus [p. 105], and Xerocomus illudens/tenax complex [p. 426]) 10b. Stalk reticulation whitish to pinkish or yellow, sometimes brown → 11 11a. Cap dark chocolate brown to dark purplish brown; stalk entirely red or reddish, reticulation yellow; occurs in sandy soil under bear oak and pitch pine; known only from Cape Cod, Massachusetts, rare → Boletus billieae (p. 95) 11b. Cap dull brick red, pinkish cinnamon, olive brown to gray brown, reddish brown, or dark yellow brown; stalk a mix of red and yellow → 12 12a. Cap light brown to grayish brown or olive brown at first, becoming paler olive brown to dark yellow brown with age; distribution in eastern North America known from the upper Midwest and New England, uncommon → Caloboletus calopus (p. 186) 12b. Cap dull brick red, pinkish cinnamon, reddish brown, or yellow brown to olive brown → Butyriboletus brunneus (p. 181), stalk reticulation fine; cap reddish brown or yellow brown to olive brown; pores instantly bruising blue; flesh pale yellow, quickly staining blue when exposed; distribution from eastern Canada to North Carolina, west to Minnesota; fairly common, especially under hemlock Xerocomus illudens/tenax complex (p. 426), stalk reticulation or pseudoreticulation coarse and wide-meshed; pores usually slowly staining cinnamon red or sometimes slowly staining bluish and then cinnamon red, rarely unchanging when bruised; cap yellow brown to pinkish cinnamon or dull brick red to reddish brown; flesh whitish to pale yellow, unchanging or slowly staining pinkish red when exposed; widely distributed in broadleaf woods with oak, in conifer woods with pine, or in mixed woods 13a. Odor of cut flesh pronounced, often described as similar to beef or chicken bouillon, maple syrup, fenugreek, curry, licorice, or fruit → Boletus pallidoroseus (p. 129), flesh slowly stains blue when exposed; cap at first uniformly pale rose pink, pale purplish pink, or mottled with darker-pink tones, golden yellow or paler yellow along the margin, becoming pale orange brownish in age Boletus sensibilis (p. 150), flesh quickly stains blue when exposed; cap dark to pale brick red, fading to dull rose or sometimes dingy cinnamon in age 13b. Odor of cut flesh not distinctive; reticulation weak or indecisive, usually limited to the very apex of the stalk → Boletus miniatopallescens (p. 119), cap dry, smooth, or somewhat velvety, red to brick red, soon fading to reddish orange or orange yellow, often cracked in age (see also Boletus miniato-olivaceus [p. 117])

32 · Boletes of Eastern North America Boletus purpureorubellus (p. 139), cap viscid when wet, shiny when dry, color variable from dark red or purplish to blood red or reddish orange, with yellow on the margin; currently known from Florida, Georgia, and Cape Cod, Massachusetts Boletus rubricitrinus (p. 145), cap dry, glabrous to slightly velvety, color variable from dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown in age, sometimes with yellow tints, usually bruising bluish black; flesh taste acidic Xerocomus sclerotiorum (p. 430), cap dry, dull or somewhat shiny, pinkish red to rose red or purplish red, sometimes with olive tints, becoming dull rose pink to brownish pink in age, slowly staining blackish blue when bruised Boletus oliveisporus (p. 127), cap dry, slightly velvety, often finely cracked in age, reddish orange overall, sometimes paler yellow orange near the margin when young, becoming dark fulvous tinged with bay, or cinnamon brown to dull brown and somewhat shiny in age, instantly staining blue black when bruised Xerocomus illudens/tenax complex (p. 426), cap yellow brown to pinkish cinnamon or dull brick red to reddish brown; pores usually slowly staining cinnamon red or sometimes slowly staining bluish and then cinnamon red when bruised 14a. Cap deeply corrugated or pitted, tan to reddish tan or ocher brown to reddish brown; reticulation delicate (when present) → Xerocomus hortonii (p. 421) 14b. Cap smooth, wavy, or irregular but not deeply corrugated or pitted → 15 15a. Stalk reticulation restricted (when present) to the very apex of the stalk; cap dull yellow brown, ocher brown, or pale cinnamon brown; flesh pale yellow; occurring with hemlock → Boletus huronensis (p. 110) (see also Caloboletus roseipes [p. 192]) 15b. Stalk reticulation (pseudoreticulation) composed of longitudinal lines that sometimes intersect to form a coarse network; flesh whitish; cap olive brown to yellow brown, often cracking with age → Boletus subtomentosus var. subtomentosus (p. 165) (see also Xerocomus illudens/ tenax complex [p. 426]) 16a. Stalk reticulation or pseudoreticulation fluted or shaggy to laceratereticulate, very slender and tall in proportion to cap diameter → 17 16b. Stalk reticulation or pseudoreticulation coarse or fine but not shaggy or lacerate → 18 (see also Suillus cavipes [p. 326], Leccinum rugosiceps [p. 262] and Boletus chippewaensis [p. 98])

Field Keys to the Boletes · 33 17a. Cap smooth, glabrous, and viscid, orange to reddish orange, red brown, or bright yellow → Heimioporus betula (p. 219) 17b. Cap velvety, dry, becoming cracked or scaly with age, yellow brown to reddish brown or cinnamon → Frostiella russellii (p. 207) (see also Boletus ferrugineus [p. 105]) 18a. Cap predominantly yellow to orange, including bright yellow, bright orange, golden orange, honey yellow, grayish, or purplish brown → 19 18b. Cap predominantly brick red, pinkish cinnamon, brown or reddish brown, yellow brown, or chestnut brown → 21 19a. Stalk reticulation prominent, typically over most or all of the stalk; handling the cap and stalk often staining fingers yellow → 20 19b. Stalk reticulation obscure or distinct, not prominent; handling the cap and stalk does not stain fingers yellow; distribution from North Carolina south → Boletus aurantiosplendens (p. 90), cap color variable from orange to brownish orange or brownish yellow, sometimes duller yellowish brown or pale brown to rusty brown with a brighter yellowish orange margin Ceriomyces aureissimus var. castaneus (see photo, p. 92), cap velvety, purplish brown, often duller in age 20a. Stalk rich golden orange, reticulation on young specimens indistinct except at the apex; cap bright orange yellow to golden yellow or brownish orange; flesh taste somewhat acidic or not distinctive → Pulveroboletus auriflammeus (p. 291) 20b. Stalk yellow to yellow brown; cap variable from yellow, mustard yellow, olive yellow to yellow brown or gray; flesh bitter tasting → Retiboletus ornatipes (p. 300) 21a. Flesh pale yellow to bright or dark yellow; cap dull brick red, pinkish cinnamon, yellow brown to chestnut brown, or grayish → 22 21b. Flesh whitish; cap reddish to dull brick red, pale to dark cinnamon brown, or reddish brown → 23 22a. Stalk golden yellow with fine yellow reticulation; cap yellowish brown to chestnut brown or grayish brown, becoming paler in age; flesh bright yellow → Boletus auripes (p. 94) (see also Retiboletus ornatipes [p. 300]) 22b. Stalk duller yellow with longitudinal rib-like lines that form a pseudoreticulum; cap pale brownish yellow when young, becoming yellow brown to pinkish cinnamon; flesh pale yellow → Xerocomus illudens/tenax complex (p. 426) 23a. Cap pale to dark cinnamon brown to dull reddish or dark reddish brown, margin with a prominent overlapping sterile band; stalk colored

34 · Boletes of Eastern North America like the cap, with a conspicuous layer of soft white hairs at the base → Aureoboletus projectellus (p. 64) 23b. Cap dull brick red to reddish brown with an olive tint, margin even; stalk woody at the base, reticulation brown, wide-meshed, basal mycelium yellow → Xerocomus illudens/tenax complex (p. 426) Key B-3 Net Stalks, with Pores Some Shade of Orange, Tan, Buff, Red, Maroon, Brown to Dark Brown, or Nearly Black 1a. Pores radially arranged and more or less decurrent, pinkish brown to golden brown or pale cinnamon brown with pinkish tones; cap brown to reddish brown, burgundy or yellow brown → Bothia castanella (p. 172) 1b. Pores not radially arranged and some shade of orange, red, maroon, or brown to nearly black → 2 2a. Pores brown to maroon or dark brown to nearly black → 3 2b. Pores red to orange → 5 3a. Pores bruising bluish to greenish blue → 4 3b. Pores unchanging or bruising brownish, color grayish orange or darker gray when mature; cap dull reddish brown to brown, dark olive brown, dark gray, brownish gray, or blackish; stalk prominently reticulate nearly overall or at least on the upper one-third → Tylopilus griseocarneus (p. 379) (see also Boletus durhamensis [p. 99], Tylopilus porphyrosporus [p. 389], Tylopilus tabacinus [p. 398], and Bothia castanella [p. 172]) 4a. Pores dark maroon, becoming reddish brown; cap pale brownish olive to yellowish olive; stalk yellow with olivaceous tones at the base, pruinose with reddish pruina that may form a pattern that simulates reticulation on the upper portion → Boletus pseudo-olivaceus (p. 135) 4b. Pores pinkish brown to dark reddish brown to blackish brown; cap dark brown to olive brown or dark vinaceous brown, sometimes staining darker brown when bruised → Tylopilus porphyrosporus (p. 389) 5a. Cap whitish to grayish, grayish brown, or grayish olive; pores pinkish to red or red orange, sometimes yellowish at the margin when young, staining blue when bruised → Caloboletus firmus (p. 189) 5b. Cap red to reddish, brick red, olive yellow, brownish red, brownish orange, or pinkish red → 6 6a. Stalk reticulation coarse and very deep; cap shiny, candy-apple red, often with a narrow, pale margin → Exsudoporus frostii (p. 205) 6b. Stalk reticulation prominent but not coarse or deep, sometimes fine to delicate → 7 7a. Flesh staining bluish when cut → 8

Field Keys to the Boletes · 35 7b. Flesh unchanging when cut; cap dull red at first, becoming pinkish red to orange red and fading to reddish orange to dull golden orange with age; known distribution in southeastern United States from North Carolina south to Florida, west to Arkansas and Louisiana → Boletus carminiporus (p. 97) 8a. Associated with oak or other broadleaf trees; stalk reticulation red → 9 8b. Associated with conifers, especially spruce, hemlock, and pine; cap dark red to brick red, rosy red, or brownish red, sometimes with tan areas near the margin in age → Boletus flammans (p. 107) (see also Suillellus luridus [p. 312]) 9a. Cap predominantly red, reddish, pinkish red, brick red, or purplish red → Exsudoporus floridanus (p. 204), occurs on sandy soil under oaks, distribution from Tennessee and North Carolina south; cap pinkish red, rose red, purplish red, or brownish red, typically yellowish or whitish along the margin; flesh pale to bright yellow, quickly staining dark blue when exposed Rubroboletus rhodosanguineus (p. 304), current known distribution from eastern Canada, western New York, Ohio, and West Virginia; cap dark reddish pink to purplish pink or brick red, usually developing brown to olive tones with age 9b. Cap color variable, from yellow to olive yellow, brownish orange, or olive brown, sometimes with pinkish or reddish tints; cap surface dry, dull or shiny, somewhat velvety → Suillellus luridus (p. 312) Key C Resin Stalks The boletes keyed here have colored resinous dots or irregularly shaped smears on the stalk that are typically tacky or somewhat sticky to the touch. This feature is characteristic of species in the genus Suillus. 1a. Stalk with a membranous or glutinous ring or distinct ring zone → see Key A, Ring Stalks (p. 23) 1b. Stalk lacking a ring or ring zone → 2 2a. Pores staining blue when bruised; cap fibrillose-scaly when young, becoming nearly glabrous in age, fibers and scales grayish, yellow ocher, dark olive yellow, or reddish brown on a yellow to orange-yellow ground color → Suillus tomentosus (p. 360) 2b. Pores unchanging or staining a color other than blue → 3 3a. Associated with broadleaf trees, especially oak or birch; cap with scattered, appressed, brownish to reddish fibrils over an apricot-orange to

36 · Boletes of Eastern North America yellow ground color, viscid beneath the appressed fibrils, somewhat shiny when dry → Suillus subaureus (p. 358) 3b. Associated with conifer trees, nearly always with pines → 4 4a. Pores at first whitish to very pale yellow, grayish, or pinkish buff → 5 4b. Pores yellow, yellowish orange, ochraceous, brown, or grayish olive → 8 5a. Resinous stalk dots and smears pinkish tan to brownish or reddish brown → 6 5b. Resinous stalk dots whitish or yellowish, inconspicuous to lacking in young specimens (use lens) → 7 6a. Cap whitish, becoming yellowish in age; pores whitish at first, soon yellowish to pale ocher yellow in age, often beaded with pinkish droplets when moist; associated with white pine → Suillus placidus (p. 345) 6b. Cap pale yellow, tan, brown, cinnamon, or orangish cinnamon; pore surface often with pinkish-cinnamon moisture beads on young specimens, staining dull cinnamon in age or when bruised or sometimes unchanging → Suillus granulatus (p. 333) 7a. Cap whitish to pale pinkish cinnamon, dull orange yellow, or yellowish tan, becoming pale brown or yellow brown; cap margin with a distinct wide sterile band when young → Suillus neoalbidipes (p. 344) 7b. Cap darker, light to dark brown, vinaceous brown, cinnamon brown, or grayish brown; stalk very short, with inconspicuous resinous dots that become visible only in age → Suillus brevipes (p. 325) (see also Suillus grisellus [p. 338]) 8a. Cap viscid → 9 8b. Cap dry, covered with scattered tufts of reddish, brownish, or grayish fibrils and scales on a yellow ground color, glabrous or nearly so in age; pores pale yellow, sometimes with whitish droplets when young, becoming dull yellow to olive yellow or dingy orange buff in age, staining vinaceous brown when bruised → Suillus hirtellus (p. 340) (see also Suillus subaureus [p. 358]) 9a. Pores bruising vinaceous brown; flesh medium yellow to mustard yellow; cap bright yellow, often with red streaks or patches; associated with white pine → Suillus americanus (p. 319) (see also Suillus hirtellus [p. 340]) 9b. Pores unchanging when bruised; flesh pale yellow; cap pale yellow to ochraceous or rusty orange; odor of flesh fruity or spicy → 10 10a. Pores grayish olive, becoming ochraceous to dull yellow and then olive brown; stalk with minute dots at the apex or glabrous; associated with Scots pine, rare → Suillus bovinus (p. 321) 10b. Pores brown, becoming orange brown to dark dingy yellow; cap viscid, covered with tufts of tiny gray or brown fibrils when young, soon becoming smooth and dull ocher orange, odor spicy; common → Suillus punctipes (p. 346) (see also Suillus subaureus [p. 358])

Field Keys to the Boletes · 37 Key D Rough Stalks, with Scabers or Punctae, or Scurfy Boletes keyed in this group have stalks that are roughened with dry, scale-like projections (scabers) or points (punctations or punctae). There is sometimes a fine line between stalks that are pruinose and stalks that are punctate. If this character is ambiguous, also consider trying the Key E series. Resin Stalks (Key C) differ in having resinous dots or asymmetrical smears on the stalks that are tacky to the touch. If the stalk is lacerated or shaggy, see Frostiella russellii (p. 207). 1a. Scabers, punctae, or scurfy punctae pinkish, red, or yellowish → 2 1b. Scabers, punctae, or scurfy punctae white, buff, pale brown, brown, reddish brown, reddish orange, or black → 8 2a. Pores soon staining when bruised → 3 2b. Pores unchanging, or very slowly staining, when bruised → 7 3a. Pores bruising blue → 4 3b. Pores bruising yellow to brownish or reddish → 5 4a. Pores red to orange Suillellus hypocarycinus (p. 310), cap dry, somewhat velvety, brown to yellow brown, sometimes with olive or cinnamon tones; distribution south from North Carolina to Florida, west to Mississippi Rubroboletus dupainii (p. 302), cap smooth, slimy-viscid when fresh, becoming shiny when dry, purplish red to pinkish red or bright red, sometimes with yellowish spots; rare, reported only from North Carolina and Iowa 4b. Pores yellow; cap color variable, dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown in age → Boletus rubricitrinus (p. 145) (compare with Xerocomellus intermedius [p. 419], Xerocomus sclerotiorum [p. 430], and Hortiboletus rubellus [p. 225]) 5a. Cap viscid to glutinous when moist, shiny when dry, smooth to slightly wrinkled or pitted, yellow orange to brownish orange or ochraceous, often developing green tones in wet weather or with age; stalk punctae yellow to reddish or reddish brown → Boletus longicurvipes (p. 112) 5b. Cap dry or only slightly viscid when moist → 6 6a. Cap dark brown to chestnut brown, smooth to slightly wrinkled or somewhat shallowly pitted; stalk white on the upper portion, pinkish on the lower portion, covered with coarse rosy-pink scabers that darken to brown with age; known only from Florida, Georgia, and Mississippi → Leccinum roseoscabrum (p. 259)

38 · Boletes of Eastern North America 6b. Cap olive brown to reddish brown, often yellow to olive gold on the margin; stalk yellow beneath a covering of reddish or reddish-brown punctae; widely distributed from Georgia northward → Xerocomus morrisii (p. 428) 7a. Pores white at first, becoming pinkish to flesh-colored; cap dry or slightly viscid when moist, pink to rose-colored when young, fading to pinkish tan and then dingy brown with age; flesh white; stalk base deep chrome yellow → Harrya chromapes (p. 217) 7b. Pores yellow at first; flesh yellow; cap furrowed or wrinkled or sometimes smooth, slightly viscid or dry, bay red to chestnut red or reddish brown → Leccinum rubropunctum (p. 261) (see also Hemileccinum subglabripes [p. 220], Leccinum rugosiceps [p. 262], and Xerocomus hortonii [p. 421]) 8a. Pores at first dark chocolate brown to purple brown, becoming reddish brown in age; cap purplish brown to grayish brown; stalk densely covered with darker purple-brown tiny scales → Sutorius eximius (p. 364) 8b. Pores at first white to pale tan, grayish, buff, olive buff, reddish, or yellow → 9 9a. Pores at first yellow to pale yellow, dingy yellow, or reddish → 10 9b. Pores at first white to pale tan, grayish, buff, olive buff, or beige → 13 10a. Pores staining brownish or unchanging when bruised → Leccinellum crocipodium (p. 240), cap dry to moist but not viscid, wrinkled, uneven, pitted, often cracked in age, dark brown to blackish brown when young, fading to pale yellow brown in age; associated with broadleaf trees, especially oak Boletus longicurvipes (p. 112), cap shiny, yellow orange to brownish orange or ochraceous, often developing green tones in wet weather or in age Boletus weberi (p. 171), cap dry, cracked, and scaly, with pale-yellow flesh showing in the cracks, brownish olive at first, becoming olive brown, sometimes with reddish tints; pores unchanging or sometimes slowly staining brown when bruised; distribution southern, from Florida west along the Gulf Coast into Texas 10b. Pores bruising blue to greenish blue or grayish blue (sometimes weakly or slowly) → 11 11a. Cap medium to dark brown, blackish brown, chocolate brown, olive brown, grayish brown, or chestnut brown → 12 11b. Cap brick red to rosy red, yellow orange, or mustard yellow, but always with red or rosy areas → Lanmaoa roseocrispans (p. 235) (see also Boletus patrioticus [p. 133], Hortiboletus campestris [p. 223], and Leccinum rugosiceps [p. 262]) 12a. Stalk conspicuously scaly-shaggy; cap medium brown to dark brown, coated with appressed fibers at first, becoming fibrillose-scaly with

Field Keys to the Boletes · 39 flattened to erect scales and yellow flesh showing between the scales; known only from Florida → Suillellus pictiformis (not illustrated) (p. 78) 12b. Stalk often scurfy to punctate but not scaly-shaggy → Caloboletus roseipes (p. 192), cap olive brown to grayish brown, becoming pale grayish brown to grayish tan with age; associated with conifers in eastern Canada, New England, and New York Boletellus chrysenteroides (p. 78), cap dark brown and velvety when young, becoming bay brown to chestnut brown and often cracked in age, with whitish to pale-yellow flesh that lacks reddish tints showing in the fissures; often growing on stumps or other woody substrates (see also Boletellus pseudochrysenteroides [p. 80], Xerocomellus chrysenteron [p. 416], and Neoboletus luridiformis [p. 277]) 13a. Pores bruising greenish, blue, brownish, yellowish, yellow brown, pinkish gray, cinnamon, or olivaceous → 14 13b. Pores unchanging when bruised → 20 (if ambiguously bruising slowly and weakly buff, see Leccinum scabrum [p. 265]) 14a. Cap at first some shade of orange to orange brown or yellow → 15 14b. Cap lacking orange, orange brown, or yellow tones → 19 15a. Cap pale yellow at first, darkening to yellow and then olive brown with age, typically wrinkled and shallowly pitted at maturity, becoming cracked in age; mycorrhizal only with American hornbeam (Carpinus caroliniana) → Leccinum luteum (p. 256) 15b. Cap orange, dull orange, orange brown → 16 16a. Pores bruising greenish → Leccinellum quercophilum (p. 244), cap glabrous, wrinkled, and pitted when young, becoming conspicuously cracked with age, whitish to grayish flesh showing in the spaces, dull orangish brown to tawny brown when young, becoming dark brown to grayish brown in age; current known distribution from Illinois and Michigan 16b. Pores bruising brownish, olive, olive fuscous, hazel, or cinnamon → 17 17a. Growing in sandy soil or dunes in northern coastal areas with beach grass, beach heather, and sedges; cap orange to yellowish orange, fading to dull cinnamon or ochraceous, wrinkled, becoming cracked in age → Leccinum arenicola (not illustrated) (p. 274) 17b. Growing with conifer or broadleaf trees → 18 18a. Associated with conifer trees, especially Norway spruce → Leccinum vulpinum (p. 274), cap dull orange to brick red, rusty red, reddish brown, or dark reddish brown, with a sterile margin that splits into hanging flaps at least on young specimens 18b. Associated with broadleaf trees → Leccinum insigne (p. 251), cap orange to rusty cinnamon or brick red, becoming brownish orange in age, with a sterile margin that splits

40 · Boletes of Eastern North America into hanging flaps at least on young specimens; flesh stains purplish gray to purplish black when exposed; associated with big-toothed aspen, quaking aspen, or birch Leccinum pseudoinsigne (p. 257), cap bright yellow, orange yellow, or bright orange to orange red, fading to orange buff in age; flesh of young specimens stains reddish cinnamon and then bluish fuscous, but reddish cinnamon stage often lacking on mature specimens; associated with aspen and birch Leccinum versipelle (p. 271), cap dry to slightly viscid, fibrillose, breaking up to form downy patches or small scales with a sterile margin that splits into hanging flaps at least on young specimens, dull orange at first, becoming tan or brownish; flesh staining pinkish and then purple gray to blackish when exposed; associated with birch 19a. Cap brown to dark brown or blackish, often mottled → Leccinum subleucophaeum (p. 269), cap dry or slightly viscid when moist, dark brown with or without paler mottling, often blackish brown over the disc, sometimes fading to yellowish brown in age; flesh slowly staining gray to grayish black; associated with broadleaf trees, especially birch or aspen (if the flesh stains blue, see Tylopilus sordidus [p. 396]) Leccinum snellii (p. 267), cap dark brown to blackish, usually mottled with cream-colored spots; stalk flesh staining orange red at the cap junction, blue green at base; associated with broadleaf trees, especially yellow birch Leccinellum griseum (p. 242), cap wrinkled, uneven, pitted, often cracked in age, dark brown to blackish brown when young, fading to pale yellow brown in age; flesh quickly staining pinkish brown and then darkening to reddish cinnamon; associated with oak Leccinum scabrum (p. 265), cap grayish brown to yellow brown or dark brown, glabrous or matted-fibrillose, breaking up into tiny flattened scales in age; flesh unchanging or slowly staining reddish, especially in the stalk, when exposed; associated with birch 19b. Cap whitish to pale pinkish buff, vinaceous buff, or pale grayish to cinnamon buff → Leccinum holopus (p. 248), cap white or whitish when young, occasionally with gray, buff, tan, or pinkish tints, often darkening and developing a greenish or bluish tinge in age Leccinum insolens (p. 253), cap dry, fibrillose to minutely scaly, sometimes finely cracked, dull white to pale pinkish buff or pale grayish cinnamon buff, typically darkening to pale yellow brown in age; flesh whitish, staining pinkish gray to purple gray and then fuscous when exposed or sometimes lacking the preliminary pinkish coloration before staining purple gray to fuscous

Field Keys to the Boletes · 41 Leccinum chalybaeum (p. 245), cap viscid when fresh, somewhat shiny when dry, smooth to slightly subtomentose, buff to pinkish buff when young, becoming dingy yellow brown, sometimes with grayish-green to bluish-gray tints, especially near the margin; distribution from Georgia south to Florida and west to Texas Leccinum glutinopallens (p. 247), cap slimy-viscid when fresh, glabrous, dingy vinaceous buff to pale pinkish gray or pale grayish brown; distribution northern, currently known from eastern Canada west to Michigan 20a. Cap dark brown to blackish, often mottled with cream-colored spots; stalk flesh staining orange red at cap junction, blue green at base → Leccinum snellii (p. 267) 20b. Cap paler → 21 21a. Cap at first dull orange, becoming tan or brownish; associated with birch → Leccinum versipelle (p. 271) 21b. Cap color variable, from whitish to pale tan grayish, brownish, pinkish buff, yellow brown, dark brown, or purplish red, but lacking orange tones → Leccinum scabrum (p. 265), cap grayish brown to yellow brown or dark brown, glabrous or matted-fibrillose, breaking up into tiny flattened scales in age; flesh unchanging or slowly staining reddish when exposed; associated with birch (see also Leccinum holopus [p. 248]) Leccinellum albellum (p. 238), cap color variable, white, whitish, pale tan, pale gray to pinkish gray, or pinkish brown to medium brown, sometimes tinged yellow or with a bluish cast, glabrous or somewhat velvety, often conspicuously pitted at maturity, sometimes cracked in age; associated with broadleaf trees (see also Leccinum holopus [p. 248]) Gyroporus purpurinus (p. 213), cap velvety, purplish red to burgundy; stalk scurfy to somewhat velvety, brittle and hollow, colored like the cap K ey E-1 Smooth and Miscellaneous Stalks, with White, Whitish, Grayish, Buff, Olive-Buff, or Pinkish-Tan Pores When Young Boletes in this group include those with a smooth or variously textured stalk that lacks reticulation, scabers, scales, rings, or resinous dots. Some may have slightly scurfy stalks or have longitudinal ribs that form a net-like pattern that mimics true reticulation. 1a. Pores staining when bruised, sometimes slowly → 7 1b. Pores unchanging when bruised → 2 2a. Cap viscid to glutinous when fresh → 3

42 · Boletes of Eastern North America 2b. Cap dry → 4 3a. Cap bright yellow to orange yellow; stalk viscid to glutinous when fresh, somewhat scurfy near the apex, nearly smooth below, pale yellow to yellow down to the base, which is sheathed with cottony white mycelium → Pulveroboletus curtisii (p. 294) 3b. Cap white to yellowish tan, yellowish brown, cinnamon, vinaceous brown, or dark brown; stalk at first with faint whitish resinous dots (use lens) that darken with age → Suillus brevipes (p. 325), cap light brown to dark brown, vinaceous brown, cinnamon brown, or grayish brown, fading to cinnamon or tan in age; stalk proportionally short, nearly glabrous, white to pale yellow, sometimes with brown stains near the base, with or without inconspicuous resinous dots that become visible only with age Suillus neoalbidipes (p. 344), cap white, pale pinkish cinnamon, dull orange yellow, or yellowish tan with a wide band of sterile tissue on the margin when young, becoming darker cinnamon to pale brown or yellowish brown 4a. Cap shaggy or scaly, often developing a network of ridges and pits, pinkish tan to golden yellow, yellow brown or salmon-tinged; pores white when young, becoming grayish pink to pinkish brown → Veloporphyrellus conicus (p. 407) 4b. Cap glabrous or fibrillose to velvety, smooth or cracking in age → 5 5a. Stalk brittle, soon becoming hollow; flesh firm and brittle → Gyroporus purpurinus (p. 213), cap vinaceous red to burgundy; widely distributed Gyroporus castaneus (p. 209), cap chestnut brown to orange brown or yellow brown; widely distributed Gyroporus subalbellus (p. 215), cap apricot buff to pinkish buff or with cinnamon tones; distribution along the Atlantic Seaboard from Cape Cod, Massachusetts, south to Florida and west to Texas Gyroporus phaeocyanescens (p. 212), cap fulvous to yellow brown or grayish brown; known distribution from Georgia south to Florida, west to Texas 5b. Stalk solid → 6 6a. Cap and stalk white, becoming whitish to pale straw-colored or tinged brownish; distribution southern, from the coastal plain of North Carolina south to Florida and west to Texas → Xanthoconium stramineum (p. 413) (see also Suillus grisellus [p. 338], Tylopilus intermedius [p. 382], and Leccinellum albellum [p. 238]) 6b. Cap and/or stalk with purplish or lilaceous tones, flesh intensely bitter → (if flesh is mild tasting, see Boletus separans [p. 153])

Field Keys to the Boletes · 43 Tylopilus plumbeoviolaceus (p. 388), cap brownish to grayish brown, at times tinged with purplish areas; stalk purple with white mycelium at the base when young, at times becoming grayish purple or purplish brown Tylopilus violatinctus (p. 402), cap grayish violet to bluish violet when young, becoming pale purplish, purplish pink, or tan to dull brown in age, staining rusty violet to dark violet when bruised; stalk concolorous with cap or paler (see also Tylopilus ferrugineus [p. 377] and Tylopilus minor [p. 384]) 7a. Cap at first bright orange to bright orange red, fading to dull orange, cinnamon, or tan in age; pores white to dingy white, becoming tan or slightly pinkish in age, staining brown when bruised → Tylopilus balloui (p. 374) 7b. Cap lacking bright-orange to bright orange-red tones → 8 8a. Cap at first white, whitish to straw yellow, buff, apricot buff, pinkish cinnamon, or tinged pinkish or ochraceous → 9 8b. Cap yellow, yellow brown, dark brown, cinnamon brown, chestnut brown, olive brown, grayish brown, maroon, purplish, or black → 12 9a. Pores bruising greenish blue to dark blue → 10 9b. Pores bruising brownish to pinkish cinnamon → 11 10a. Cap, stalk, pores, and flesh instantly staining dark blue to indigo blue when bruised or handled; stalk brittle, stuffed with pith, becoming chambered or hollow with age → Gyroporus cyanescens (p. 211) 10b. Cap and stalk not staining dark blue when bruised or handled; pores bruising greenish blue (sometimes weakly) and then grayish brown; stalk solid → Boletus pallidus (p. 131) (see also Phylloporus boletinoides [p. 285]) 11a. Cap at first apricot buff to pinkish buff; stalk becoming chambered or hollow; pores slowly staining pinkish cinnamon when bruised or in age; flesh unchanging when exposed → Gyroporus subalbellus (p. 215) 11b. Cap at first whitish to pale ochraceous, becoming tan to brownish or orangish brown; stalk solid → Tylopilus rhodoconius (p. 392), cap pale ochraceous to brownish orange at first, becoming pale brown and then darker brown with a paler margin; flesh white to pale creamy white with hyaline marbling, slowly staining pale pinkish and then dingy pinkish red when exposed; odor not distinctive or faintly of chlorine; taste mild at first and then very bitter Tylopilus peralbidus (p. 386), cap white when young, soon becoming ochraceous tan to chamois and finally brownish; flesh white, sometimes staining pale pinkish brown or buff when exposed, often

44 · Boletes of Eastern North America slowly; odor bleach-like or unpleasant; taste bitter (see also Tylopilus intermedius [p. 382] and Boletus pallidus [p. 131]) 12a. Cap yellow, yellow brown, chestnut brown, grayish brown, olive brown, or blackish → 13 12b. Cap maroon to dark brown, cinnamon brown, reddish brown, purple violet, or purplish → 16 13a. Cap black to dark grayish brown, brown, or olive brown → 14 13b. Cap yellow brown, chestnut brown, orange brown, reddish brown, or rusty red → 15 14a. Pores bruising dark blue to dark blue green and then brownish red; cap gray brown to olive brown or dark brown, often with dark greenish or bluish tints along the margin → Tylopilus sordidus (p. 396) 14b. Pores bruising blackish or reddish and then slowly black → Tylopilus alboater (p. 366), cap dry, somewhat velvety, occasionally finely cracked in age, black to dark grayish brown, often covered with a thin whitish bloom when young; stalk sometimes slightly reticulate at the very apex Tylopilus atronicotianus (p. 369), cap dry, glabrous or nearly so, somewhat shiny, olive brown, bronze brown, grayish brown, or pale brownish, at times imbued with yellowish tones, darker in age; stalk occasionally finely reticulate at the very apex 15a. Pores bruising pale cinnamon; cap yellow brown or dark rusty red to reddish brown at first, becoming dull orange to dark orange yellow → Aureoboletus roxanae (p. 67) 15b. Pores bruising dull yellow to brown → Tylopilus appalachiensis (p. 367), cap yellow brown over the disc, fading to pale yellow brown to dull tan on the margin or darker brown overall, at times with an orange or grayish tinge; pores becoming pinkish with age; stalk usually lacking reticulation but sometimes finely reticulate at the apex Xanthoconium affine (p. 409), cap dark brown to chestnut brown or ocher brown, sometimes with white to pale-yellow spots or patches; pores becoming yellowish to dingy yellow brown with age 16a. Flesh staining pinkish to brown when cut (sometimes slowly) → Tylopilus ferrugineus (p. 377), cap somewhat velvety, dark reddish brown; pores whitish at first, becoming pale pinkish buff, bruising dull brown; stalk dull brown to reddish brown, often whitish near the apex, with white basal mycelium; associated with oak Tylopilus badiceps (p. 371), cap maroon and velvety when young and fresh, soon purplish brown to dark reddish brown; pores white for a long time and then dingy white to brownish but not pinkish at maturity;

Field Keys to the Boletes · 45 stalk colored like the cap but often with more pronounced violaceous tones or sometimes violaceous nearly overall, usually white at the apex and base 16b. Flesh unchanging when cut → 17 17a. Flesh taste decidedly bitter → Tylopilus williamsii (p. 405), cap dark magenta to purple violet or brownish purple when young, fading to pinkish gray to pale yellow brown; known only from coastal Georgia and Florida west to Mississippi (see also Tylopilus plumbeoviolaceus [p. 388]) Tylopilus rubrobrunneus (p. 395), cap dark to bright purple when young, becoming purple brown, dark reddish brown, dull brown, or cinnamon; widely distributed 17b. Flesh taste mild to slightly tart but not bitter → Austroboletus gracilis var. gracilis (p. 70), cap maroon to reddish brown or cinnamon, at times tawny to yellow brown; pores becoming pinkish to pinkish brown with age Xanthoconium purpureum (p. 412), cap dark red to purple red or maroon, sometimes with pale-brownish tints; pores becoming ochraceous yellowish to rusty ochraceous Key E-2 Smooth and Miscellaneous Stalks, with Yellow to Pale-Yellow or Olive-Buff Pores at First Boletes keyed here lack scales, scabers, resin dots, stalk rings, or reticulation on the stalk. Some species may have longitudinal ribs on the stalk that intersect and form a net-like pattern (pseudoreticulation). If this feature is strongly expressed, these species may be placed with Net Stalks (Key B, p. 26). If the stalk is shaggy-lacerate, see Frostiella russellii (p. 207). 1a. Fruitbody attached to the earthball, Scleroderma citrinum; cap ocher brown to tawny olive or like tarnished brass; pores yellow, becoming olivaceous with age, sometimes with reddish or rusty stains, unchanging or very rarely becoming slightly bluish when bruised → Pseudoboletus parasiticus (p. 288) 1b. Fruitbody growing on soil or decaying wood → 2 2a. Pores elongated, decurrent, and radially arranged, at times nearly lamellate and intervenose → 3 2b. Pores not elongated, decurrent, radially arranged or at times nearly lamellate and intervenose → 6

46 · Boletes of Eastern North America 3a. Fruitbody occurring strictly with ash trees, stalk usually eccentric; cap slightly viscid and shiny when moist, yellow brown to olive brown or reddish brown → Boletinellus merulioides (p. 82) 3b. Fruitbody occurring with other broadleaf trees or conifers → 4 4a. Pores pale olive buff, becoming dark olive buff with age, gill-like with numerous cross-veins, or poroid; cap dry, somewhat velvety to minutely scaly, becoming nearly smooth with age, cinnamon to dark pinkish brown, fading to dull yellow brown in age → Phylloporus boletinoides (p. 285) 4b. Pores pale to bright yellow → 5 5a. Pores bright yellow, becoming golden yellow with age, strongly decurrent, typically gill-like with cross-veins, often wrinkled and sometimes forked; cap dull red to reddish brown, reddish yellow, or olive brown → Phylloporus rhodoxanthus (p. 286) (see also Boletus subfraternus [p. 159]) 5b. Pores pale yellow, becoming golden yellow and then brownish yellow; cap pinkish purple to reddish purple, covered with minute scales or hairs; associated with larch trees → Boletus paluster (p. 132) 6a. Fruitbody occurring on or around tree stumps and roots or other woody substrates → 7 6b. Fruitbody occurring on soil → 9 7a. Cap blackish brown, becoming chocolate brown to chestnut brown, velvety at first and often cracked in age; pores pale yellow, becoming yellow to greenish yellow, quickly staining blue when bruised; associated with broadleaf trees → Boletellus chrysenteroides (p. 78) (see also Xerocomus hypoxanthus [p. 422]) 7b. Cap bright yellow to golden yellow, orange red, or reddish brown; associated with pines → 8 (see also Boletellus ananas [p. 76] and Imleria badia [p. 227]) 8a. Cap dry, velvety, and suede-like, sometimes finely cracked in age, reddish brown to yellow brown or rusty golden yellow, often darker brown at the margin; pores typically decurrent, yellow to golden yellow at first, becoming brownish yellow in age, bruising dark greenish blue to blackish blue and then fading to reddish brown; flesh pale yellow, staining blue when exposed, sometimes slowly; odor sweet, somewhat minty or citrus-like; associated with and often found with the polypore Phaeolus schweinitzii → Buchwaldoboletus lignicola (p. 177) (see also Boletus oliveisporus [p. 127] and Buchwaldoboletus hemichrysus [p. 174]) 8b. Cap dry to slightly viscid, glabrous, often cracked in age, sulfur yellow when young, becoming pale yellow to whitish with age; flesh sulfur yellow to pale yellow, typically staining blue when exposed but sometimes slowly or not at all; odor not distinctive; pores bruising blue then slowly brownish → Buchwaldoboletus sphaerocephalus (p. 179)

Field Keys to the Boletes · 47 9a. Cap covered with coarse, purplish-red to dark-red scales and having whitish veil remnants on the margin; associated with oak or pine; southern in distribution → Boletellus ananas (p. 76) 9b. Cap lacking coarse scales → 10 10a. Cap viscid to glutinous when fresh and moist, shiny when dry → 11 10b. Cap dry, more or less glabrous or velvety, dull or shiny, with or without cracks or fissures → 14 11a. Pores bruising dark bluish gray to blackish blue; cap color variable, dark red or purplish to blood red or reddish orange, yellow on the margin → Boletus purpureorubellus (p. 139) 11b. Pores bruising brownish to dull brick red or unchanging when bruised → 12 12a. Cap bright yellow at first, becoming duller yellow with age; stalk viscid to glutinous when fresh, somewhat scurfy near the apex, smooth below, pale yellow to yellow down to the base, which is sheathed with cottony white mycelium → Pulveroboletus curtisii (p. 294) 12b. Cap light to dark brown, reddish brown, ochraceous to rusty orange or cinnamon → 13 13a. Pores bright golden yellow and remaining so well into age; cap pinkish brown to vinaceous brown when fresh, fading in age or when dry; taste of the cap surface is acidic; stalk viscid when fresh, pale yellow at the apex, streaked and flushed pale pinkish brown downward, with copious white mycelium at the base → Aureoboletus auriporus (p. 60) 13b. Pores pale yellow, becoming olivaceous brown; cap reddish brown to reddish cinnamon, becoming much darker brown in age, often with a pinkish tinge; stalk viscid when wet, glabrous, often longitudinally ridged → Pulveroboletus atkinsonianus (p. 289) (see also Suillus brevipes [p. 325] and Suillus bovinus [p. 321]) 14a. Cap corrugated to wrinkled or with pits or depressions → 15 14b. Cap more or less smooth, with or without cracks or fissures → 16 15a. Pores quickly bruising greenish blue to grayish blue; cap color variable, brick red or rosy red over a yellow ground color when young, becoming yellow orange or mustard yellow with dull pinkish or rosy tones in the folds, burgundy-red or brownish-red splashes on portions of mature specimens, slowly bruising greenish blue to grayish blue especially along the margin; known only from Florida → Lanmaoa roseocrispans (p. 235) 15b. Pores unchanging when bruised, widely distributed; cap tan to reddish tan or ocher brown to reddish brown → Xerocomus hortonii (p. 421) (see also Hemileccinum subglabripes [p. 220]) 16a. Pores bruising blue to greenish blue (sometimes slowly) → 21 16b. Pores unchanging when bruised or changing to colors other than blue or greenish blue → 17

48 · Boletes of Eastern North America 17a. Cap uniformly pinkish red or purplish red to blood red or dark red → Boletus roodyi (p. 141) 17b. Cap tan, yellow, brown, yellow brown, reddish brown, orange brown, grayish brown, or olive brown → 18 18a. Flesh instantly deep indigo blue when cut, brittle; cap fulvous to yellow brown or grayish brown; stalk brittle and soon becoming hollow; current known distribution from Georgia south to Florida and west to Texas → Gyroporus phaeocyanescens (p. 212) 18b. Flesh unchanging or staining weakly when cut → 19 19a. Distribution restricted to the Deep South, currently known only from sandy coastal woodlands in southern Georgia and the Gulf Coast of Florida; cap reddish brown to dark brown, cinnamon brown, or purplish brown; stalk typically abruptly bulbous at the base → Boletus abruptibulbus (p. 84) (see also Xerocomus hypoxanthus [p. 422] and Ceriomyces aureissimus var. castaneus [see photo, p. 92]) 19b. Distribution from South Carolina and northward → 20 20a. Cap glabrous or nearly so → Boletus alutaceus (p. 87), cap pinkish cinnamon to pinkish brown or pinkish tan; pores slightly greenish yellow (see also Pulveroboletus atkinsonianus [p. 289]) Aureoboletus roxanae (p. 67), cap yellow brown to reddish brown, becoming dull orange to dark orange yellow or tawny; pores whitish at first, soon becoming pale yellow and finally pale golden yellow (see also Boletus aureissimus var. aureissimus [p. 92] and Hemileccinum subglabripes [p. 220]) 20b. Cap suede-like to somewhat velvety → Aureoboletus innixus (p. 62), cap dry and somewhat velvety, frequently cracked in age, dull reddish brown to dull cinnamon or yellow brown, sometimes with purplish or reddish tones near the margin; stalk enlarged downward to a pinched base (see also Boletus alutaceus [p. 87] and Aureoboletus roxanae [p. 67]) Boletus subtomentosus var. subtomentosus (p. 165), cap olive brown to yellow brown, often cracked in age Pulveroboletus auriflammeus (p. 291), cap and stalk deep orange yellow when young to golden yellow or brownish orange in age; stalk with longitudinal ribs or reticulation toward the apex 21a. Cap predominantly red, pinkish red, rose red, purplish red, brick red, rusty brown, tawny brown, reddish brown, olive brown, mahogany, or with reddish tones → 25 21b. Cap predominantly whitish to pale buff, bright yellow, ochraceous yellow, orange ocher, yellow brown, ocher brown, or pale cinnamon brown → 22

Field Keys to the Boletes · 49 22a. Cap whitish to pale buff or pale brownish; pores whitish to pale yellow when young, becoming yellow to greenish yellow in age, quickly staining greenish blue and then grayish brown when bruised → Boletus pallidus (p. 131) 22b. Cap more highly colored → 23 23a. Cap bright yellow, ochraceous yellow, orange ocher, yellow brown, ocher brown, or pale cinnamon brown → 24 23b. Cap lacking yellow, orange-ocher, yellow-brown, ocher-brown, or cinnamon-brown tones → 25 24a. Distribution southern from Georgia and Florida west to Mississippi; cap ochraceous yellow to orange ocher or pale pinkish cinnamon with yellow on the margin at first, becoming dull brownish yellow in age → Boletus ochraceoluteus (p. 125) 24b. Distribution northern or more widely distributed than just from Georgia and Florida west to Mississippi → Boletus huronensis (p. 110), associated with hemlock trees from eastern Canada south to New York and Connecticut; cap dull yellow brown to pale cinnamon brown (compare with Cyanoboletus pulverulentus [p. 202], Caloboletus roseipes [p. 192], Boletus subtomentosus var. subtomentosus [p. 165], and Hemileccinum subglabripes [p. 220]) Neoboletus pseudosulphureus (p. 279), cap bright yellow at first, becoming reddish brown to brown in age; all parts instantly bruising blue to blackish blue (see also Lanmaoa pseudosensibilis [p. 233]) 25a. Cap olive to olive brown or grayish brown at first, possibly developing pinkish or reddish tones with age; cap surface velvety → 26 25b. Cap lacking olive tones at first → 27 26a. Cap dark olive to olive brown or grayish brown, developing surface cracks, with a red to pinkish color in the cracks, at least on the margin, velvety at first; stalk finely granular to somewhat scurfy, sometimes with longitudinal streaks, often yellow on the upper portion, usually reddish near the base → Xerocomellus chrysenteron (p. 416) 26b. Cap olive when very young, soon pinkish to brick red or dark red, often brownish red toward the margin, usually with olive to tarnished-brass tints, smooth, somewhat velvety; taste of flesh tart to acidic → Boletus patrioticus (p. 133) 27a. Cap at first predominantly brown, dull rusty brown to dull reddish brown, tawny brown, mahogany, cinnamon brown, dark yellow brown, or blackish brown → 28 27b. Cap at first predominantly red, brick red, purplish red, rose red, reddish brown, or pinkish red → 30 28a. Cap mahogany to reddish brown or rose brown, paler toward the margin, margin bruising blue; known only from Mississippi → Boletus mahogonicolor (p. 115)

50 · Boletes of Eastern North America 28b. Cap color variable; more widely distributed → 29 29a. All parts of the fruitbody instantly staining dark blue when handled or cut → Cyanoboletus pulverulentus (p. 202), cap dark yellow brown to blackish brown or dark cinnamon brown Boletus oliveisporus (p. 127), cap dark fulvous tinged with bay or cinnamon brown to dull brown 29b. Cap not staining blue when bruised → Lanmaoa pseudosensibilis (p. 233), cap dull rusty brown to dull reddish brown, fading to dull cinnamon or dingy yellow brown Boletus rufomaculatus (p. 148), cap at first dull rusty brown to ocher brown, becoming cracked, mottled with brick red or brownish red in age; usually occurring with beech 30a. Flesh decisively and fairly quickly staining blue when exposed → 31 30b. Flesh slowly and weakly staining blue when exposed or unchanging → 34 31a. Cap small (often 5 cm wide or less), rose red to pinkish red, typically cracking with age → Hortiboletus campestris (p. 223) (see also Boletus subfraternus [p. 159] and Xerocomellus intermedius [p. 419]) 31b. Cap typically broader than 5 cm → 32 32a. Flesh with pronounced odor of curry, fenugreek, maple syrup, or licorice; cap somewhat velvety when young, becoming nearly glabrous in age, dark to pale brick red, fading to dull rose or sometimes dingy cinnamon → Boletus sensibilis (p. 150) (see also Boletus miniatopallescens [p. 119]) 32b. Flesh odor not distinctive → 33 33a. Taste of flesh acidic or tart; occurring in sandy soil under oak or in oak–pine woods; southern in distribution from Florida to Texas but also reported as far north as New Jersey; cap color variable, dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown → Boletus rubricitrinus (p. 145) 33b. Taste of flesh not distinctive; more broadly distributed in eastern North America → Boletus miniatopallescens (p. 119), cap red to brick red, soon fading to reddish orange or orange yellow, often cracked in age Lanmaoa carminipes (p. 231), cap pinkish red to rose red at first, becoming dingy brownish, often yellowish or paler pinkish red along the margin, quickly staining dark blue when bruised or handled Imleria badia (p. 227), cap chestnut brown, reddish brown, or yellow brown, sometimes with olive tones, dry to slightly viscid, smooth or somewhat velvety Xerocomellus intermedius (p. 419), cap red to rose red, often soon developing grayish to olive-brown tones, surface dry, subtomentose, becoming cracked, with pink tints in the cracks (see also Boletellus pseudochrysenteroides [p. 80])

Field Keys to the Boletes · 51 34a. Stalk completely glabrous and smooth or in rare instances having slight reticulation at the apex; tube layer shallow → Baorangia bicolor complex (p. 74), cap dark red to rose red, fading to ochraceous or tan, often cracking in dry weather or in age; flesh unchanging or slowly and weakly staining blue when exposed; odor not distinctive or reminiscent of cheese in older specimens Boletus rufomaculatus (p. 148), cap red to dull red, becoming cracked, mottled with brick red or brownish red; flesh slowing staining blue, especially above the tubes, or sometimes unchanging; odor unpleasant or not distinctive Boletus pallidoroseus (p. 129), cap uniformly pale rose pink, pale purplish pink, or mottled with darker-pink tones, golden yellow or paler yellow along the margin, becoming pale orange brownish with age; flesh slowly staining blue when exposed; odor reminiscent of beef or chicken bouillon or of spoiled meat 34b. Stalk somewhat roughened or dotted or with darker flakes → 35 35a. Stalk bruising dark blue or darkening brown when handled → Xerocomus sclerotiorum (p. 430), cap pinkish red to rose red or purplish red, sometimes with olive tints, becoming dull rose pink to brownish pink in age, slowly staining blackish blue when bruised; known from New England south to Florida, west to Tennessee Hortiboletus rubellus (p. 225), cap dark red, fading with age to brick red or olivaceous brown, often becoming finely cracked in age; widely distributed (see also Boletus harrisonii [see photo, p. 222], Hortiboletus campestris [p. 223], and Boletus miniato-olivaceus [p. 117]) 35b. Stalk not bluing or discoloring when handled → Boletus miniato-olivaceous (p. 117), cap red to rosy red when young, becoming pale rose pink and developing olive tones, finally becoming rosy tan to olive yellow with rosy tints, often staining dark blue when bruised; widely distributed (see also Boletus patrioticus [p. 133]) Xerocomus hypoxanthus (p. 422), cap tawny brown to pale red brown or pale yellow brown to orange brown; known only from South Carolina and Florida west to Mississippi Key E-3 Smooth and Miscellaneous Stalks, with Red to Rosy-Red or Coppery-Red, Orange, Maroon, Cinnamon-Brown, Reddish-Brown, or Coffee-Brown Pores 1a. Pores radially arranged, at times gill-like, buff to olive buff; cap somewhat velvety to minutely scaly, becoming nearly smooth in age, cinnamon

52 · Boletes of Eastern North America to dark pinkish brown, fading to dull yellow brown → Phylloporus boletinoides (p. 285) 1b. Pores not radially arranged → 2 2a. Pores bruising blue, greenish blue, or blackish blue → 8 2b. Pores not bruising blue, greenish blue, or blackish blue → 3 3a. Flesh taste decidedly acrid-peppery; base of stalk bright yellow → Chalciporus piperatus (p. 195) 3b. Flesh lacking acrid-peppery taste → 4 4a. Flesh taste tart and lemony; cap brownish olive at first, becoming olive brown, sometimes with reddish tints, cracked and scaly with pale yellow flesh showing in the cracks; occurring on sandy soil with longleaf pine from Florida west to Texas → Boletus weberi (p. 171) 4b. Flesh taste not distinctive or bitter to slightly acidic → 5 5a. Pore surface uniformly dark cinnamon brown to reddish brown; cap medium brown to rusty brown, becoming paler brown to yellowish brown with age, typically yellowish along the margin; known only from the eastern piedmont region of North Carolina → Boletus durhamensis (p. 99) (see also Tylopilus porphyrosporus [p. 389] and Buchwaldoboletus hemichrysus [p. 174]) 5b. Pores at first bright rose red to grayish red; cap less than 6 cm wide → 6 6a. Pores at first grayish red to dull red; cap dark orange yellow and staining brown when young, becoming reddish brown to orange brown at maturity; known only from Florida → Chalciporus rubritubifer (p. 199) 6b. Pores bright rose red, becoming dull rose, orangish, or yellow brown with age; widely distributed north of North Carolina → 7 7a. Stalk finely pruinose, mostly rose pink, and typically yellow near the base, with yellow basal mycelium; cap yellowish to pinkish red, becoming pinkish cinnamon in age → Chalciporus pseudorubinellus (p. 196) 7b. Stalk red or reddish, sometimes mixed with yellow, lacking yellow mycelium at the base; cap red or reddish when young, more yellow and becoming cracked with age → Chalciporus rubinellus (p. 198) 8a. Pores dark brown to coffee brown, maroon or brownish yellow; flesh quickly staining blue when cut; odor pungent → Boletus vermiculosoides (p. 169), cap yellow when young, becoming brown with maturity; stalk brownish pruinose over a dull-whitish or pale-yellow ground color (compare with Boletus vermiculosus [see photo p. 169]) Boletus subgraveolens (p. 161), cap dull yellow brown overall or mottled with darker or paler shades of yellow or brown, usually staining dark blue and then darker brown when fresh, becoming cracked in dry weather; stalk dull pale yellow and pruinose on the lower portion with a dark vinaceous-red to maroon base

Field Keys to the Boletes · 53 8b. Pores at first red to dark red, pinkish red, orange red, coppery red, or reddish maroon, sometimes with a yellowish margin → 9 (see also comments under Chalciporus piperatus [p. 195] and the photos of Chalciporus piperatoides [p. 195]) 9a. Flesh staining blue when exposed, sometimes slowly → 10 9b. Flesh unchanging or becoming darker yellow when exposed; cap dull red at first, becoming pinkish red to orange red, color fading with age to reddish orange to dull golden orange; current known distribution from North Carolina south to Florida, west to Arkansas and Louisiana → Boletus carminiporus (p. 97) 10a. Stalk glabrous to faintly longitudinally striate; cap dark reddish salmon to bright orange, becoming rosy brick red to reddish brown, fading to brownish orange with yellow tints and a whitish margin; recorded only from McComb, Mississippi → Boletus roseolateritius (p. 143) (see also Boletus fairchildianus [p. 104]) 10b. Stalk more or less glabrous to faintly or at least partially pruinose, scurfy, or punctate → 11 11a. Distribution from North Carolina south to Florida and west to southeast Texas → Suillellus hypocarycinus (p. 310), cap brown to yellow brown, sometimes with olive or cinnamon tones (compare with Boletus subluridellus [p. 163]) Suillellus subluridus (p. 314), cap orange pink to orange yellow with vinaceous tints or red to purplish red, sometimes rusty red to cinnamon red, developing more brownish tones with age 11b. Distributed from South Carolina northward → 12 (see also Suillellus subvelutipes [p. 315]) 12a. Cap brownish to dark olive brown, dark reddish brown, or nearly blackish brown, staining blue to blackish blue when bruised → Neoboletus luridiformis (p. 277), cap velvety when young, dark brown to nearly blackish brown, becoming dark reddish brown to olive brown; known distribution from eastern Canada south to South Carolina and west to Minnesota Boletus pseudo-olivaceus (p. 135), cap pale brownish olive to yellowish olive; stalk with reddish pruina that may form a pattern that simulates reticulation on the upper portion; known distribution from New England south to North Carolina and west into the Great Lakes region 12b. Cap bright red to pinkish red, orange red, brick red, purplish red, reddish brown, or cinnamon brown, all parts staining blue to blackish blue when cut or bruised → 13

54 · Boletes of Eastern North America 13a. Stalk yellow, pruinose; cap purplish red, brownish red, pinkish red, or orange red; distribution from New England and New York south to Georgia and west to Minnesota → Boletus subluridellus (p. 163) 13b. Stalk reddish to orange cinnamon (may be yellow on the upper portion) → Suillellus subvelutipes (p. 315), cap color variable, cinnamon brown to yellow brown, reddish brown, or reddish orange to orange yellow; stalk base often with stiff yellow or reddish hairs; all parts instantly bruising blackish blue Boletus rufocinnamomeus (p. 146), cap dull brick red with an ochraceous overtone from an evanescent bloom when young, becoming reddish brown to dull cinnamon brown with age Neoboletus luridiformis (p. 277), cap velvety when young, dark brown to nearly blackish brown, becoming dark reddish brown to olive brown; known distribution from eastern Canada south to South Carolina and west to Minnesota Lanmaoa borealis (p. 229), cap dark red to bright apple red, becoming dull brick red to rusty rose in age; yellow flesh that slowly stains blue at the stalk apex; orange red to red pores that stain greenish blue; and a red stalk that may be reticulate at the apex

55 Descriptions and Illustrations of Bolete Genera and Species Genus Alessioporus Gelardi, Vizzini & Simonini The genus Alessioporus was erected in 2014 to accommodate species formerly classified in several genera, including Boletellus and Boletus. Alessioporus was named in honor of the late Carlo Luciano Alessio (1919–2006), who dedicated most of his mycological studies to the investigation of Italian boletes. Species in this genus have dry caps, reticulate stalks, and yellow flesh that quickly stains dark blue when exposed. At present, it is a small genus consisting of two European and one American species that have been shown to be closely related when multigene molecular analysis was performed. Alessioporus rubriflavus (A)

56 · Boletes of Eastern North America Alessioporus rubriflavus J. L. Frank, A. R. Bessette & A. E. Bessette, nom. prov. common name(s): none. overview: The term rubriflavus means “red and yellow,” a reference to the colors of the fruitbody. Red portions of the cap cuticle immediately stain amber with KOH, and the flesh immediately stains orange with KOH. Cap: 5–14(21) cm wide, convex, becoming broadly convex to nearly plane at maturity, margin with a narrow band of sterile tissue when young, becoming even; surface slightly viscid or dry, slightly velvety, sometimes finely cracked in age, dark wine red on very young buttons, soon developing a yellow ground color covered with streaks and splashes of various shades of wine red, red brown, and ocher, sometimes retaining wine-red coloration well into maturity, becoming olive to brownish over the disc in age, staining greenish blue to bluish black when bruised; flesh firm, bright yellow, quickly staining dark blue when exposed; odor not distinctive or mildly unpleasant; taste unpleasant, sour, or sometimes astringent. pores and tubes: yellow at first, soon becoming pale orange yellow and then olive brown, deeply depressed at maturity, quickly bruising blue then slowly reddish brown; tubes 8–25 mm deep. stalk: 4.5–11 cm long, enlarged downward, ventricose to clavate, typically with a pinched base, solid; surface dry, conspicuously reticulate over at Alessioporus rubriflavus (B)

Aureoboletus · 57 least the upper half, finely velvety over the lower half, bright yellow near the apex or nearly overall, with dark wine-red to red-brown streaks and splashes below, especially near the base, quickly staining blue and then slowly brownish when bruised; reticulation yellow toward the apex, yellow brown below, darkening when handled, with a whitish mycelium; flesh bright yellow, quickly staining blue when exposed, usually dark wine red near the base. habitat and season: solitary, scattered, or in groups on the ground in mixed pine and oak woods; summer and fall. distribution and frequency: Georgia and South Carolina northward to southern New York, distribution limits yet to be determined; occasional to locally common. spore print: olive brown. microscopic features: spores 13–19 × 4–6 µm, subellipsoid to subfusiform, smooth, pale brownish yellow. edibility: unknown. lookalikes: Boletus flavissimus (Murrill) Murrill (not illustrated), originally described from Florida, has a bright-yellow cap, lacks red coloration in all stages of development, does not stain when bruised, and its stalk lacks reticulation. Butyriboletus brunneus (p. 181) has a reddish-brown or yellow-brown to olive-brown cap; pale-yellow flesh that quickly stains blue; a yellow, finely reticulate stalk with red on the lower portion; and smaller spores, 10–15 × 3–5.5 µm. Neoboletus pseudosulphureus (p. 279) has a bright-yellow cap that becomes duller yellow to tawny at maturity and typically develops brown to brownish-red tints, especially over the disc, in age. It has a yellow stalk that sometimes develops reddish tints, especially near the base, and usually lacks reticulation or is sometimes reticulate on the upper portion. It has smaller spores that measure 10–16 × 4–6 µm. Genus Aur eobolet us Pouzar The genus Aureoboletus was erected in 1957 to accommodate species formerly classified in the genus Boletus. The name Aureoboletus means “brilliant golden yellow,” a reference to the color of the pores of the species in this genus. It is a small genus of medium-size terrestrial boletes, with only five species occurring in eastern North America. They have viscid or dry caps that sometimes have a projecting sterile margin. A white mycelium is often present at the base of the stalk. The pores are brilliant golden yellow at first and become dull golden yellow to greenish yellow or brownish yellow in age. The cap and stalk flesh is white and does not stain when cut or bruised. These species have olive-brown spore prints.

Aureoboletus auriporus (A) Aureoboletus auriporus (B)

Aureoboletus auriporus (C) Aureoboletus gentilis

60 · Boletes of Eastern North America Aureoboletus auriporus (Peck) Pouzar = Boletus auriporus Peck common name(s): Acidic Golden-pored Bolete, Golden-pored Bolete overview: The combination of the pinkish-brown to vinaceous-brown cap and brilliant golden-yellow pores is very striking. Both the cap cuticle, which is distinctly acidic tasting, and the tube layer of this bolete are easily removed. The term auriporus means “golden pores,” in reference to the color of this beautiful bolete’s pores. cap: 2–8 cm wide, convex, becoming broadly convex to nearly plane, margin with a narrow band of sterile tissue; surface coated with tiny appressed fibrils, smooth or sometimes bumpy and appearing blistered, moist and viscid when fresh, becoming dull when dry, pinkish brown to vinaceous brown when fresh, fading in age or when dry; taste of the cap surface is acidic; flesh white to pale yellow, except vinaceous under the cap surface, typically not staining blue when cut or bruised, sometimes staining weakly blue or pinkish; odor somewhat pungent or not distinctive; taste acidic or not distinctive. Pores and tubes: brilliant golden yellow when young and fresh and remaining so well into maturity, becoming dull golden yellow to greenish yellow and then brownish yellow in age, usually slowly staining dull brick red when bruised or sometimes unchanging; tubes 6–15 mm deep. Stalk: slightly enlarged downward or nearly equal, typically narrowed abruptly at the base; surface viscid to tacky when fresh, pale yellow at the apex, streaked and flushed pale pinkish brown downward, with copious white mycelium at the base; flesh concolorous with the cap flesh. Habitat and season: solitary to scattered or in groups under oak; late spring, summer, and fall. Distribution and frequency: New Jersey to Florida, west to Mississippi and Texas; fairly common. Spore print: olive brown. Microscopic features: spores 11–16 × 4–6 µm, fusiform-ellipsoidal, smooth, pale brown. Edibility: edible and somewhat acidic tasting. Lookalikes: Aureoboletus gentilis (Quél.) Pouzar (see photo, p. 59) has a pale gray-pink to pinkish-brown streaked cap and grows under broadleaf trees, especially oak and beech. Aureoboletus innixus (p. 62) has a darkerbrown dry cap, a cap surface lacking an acidic taste, and flesh that has an odor reminiscent of witch hazel. It often grows in clusters fused at the stalk bases.

Aureoboletus innixus (A) Aureoboletus innixus (B)

62 · Boletes of Eastern North America Aureoboletus innixus (Frost) Halling, A. R. Bessette & Bessette = Boletus innixus Frost = Boletus caespitosus Peck = Pulveroboletus innixus (Frost) Singer Common name(s): Clustered Brown Bolete Overview: This common small- to medium-size brown bolete, which lacks any prominent distinguishing feature, is often present during dry periods when other boletes are scarce. The term innixus means “reclining,” alluding to the fruitbodies, which are often leaning, especially when tightly clustered. The cap surface stains mahogany red to reddish brown with KOH, develops a green flash and then stains dull orange red with NH4OH, and is pale olive with FeSO4. The flesh stains pale dull pinkish orange with KOH or NH4OH and pale gray with FeSO4. Cap: 3–7.5 cm wide, convex, becoming broadly convex to nearly plane at maturity, margin with a narrow band of sterile tissue; surface dry and somewhat velvety, frequently cracked in age, dull reddish brown to dull cinnamon, or yellow brown, sometimes with purplish or reddish tones near the margin; flesh white to pale yellow or tinged vinaceous beneath the cuticle; odor pungent, reminiscent of witch hazel; taste slightly lemony or not distinctive. Pores and tubes: bright yellow when young, becoming dull yellow, unchanging when bruised; tubes 3–10 mm deep. Stalk: short and stout, typically enlarged downward, club-shaped and distinctly swollen above a tapered base, solid; surface moist or dry on the upper portion, viscid near the base, yellowish and streaked with darkbrown tones, with yellow basal mycelium sometimes visible; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or often in caespitose clusters in broadleaf woods, especially with oak; late spring through fall. Distribution and frequency: eastern Canada south to Georgia, west to Michigan; occasional to locally common. Spore print: olive brown. Microscopic features: spores 8–11 × 3–5 µm, ellipsoidal, smooth, pale brown. Edibility: edible but not highly regarded. Lookalikes: Aureoboletus auriporus (p. 60) has a viscid cap with a peelable cuticle that tastes acidic and flesh that lacks a distinctive odor.

Aureoboletus projectellus (A)

64 · Boletes of Eastern North America Aureoboletus projectellus (Murrill) Halling = Boletellus projectellus (Murrill) Singer = Boletus projectellus (Murrill) Murrill = Ceriomyces projectellus Murrill Common name(s): none Aureoboletus projectellus (B)

Aureoboletus · 65 Overview: Although not common for much of its range, this tall, slender-stalked bolete is not difficult to recognize. The dull-pinkish to cinnamon-brown cap, concolorous coarsely ribbed stalk, and association with pines are distinctive features. The term projectellus means “extending or projecting,” a reference to the cap cuticle, which conspicuously projects beyond the tube layer. The identification features include a large pinkish to reddish-brown cap with an overlapping cap cuticle, a long reddish-brown reticulate stalk with a copious white basal mycelium, growth under pines in late summer and fall, and very large spores. Cap: 4–20 cm wide, convex, becoming broadly convex to nearly plane in age, margin with a distinct band of sterile tissue; surface dry, somewhat velvety when young, often cracked in age, pale to dark cinnamon brown or dark reddish brown, occasionally with gray or olive shades, especially when young; flesh whitish, often with a rosy tinge, not staining blue when exposed but slowly changing to yellow brown; odor not distinctive; taste somewhat acidic. Pores and tubes: pale yellow to olive yellow when fresh, becoming brownish olive in age, staining lemon yellow when bruised, depressed at the stalk in age; tubes 1–2.5 cm deep. Stalk: unusually long, up to 24 cm, and up to 5 cm thick, equal or enlarging downward, solid; concolorous with the cap or somewhat paler, often with a conspicuous covering of soft white mycelium at the base and with prominent coarse reticulation overall or at least on the upper two-thirds, distinctly viscid at the base in wet weather; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or scattered under pines, especially two-needle Scots pine; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to locally common. Spore print: olive brown. Microscopic features: spores 18–33 × 7.5–12 µm, oval to ventricose, smooth, pale brown. Edibility: edible and good when fresh, flavorless when dried and reconstituted. Lookalikes: Perhaps the bolete that is most likely to be confused with Aureoboletus projectellus is Frostiella russellii (p. 207), which has a scaly cap and a more lacerated stalk. Imleria badia (p. 227) is similarly colored but smaller and has a nonreticulate stalk and pale-yellow pores that stain blue when bruised.

Aureoboletus roxanae (A) Aureoboletus roxanae (B)

Aureoboletus · 67 Aureoboletus roxanae (Frost) Klofac = Boletus roxanae Frost = Boletus roxanae var. auricolor Peck = Ceriomyces roxanae (Frost) Murrill = Xerocomus roxanae (Frost) Snell Common name(s): Roxanna’s Bolete Overview: Although this small and relatively nondescript bolete lacks dramatic features, it is easily recognized once encountered a time or two. Its distinguishing features include a granulose yellow-brown cap and a club- or bowling-pin-shaped yellow to pale orange-yellow stalk, typically with a distinct dull-orange zone at the apex. The term roxanae means “Roxanna’s”: this bolete is named after the wife of American mycologist Charles C. Frost (1805–1880). Cap: 3–9 cm wide, convex to broadly convex, becoming nearly plane in age, margin even; surface dull, dry, granular-scaly when young, becoming nearly glabrous, sometimes cracking at maturity, yellow brown or dark rusty red to reddish brown at first, becoming dull orange to dark orange yellow, fading somewhat in age, margin often paler than the disc; flesh whitish to pale yellow, not staining when exposed; odor weakly pungent or not distinctive; taste unpleasant. Pores and tubes: whitish at first, soon becoming pale yellow and finally pale golden yellow in age, darkening somewhat or developing pale cinnamon stains when injured; tubes 6–10 mm deep. Stalk: enlarged downward, solid; surface dry, faintly pruinose and longitudinally striate, at least on the upper portion, yellow to pale orange yellow or tawny, typically with a distinct dull-orange zone at the apex; flesh concolorous with the cap flesh or at times having pinkish tones. Habitat and season: solitary or scattered under oaks, usually in mixed oak and pine woods; summer and fall. Distribution and frequency: eastern Canada south to South Carolina, west to Minnesota; occasional. Spore print: dark olive brown. Microscopic features: spores 8–13 × 3–5 µm, fusiform-ellipsoid, smooth, yellowish. Edibility: edible but usually having an unpleasant taste. Lookalikes: Boletus alutaceus (p. 87) has been reported from New England south to Kentucky and west to Ohio and Wisconsin, and it grows in broadleaf and mixed woods, especially with oak and beech. It initially has a yellowish-brown cap that becomes pale tan with a pinkish tint, pale greenish-yellow pores that do not stain when bruised, and whitish to pale-tan flesh that does not stain when exposed. Aureoboletus innixus (p. 62) has

68 · Boletes of Eastern North America much brighter-yellow pores and a brownish stalk, and it usually grows in clusters fused at the stalk bases. Genus Aust robolet us (Corner) Wolfe The genus Austroboletus was erected in 1980 to accommodate species formerly classified in the genus Tylopilus. The name Austroboletus means “southern bolete.” These boletes are medium-size terrestrial boletes with viscid or dry caps that sometimes have a projecting sterile margin and often have unusually long stalks in relation to their cap diameters. A conspicuous white mycelium is usually evident at the base of the stalks. The cap and stalk flesh is white to pale vinaceous and does not change color when cut or bruised. These mushrooms’ pores are white at first, darken to pinkish brown at maturity, and do not stain blue; they may darken or stain brownish when bruised. The colors of their spore prints vary from vinaceous pink to pinkish brown or reddish brown. Microscopically, their spores are obscurely to distinctly pitted or sinuously pitted. Two species of Austroboletus currently occur in eastern North America. Austroboletus gracilis var. gracilis

Austroboletus gracilis var. flavipes Austroboletus gracilis var. pulcherripes

70 · Boletes of Eastern North America Austroboletus gracilis var. gracilis (Peck) Wolfe = Tylopilus gracilis (Peck) Henn. = Porphyrellus gracilis (Peck) Singer Common name(s): Graceful Bolete Overview: The Graceful Bolete is not uncommon, though it seldom occurs in large numbers. It is typically solitary or scattered on the ground but occasionally will fruit on decaying logs and stumps. The term gracilis means “slender,” a reference to the stalk. Cap: 3–10 cm wide, convex to broadly convex, margin even; surface dry, finely velvety when young, sometimes cracked in age, maroon to reddish brown or cinnamon, at times tawny to yellow brown; flesh white or tinged pink, not staining when exposed; odor not distinctive; taste slightly tart or not distinctive. Pores and tubes: white when young, becoming pinkish to pinkish brown or burgundy tinged in age, darkening or staining brownish when bruised; tubes 1–2 cm deep. Stalk: long and slender in relation to the cap diameter, solid, enlarged downward or nearly equal, often curved; surface dry, concolorous with the cap or paler, with elevated and anastomosing rib-like lines that sometimes form an obscure, narrow reticulation overall or at least on the upper half, basal mycelium white; flesh concolorous with the cap flesh or sometimes yellow toward the base. Habitat and season: solitary, scattered, or in groups on the ground or sometimes on decaying wood in conifer and broadleaf forests; late spring, summer, and fall. Distribution and frequency: eastern Canada south to Florida, west to Tennessee and Michigan; occasional. Spore print: pinkish brown to reddish brown. Microscopic features: spores 10–17 × 5–8 µm, narrowly ovoid to subellipsoidal, pitted, pale brown. Edibility: edible. Lookalikes: Some species of Tylopilus are similar and have the same sporeprint colors. However, they have smooth spores. Gyroporus castaneus (p. 209) and Gyroporus purpurinus (p. 213) have white pores that do not become pinkish brown at maturity, hollow stalks, yellow spore prints, and smooth spores. Austroboletus gracilis var. flavipes T. J. Baroni, Halling & Both (see photo, p. 69), differs from Austroboletus gracilis var. gracilis by having more yellowish colors overall and by lacking both anastomosing riblike lines on the stalk and reticulation. Austroboletus gracilis var. pulcherripes Both & Bessette (see photo, p. 69), differs from var. gracilis by having darker colors overall and a stouter stalk that is coarsely reticulate over the upper two-thirds.

Austroboletus · 71 Austroboletus subflavidus (Murrill) Wolfe = Tylopilus subflavidus Murrill = Porphyrellus subflavidus (Murrill) Singer Common name(s): White Coarsely-ribbed Bolete Overview: The term subflavidus means “nearly yellow,” a reference to the color of this bolete’s cap and stalk. Specimens of this beautiful bolete are often stunted because they dry out quickly in their sandy habitat. The white to pale-yellow cap and stalk, coarsely ribbed reticulation, and bitter-tasting flesh are the key identification features. Cap: 3–10 cm wide, convex, becoming nearly plane and usually slightly depressed in age, margin even; surface dry, finely velvety at first soon becoming cracked, white at first, becoming buff or yellowish to grayish, often with a pale pinkish-orange tinge in age; flesh white, unchanging when exposed or bruised; odor somewhat fruity; taste bitter. Pores and tubes: white to grayish, sometimes beaded with clear droplets at first, becoming pinkish at maturity, not staining when bruised; tubes 1–2 cm deep. Stalk: nearly equal or tapered downward, rarely enlarged at the base, solid; surface dry, concolorous with the cap, with thick, raised, and coarsely ribbed reticulation that has a pitted appearance; flesh yellow in the base and white above. Habitat and season: scattered or in groups under oak and pine; late spring, summer, and fall. Austroboletus subflavidus

72 · Boletes of Eastern North America Distribution and frequency: New Jersey Pine Barrens south to Florida, west to Mississippi along the Gulf Coast; infrequent or sometimes fairly common locally. Spore print: reddish brown. Microscopic features: spores 15–20 × 6–9 µm, fusoid, minutely pitted, pale brown. Edibility: inedible because of the bitter taste. Lookalikes: Tylopilus rhoadsiae (p. 391) is similarly colored and has bitter-tasting flesh. Its stalk is typically clavate with a pinched base and prominent brown reticulation that is not coarsely ribbed, thick, raised, or appearing pitted, and its spores are smaller and smooth. Genus Baor angia G. Wu & Zhu L. Yang The genus Baorangia was erected in 2015 to accommodate species formerly classified in the genus Boletus. The name Baorangia is a combination of the Chinese words bao, meaning “thin,” and rang, meaning “hymenium,” a reference to short tubes. Species in this genus have tubes that are one-fifth to one-third the thickness of the cap flesh at a position halfway to the cap center. They have yellow tubes and pores that stain blue when bruised and yellow flesh that is unchanging or that slowly and weakly stains blue when exposed. At this time, there are four known species of Baorangia worldwide, with only one occurring in eastern North America. Baorangia bicolor complex (A)

Baorangia bicolor complex (B) Baorangia bicolor complex (C)

74 · Boletes of Eastern North America Baorangia bicolor complex (Kuntze) G. Wu, Halling & Zhu L. Yang = Baorangia rubelloides G. Wu, Halling & Zhu L. Yang = Boletus bicolor Peck = Boletus rubellus subsp. bicolor (Kuntze) Singer = Boletus bicolor var. subreticulatus A. H. Sm. & Thiers Boletus bicolor Massee and Boletus bicolor Raddi are names assigned to different species. Common name(s): Red and Yellow Bolete, Two-toned Bolete Overview: The term bicolor means “two colors.” The Red and Yellow Bolete is generally considered to be a good edible but is easily confused with similar species that are suspected to be poisonous or of uncertain edibility. Its distinguishing features are a red cap and stalk, bright-yellow pores that quickly stain blue, a tube layer that is shallow in proportion to the thickness of the flesh, and yellow flesh that is unchanging or slowly and weakly stains blue when exposed. Although most collections exhibit the macroscopic and microscopic features that closely agree with those provided in the original description, some collections display a wide variation in color and staining of the bruised cap surface, the degree of blue staining of the flesh, and the amount of the red color present on the stalk. Odor of the flesh is also inconsistently interpreted. These differences strongly suggest a group of closely allied species that ultimately will require molecular analysis to confirm their relationship. Until more information is available, we have Baorangia bicolor complex (D)

Baorangia · 75 decided to treat Baorangia bicolor as a complex and have included photos of several examples that illustrate this variability. Cap: 5–12.5 cm wide, convex to broadly convex, becoming nearly plane in age, margin incurved at first; surface dry, somewhat velvety when young, often cracking in dry weather or with age, dark red to rose red or brick red, rarely yellow, fading to ochraceous or tan with age, sometimes staining blue when bruised; flesh yellow, unchanging or slowly and weakly staining blue when exposed; odor not distinctive or reminiscent of cheese in older specimens; taste not distinctive. Pores and tubes: bright yellow, becoming olive yellow in age, quickly staining blue when bruised; tubes 3–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, smooth, yellow at the apex, red or rosy red on the lower two-thirds or more, unchanging or sometimes slowly staining blue when bruised, typically lacking reticulation but sometimes reticulate on the upper portion; flesh usually bright golden yellow, deeper yellow than the cap, unchanging or erratically staining blue. Habitat and season: solitary, scattered, or in groups under oaks; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: olive brown. Microscopic features: spores 8–12 × 3.5–5 µm, oblong to slightly ventricose, smooth, pale brown. Edibility: edible with caution and very good. Lookalikes: Boletus bicoloroides A. H. Sm. & Thiers (not illustrated), reported from Michigan, has a deep vinaceous-red cap that slowly fades to dull ochraceous with red splashes and does not become cracked in age. It has yellow flesh that stains blue when exposed, yellow pores that stain blue when bruised, tubes up to 15 mm deep, and a dark-red stalk from the base to the apex. Microscopically, it has large strongly amyloid spores that measure 14–17 × 4–5.5 µm and strongly dextrinoid pileus trama. The combination of the strongly amyloid spores with the strongly dextrinoid pileus trama is most unusual and clearly separates this species from other similar boletes. Boletus sensibilis (p. 150) quickly bruises blue on all parts and has a mostly yellow stalk and longer tubes, and its flesh has a sweet odor of fenugreek or curry. Boletus pallidoroseus (p. 129) is somewhat similar. It has a pale rose-pink to pale purplish-pink cap typically with a yellow margin and flesh with an odor of beef or chicken bouillon. Also compare with Boletus rufomaculatus (p. 148), which has a mottled brick-red to brownish-red cap with a narrow band of sterile tissue and paler-yellow stalk flesh and grows under beech.

76 · Boletes of Eastern North America Genus Bolet ellus Murrill The genus Boletellus was erected in 1909 to accommodate species formerly classified in the genera Boletus and Suillus. The name Boletellus means “small Boletus.” It is a fairly small genus with less than a dozen species known from North America. They are small- to medium-size terrestrial or lignicolous boletes that closely resemble species of the genus Boletus. Their caps are typically dry, somewhat velvety or scaly, and often cracked in age, sometimes with a sterile projecting margin. Stalks are typically solid at maturity and lack scabers and resinous dots. They have white to yellow flesh that often stains blue when cut. Spore-print colors may be olive brown or dark rusty brown to dark brown. Microscopically, their spores are ornamented with longitudinal ridges, striations, or wings. Boletellus ananas (M. A. Curtis) Murrill = Boletus ananas M. A. Curtis Common name(s): Pineapple Bolete Overview: The Pineapple Bolete is a distinct species unlikely to be confused with any other southern bolete. The term ananas refers to the genus of pineapple because of the scales on the cap that extend beyond the cap margin, which give this bolete its unique appearance. Boletellus ananas (A)

Boletellus · 77 Cap: 3–10 cm wide, obtuse at first, becoming convex to broadly convex; surface dry, coated with coarse and overlapping purplish-red to dark-red scales that become dull pinkish tan to dingy yellow in age and extend beyond the margin; flesh whitish on exposure, quickly becoming yellowish and then blue and finally bluish gray; odor and taste not distinctive. Pores and tubes: yellow when fresh, sometimes tinged reddish brown in age, staining blue when bruised, covered when young by a whitish partial veil that usually leaves remnants on the cap margin and normally does not form a ring on the stalk; tubes 9–16 mm deep. Stalk: enlarged downward or nearly equal; surface dry, glabrous to slightly fibrillose, white to pale tan, sometimes with a reddish zone near the apex; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups, often on the bases of trees or on the ground, under oaks and pines; year-round. Distribution and frequency: North Carolina south to Florida, west to Texas; occasional. Spore print: dark rusty brown to dark brown. Microscopic features: spores 15–24 × 7–11 µm, fusoid, with spirally arranged, conspicuous, longitudinally ridged, thin wings and often with an indistinct apical pore, pale brown. Boletellus ananas (B)

78 · Boletes of Eastern North America Edibility: edible. Lookalikes: Suillellus pictiformis Murrill = Boletellus pictiformis (Murrill) Singer (not illustrated) is a similar and rare species known only from Florida. It has a dry cap coated with appressed to erect, shaggy, medium-brown to darkbrown scales. Its cap becomes cracked in age, with yellow flesh showing in the fissures. Its stalk is sheathed with shaggy to fibrillose, medium-brown to dark-brown scales, and its greenish-yellow to olive-yellow pores stain blue when bruised. Boletellus flocculosipes (Murrill) Perr.-Bertr. (not illustrated), known only from Gainesville, Florida, has a dry, coarsely shaggy, dark grayish-brown to grayish-black cap, pale-yellow flesh with a pinkish zone near the cuticle that stains blue when exposed; sulfur-yellow pores that become dingy yellow to brownish at maturity and rapidly stain dark blue green when bruised; and a dry, shaggy, reddish-brown stalk with a glabrous yellow apex and a dark grayish-brown to grayish-black base. It has spores that measure 12–14 × 6–7 µm, and it grows in broadleaf woods near hickory. Boletellus chrysenteroides (Snell) Snell = Boletus chrysenteroides Snell Common name(s): none. Overview: This small, dark bolete is unusual in that it most often grows on decaying wood rather than in soil. The epithet chrysenteroides means Boletellus chrysenteroides

Boletellus · 79 “resembling chrysenteron,” a reference to its similarity with Xerocomellus chrysenteron (p. 416). Cap: 3–6 cm wide, convex, becoming broadly convex, margin even; surface dry, dark brown and velvety when young, becoming bay brown to chestnut brown and often cracked in age, with whitish to pale-yellow flesh that lacks reddish tints showing in the fissures; flesh whitish to pale yellow, staining blue, often slowly, when exposed; odor and taste not distinctive. Pores and tubes: pale yellow, becoming yellow to greenish yellow, quickly staining blue when cut or bruised; tubes 1.2–1.8 cm deep. Stalk: equal or enlarged slightly downward, solid; surface dry, reddish brown to blackish brown, often yellowish at the apex, often decorated with punctae that may align to simulate a net-like pattern, staining blue then slowly reddish when bruised or handled; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or in groups on decaying logs, stumps, at the base of standing trunks, or sometimes on the ground, in broadleaf or mixed oak and pine woods, especially with oak; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to fairly common. Spore print: olive brown to dark brown. Microscopic features: spores 10–16 × 5–8 µm, narrowly ovate to nearly oblong, longitudinally striate, yellowish to brownish. Edibility: edible. Lookalikes: Xerocomellus chrysenteron (p. 416) grows on the ground and has a dark-olive to olive-brown or grayish-brown cap, often with a reddish zone at the margin. Its cap soon becomes cracked and shows a red to pinkish color in the fissures. Xerocomellus intermedius (p. 419) has a cap that is grayish olive to brownish olive or red to rose red with olive tints and becomes cracked in age, with pinkish tints showing in the fissures. It grows on the ground. Boletellus pseudochrysenteroides (p. 80) grows on the ground or on decaying wood in broadleaf woods, especially with beech or oak. It has a dark rose-red to red-brown cap when young that typically becomes olivaceous to ochraceous brown and develops prominent cracks as it ages. It has whitish to pale-yellow flesh that quickly stains blue when exposed, and its stalk base is sometimes coated with yellowish mycelium.