Table 1.—Beetle- and host tree-associated compounds found to affect southern pine beetle behavior.
COMPOUND / REFERENCE / SOURCE / BEHAVIORAL EFFECT / METHOD OF INVESTIGATIONFrontalin (racemic)(+)(-) / 1, 2 / A, B, C, D / Attractant (1, 14, 15) / Field & laboratory bioassay
Endo-brevicomin / 3, 4 / A, C, D / Inhibitor (14, 16, 17) / Field & laboratory bioassay
Trans-verbenol / 5, 2 / A, B, C, D / Synergist / Field & laboratory bioassay
Cis-verbenol (racemic)(R)(S) / 6, 2 / B, C, D
Verbenone / 5, 2 / A, B, C, D / Multifuntion Inhibitor/Synergist (14, 15, 17, 18) / Field & laboratory bioassay
Alpha-pinene / 7, 2 / C, D / Synergist (1, 7, 14, 15) / Field & laboratory bioassay
Myrcene / 2 / C, D / Attractant (19) / Laboratory bioassay
Myrtenol / 8,9 / A, B, C, D / Synergist (20) / Laboratory bioassay
Cis-myrtanol / 2 / B, D, C
Isoamyl acetate / 10 / E / Synergist (10) / Laboratory bioassay
Isoamyl alcohol / 10, 11 / E, F / Synergist (10) / Laboratory bioassay
2-phenylethanol / 10 / E / Synergist (10) / Laboratory bioassay
2-phenylethyl acetate / 10, 11 / E / Synergist (10) / Laboratory bioassay
6-methyl-5-hepten-2-one / 11 / F
6-methyl-5-hepten-2-ol / 11 / F
4-methyl-2-pentanol / 9 / A, B
6-hydroxycamphene / 12, 2 / A, B, C, D
Pinocarvone / 9, 2 / A, B, C, D
Trans-pinocarveol / 9, 2 / A, B, C, D
Myrtenal / 9, 2 / A, B, C, D
Acetophenone / 2 / A, C, D
Chrysanthenone / 2 / A, C, D
4-Allylanisole / 2 / C, D
Cis-3-pinen-2-ol / 13 / B
3-methyl-2-cyclohexen-l-ol (seudenol) / 13 / A, B
1-methyl-2-cyclohexen-l-ol / 13 / A, B
1-cyclohexenemethanol / 13 / A, B
3-methyl-2-cyclohexen-1-one (MCH) / 13 / A, B
2-methyl-2-cyclohexen-1-ol / 13 / A, B
1—Kinzer et al. 1969
2—R.M. Silverstein and J.R. West, personal communication
3—Silverstein et al. 1968
4—Pitman et al. 1969
5—Renwick 1967
6—Silverstein et al. 1966
7—Renwick and Vité 1969
8—Hughes 1973
9—Renwick et al. 1973
10—Brand et al. 1977
11—Brand and Barras 1977
12—Renwick et al. 1976
13—Renwick and Hughes 1975
14—Payne et al. 1978a
15—McCarty et al. 1980
16—Vité and Renwick 1971
17—Richerson and Payne 1979
18—Rudinsky 1973
19—P.D. Billings, personal communication
20—Rudinsky et al. 1974
A—Male hindguts
B—Female hindguts
C—Beetle-infested tree parts
D—Female frass
E—Yeast metabolite
F—Basidiomycete
Table 2.—Arthropod predators in the southern pine beetle (References indicated in bold type specify predatory role with the SPB. Others recorded only as predators of bark beetles or predators under bark. Unconfirmed predatory roles indicated by [?].) Reference designated by ( ).
Hemiptera
Anthocoridae
Lyctoris campestris (Fab.)(6)
Lyctoris elongatus (Reuter)(1)(2)(3)(5)(6)(7)(8)(9)
Scoloposcelis flavicornis (Reuter)(2)(9)
Scoloposcelis mississippensis Drake & Harris (1)(3)(5)(6)(7)(8)
Aradidae
Aradus cinnamomeus (Panzer)(8)
Pentatomidae
Diolcus chryssorhoesus (Fab.)(8)
Coleoptera
Carabidae
Apristus subsulcatus (Dejean)(6)
Dromius piceus Dejean (6)
Pinacodera limbata Dejean (3)
Pinacodera platicollis (Say)(6)
Stenolophus lineola (3?)
Tachyta pavicornis Notman (3)
Histeridae
Abraeus sp. (3)
Cylistix attenuata Lec. (3)(7)
Cylistix cylindrica (Payk.)(1)(3)(5)(6)(7)
Epierus pulicarius Erichson (7)
Hister sp. (3)
Platysome parallelum Say (3)(5)(7)
Plegaderus pusillus Lec. (7)
Plegaderus transversus Say (7)
Plegaderus sp. (2 spp.?)(5)(6)
Staphylinidae
Aleocharinae (3)
Leptacinus paurumpunctatus (Gyll.)(3?)(6)
Nacaeus tenellus Erichson (3?)
Pseudolispinodes tenellus Erichson (7)
Silusa sp. (6)
sp. (undetermined)(4 spp.?)(3)(7)
Orthoperidae
Molamba sp. (3)
Sacium sp. (7?)
Trogositidae
Corticotomus parallelus Melsh. (3)
Temnochila virescens (Fab.)(1)(2)(3)(6)(7)(8)(9)
Tenebroides collaris (Sturm)(1)(2)(3)(7)(8)(9)
Tenebroides marginatus (P. de B.)(3)(7)
Tenebroides nanus (Melsh.)(3)
Tenebroides sp. (6)
Cleridae
Cregya oculata (Say)(3?)
Cymatodera undulata (Say)(3)
Enoclerus quadriguttatus Oliv. (2)(9)
Phyllobaenus pallipennis Dejean (3?)
Priocera castanea (Newm.)(2)(3)(9)
Thanasimus dubius (Fab.)(1)(2)(3)(5)(6)(7)(8)(9)
Tillus collaris Spin. (3?)
Elateridae
Anchasius signaticollis (Germ.)(3)
Athous sp. (3)
Elateridae sp. (9)
Glyphonyx sp. (3)
Lacon impressicollis (Say)(3)(8)
Melanotus sp. (2 spp.?)(3)(8)
sp. (undetermined)(3)
Eucnemidae
Dirhagus triangularis (Say)(3)
Dirhagus sp. (3)
Throscidae
Trixagus sp. (3)
Cucujidae
Ahasverus advena (Waltl)(8)
Carthartosilvanus imbellis (Lec.)(3)
Nausibus clavicornis (Kug.)(3)
Silvanus bidentatus (Fab.)(7?)
Silvanus muticus Sharp (3)
sp. (unidentified)(2 spp.?)(3)
Colydiidae
Aulonium ferrugineum Zimm. (3)(5)(6)(7?)
Aulonium tuberculatum Lec. (3)(5)(6)
Colydium lineola Say (3?)
Colydium nigripenne Lec. (3)
Lasconotus pusillus Lec. (3)(6)
Lasconotus referendarius Zimm. (3)(7)
Pycnomerus sulcicollis Lec. (3)
Cerylonidae
Cerylon castaneum Say (3)
Mordellidae
sp. (unidentified)(2 spp.?)(3?)
Tenebrionidae
Corticeus glaber (Lec.)(3)(7)
Corticeus parallelus Melsh. (7)(9 as Hypophloeus parallelus)
Corticeus sp. (2 spp.?)(5)(6)
Coccinelidae
Nephus bioculatus Mulsant (8)
Melyridae
Melyrodes cribratus Lec. (7)
Rhizophagidae
Rhizophagus sp. (7)
Diptera
Stratiomyidae
Zabrachia sp. (3)
Empidae
Euhybus sp. near gentivus Melander (undescribed)(3)
Syndas polita (Loew)(3)
Dolichopodidae
Medetera bistriata Parent (1)(5)(7)(recorded as parasite in 6)
Medetera maura Wheeler (8)
Medetera sp. (2 spp.?)(2)(recorded as parasite in 6)
Phoridae
Dorniphora sp. (3?)
Lonchaeidae
Lonchaea auranticornis McAlpine (7?)
Lonchaea polita Say (7?)
Lonchaea sp. (3)
Hymenoptera
Formicidae
Camponotus clarus Mayr (7)
Camponotus nearcticus Emery (7)
Camponotus sayi Emery (3)
Crematogaster ashmeadi Mayr (3)(7)
Crematogaster sp. (3)
Cryptopone gilva (Roger)(3)
Dorymyrmex pyramicus (Roger)(7)
Hypoponera opacior (Forel)(3)
Leptothorax schaumi Roger (7)
Monomorium minimum (Buckley)(7)
Paratrechina parvula (Mayr)(7)
Pheidole metallescens metallescens Emery (7)
Pheidole sp. (3)
Proceratium croceum (Roger)(3)
Solenopsis picta Emery (3)
Solenopsis sp. (7)
Sp. (undetermined)(6)
References for Table 2
(1) Coulson, R.N., T.L. Payne, J.E. Coster, and M.W. Houseweart, 1972. The southern pine beetle Dendroctonus frontalis Zimmerman (Coleoptera: Scolytidae) 1961-1971. Tex. For. Serv., Pub. 108. College Station, Tex.
(2) Dixon, J.C., and E.A. Osgood. 1961. Southern pine beetle: a review of present knowledge. U.S. Dep. Agric. For. Serv., Res. Pap. SE-128. Southeast For. Exp. Stn., Asheville, N.C.
(3) Dixon, W.N., and T.L. Payne. 1979. Sequence of arrival and spatial distribution of entomophagous and associated insects on southern pine beetle-infested trees. Tex. Agric. Exp. Stn., MP-1432. College Station, Tex.
(4) Lenhard, G.J., and R.A. Goyer. 1979. The relative abundance and seasonal distribution of the major predators of the southern pine beetle in loblolly pine. [Unpublished ms.]
(5) Moore, G.E. 1972. Southern pine beetle mortality in North Carolina caused by parasites and predators. Environ. Entomol. 1:58-65.
(6) Moser, J.C., R.C. Thatcher, and L.S. Pickard. 1971. Relative abundance of southern pine beetle associates in east Texas. Ann. Entomol. Soc. Am. 64:72-77.
(7) Overgaard, N.A. 1968. Insects associated with the southern pine beetle in Texas, Louisiana, and Mississippi. J. Econ. Entomol. 61:1197-1201.
(8) Thatcher, R.C. 1960. Bark beetles affecting southern pines: a review of current knowledge. U.S. Dep. Agric. For. Serv., South. For. Exp. Stn., Occas. Pap. 180. [Discontinued South. Stn. series.]
Table 3.—Arthropod parasitoids of the southern pine beetle (References indicated in bold type specify parasitic role with the SPB. Others recorded only as parasitoids of bark beetles. Unconfirmed parasitic roles indicated by [?].) References designated by ( ).
Hymenoptera
Braconidae
Atanycolus comosifrons Shenefelt (6)(7?)
Atanycolus ulmicola (Vier.)(3?)
Cenocoelius nigrisoma (Rohwer)(1)(3)(7)
Cenocoelius sp. (6)
Coeloides pissodis (Ashm.)(1)(2)(3)(4)(6)(7)(9)
Compyloneurus movoritus (Cress.)(2)(9)
Dendrosoter sulcatus Mues. (1)(2)(3)(4)(6)(7)(9)
Doryctes sp. (1)(2)(3)(7)(9)
Heterospilus sp. (6)
Meteorus hypophloei Cushman (6)
Spathius canadensis Ashm. (1)(2)(9)
Spathius pallidus Ashm. (1)(3)(4)(6)(7)(9)
Vipio rugator (Say)(1)
Ichneumondiae
Cremastus sp. (6)
sp. (undetermined)(3)
Eupelmidae
Arachnophaga sp. (3)
Eupelmus cyaniceps cyaniceps (Ashm.)(7)
Lutnes sp. (3)(7?)
Torymidae
Liodontomerus sp. (1)
Lochites sp. (2)(9)
Roptrocerus eccoptogastri (Ratz.)(1)(2)(3)(8)(9)
Roptrocerus xylophagorum Ratz. (1)(4)(6)(7)(8)
Roptrocerus sp. (6)
Pteromalidae
Dinotiscus (=Cecidostiba) dendroctoni (Ashm.)(1)(2)(3)(4)(6)(9)
Heydenia unica Cook & Davis (1)(2)(3)(4)(6)(7)
Rhopalicus pulchripennis (Crawford)(3)(7)
Eurytomidae
Eurytoma cleri (Ashm.)(6)
Eurytoma tomici Ashm. (1)(7)
Eurytoma sp. (3)
Scelionidae
Gyron sp. (8)
Idris sp. (8)
Leptoteleia sp. (6)
Probaryconus heidemanni Ashm. (8)
Telenonus podisi Ashm. (8)
Bethylidae
Parasierola sp. (3)
References for Table 3
(1) Coulson, R.N., T.L. Payne, J.E. Coster, and M.W. Houseweart. 1972. The southern pine beetle Dendroctonus frontalis Zimmerman (Coleoptera: Scolytidae) 1961-1971. Tex. For. Serv., Pub. 108. College Station, Tex.
(2) Dixon, J.C. and E.A. Osgood. 1961. Southern pine beetle: a review of present knowledge. U.S. Dep. Agric. For. Serv., Res. Pap. SE-128. Southeast. For. Exp. Stn., Asheville, N.C.
(3) Dixon, W.N., and T.L. Payne. 1979. Sequence of arrival and spatial distribution of entomophagous and associated insects on southern pine beetle-infested trees. Tex. Agric. Exp. Stn., MP-1432. College Station, Tex.
(4) Goyer, R.A., and C.K. Finger. 1980. The relative abundance and seasonal distribution of the major hymenopterous parasites of the southern pine beetle, Dendroctonus frontalis Zimmerman, on loblolly pine. Environ. Entomol. 9:97-100.
(5) Lenhard, G.J., and R.A. Goyer. 1979. The relative abundance and seasonal distribution of the major predators of the southern pine beetle in loblolly pine. [Unpublished ms.]
(6) Moore, G.E. 1972. Southern pine beetle mortality in North Carolina caused by parasites and predators. Environ. Entomol. 1:58-65.
(7) Moser, J.C., R.C. Thatcher, and L.S. Pickard. 1971. Relative abundance of southern pine beetle associates in east Texas. Ann. Entomol. Soc. Am. 64:72-77.
(8) Overgaard, N.A. 1968. Insects associated with the southern pine beetle in Texas, Louisiana, and Mississippi. J. Econ. Entomol. 61:1197-1201.
(9) Thatcher, R.C. 1960. Bark beetles affecting southern pines: a review of current knowledge. U.S. Dep. Agric. For. Serv., South. For. Exp. Stn., Occas. Pap. 180. [Discontinued South. Stn. series.]
Table 4.—Key to the final instar larvae of the major parasites of the southern pine beetle. (From Finger and Goyer 1978).
1 / Body covered with many microspines; head with many sclerites; spiracles on segments 1 and 4-11…21’ / Body with some setae but without microspines; head with few if any sclerites; spiracles on segments 2-10...4
2 / Labial sclerite very thick and rounded, often with slight projection on ventral surface and flat on dorsal surface between arms; silk orifice on wide oval sclerite…Dendrosoter sulcatus (figs. 1B, 2B, 3B)
2’ / Labial sclerite not as above…3
3 / Thickness of ventral part of labial sclerite about 2 times as wide as where dorsal arms start; area inside labial sclerite more circular than ovoid; silk orifice forming a straight line (figs. 1A, 2A, 3A)… Coeloides pissodis
3’ / Thickness of ventral part of labial sclerite at least 3 times as thick as where arms start; area inside labial sclerite more ovoid; silk orifice often with slight “V” in middle (figs. 1C, 2C, 3C)…Spathius pallidus
4 / Head with very long setae; stalk of spiracle with over 20 chambers (figs. 1D, 2E, 3D)….Heydenia unica
4’ / Head with short setae; stalk of spiracle with less than 20 chambers……5
5 / Stalk of spiracle with less than 9 chambers, each decreasing in size from the atrium, forming a continuous funnel-shaped spiracle (figs. 2F, 3E)…Dinotiscus dendroctoni
5’ / Stalk of spiracle with more than 9 chambers, only first 3 chambers and atrium forming enlarged club-shaped structure (fig. 2G)…Roptrocerus eccoptogastri
Table 5.—Mites that prey on the southern pine beetle
Kleemannia sp.
Lasioseius dentatus (Fox)
Lasioseius epicriodopsis DeLeon
Lasioseius tubiculiger (Berlese)
Proctolaelaps bickleyi (Bram)
Proctolaelaps dendroctoni Lindquist and Hunter
Proctolaelaps fiseri Samsinak
Proctolaelaps hystricoides Lindquist and Hunter
Proctolaelaps hystrix (Vitzthum)
Dendrolaelaps isodentatus Hurlbutt
Dendrolaelaps neocornutus Hurlbutt
Dendrolaelaps neodisetus Hurlbutt
Dendrolaelaps rotoni Hurlbutt
Dendrolaelaps varipunctatus Hurlbutt
Androlaelaps casalis (Berlese)
Hypoaspis sp. nr. praesternalis Willman
Hypoaspis vitzthumi (Womersley)
Pseudoparasitus thatcheri Hunter and Moser
Macrocheles boudreauxi Krantz
Macrocheles mammifer Berlese
Eugamasus lyriformis McGraw and Farrier
Gamasolaelaps subcorticalis McGraw and Farrier
Cercoleipus coelonotus Kinn
Pleuronectocelaeno drymoecetes Kinn
Trichouropoda lamellose Hirschmann
Uroobovella americana Hirschmann
Histiogaster arborsignis Woodring
Histiogaster rotundus Woodring
Pyemotes parviscolyti Cross and Moser
Iponemus calligraphi calligraphi Lindquist
Iponemus confuses oriens Lindquist
Iponemus truncates eurus Lindquist
Table 6.—Key to mites commonly associated with the southern pine beetle. (From D.N. Kinn 1976.)
1 / Discernible without the aid of a lens; often red-brown in color; body hardened, with many shields or plates; stigmata (respiratory openings) located lateral to the bases of legs III and IV; tritosternum present; special sensory hairs not present on dorsal surface (fig. 1)…Order Parasitiformes—Suborder Mesostigmata…21’ / Usually small and light in color; body without numerous plates, stigmata not located lateral to bases of legs III and IV; tritosternum absent; special sensory hairs may be present on the anterior dorsal surface (fig. 4)…Order Acariformes…8
2 / Turtle shaped; leggs can be withdrawn into grooves (fig. 2); attached to beetle by anal pedicle…Superfamily Uropodoidea—Family Uropodiadae…3
2’ / Not turtle shaped; leg grooves absent; attached to beetle by the mouthparts and/or leg claws…Superfamily Parasioidea…4
3 / Anal shield with 14 hairs (fig. 2A); length about one-tenth of host’s length…Trichouropoda australis
3’ / Anal shield with 10 hairs (fig. 2B); length about one-third of host’s length…Trichouropoda hirsuta
4 / Large red-brown mite lacking claws on leg I; peritremes looped, joining stigmata posteriorly (fig. 1C)…Family Macrochelidae…Marocheles boudreauxi
4’ / Leg I with claws; peritremes not looped; joining stigmata anteriorly (fig. 1B)…5
5 / Dorsal shield entire; posterior end rounded (fig. 3A)…Family Ascidae…6
5’ / Dorsal shield divided into two plates; posterior end more or less truncated (fig. 3B)…Family Digamasellidae…7
6 / Ventral surface with four shields (sternal, genital, ventral, and anal)(fig. 1A)…Proctogastrolaelaps libris
6’ / Ventral surface with three shields (sternal, genital, and anal)(fig. 1B)…Proctolaelaps dendroctoni
7 / Body about two times longer than wide (fig. 3B)…Dendrolaelaps neodisetus
7’ / Body about four times longer than wide (fig. 3C)…Longoseius cuniculus
8 / Mouthparts functional; anal suckers absent; special sensory hairs present on anterior dorsal surface (fig. 4)…Suborder Prostigmata…9
8’ / Mouthparts vestigial; anal suckers present; special sensory hairs not present on anterior dorsal surface (fig. 5)…Suborder Astigmata…13
9 / Small, slow-moving mites often found under the beetle’s wing covers or around the leg bases; mouthparts indistinct; sensory hairs club-shaped; legs short in relation to body (figs 4B and 6)…Superfamily Tarsonemoidea…10
9’ / Fast-moving, orange-colored mite; mouthparts distinct; sensory hairs long and barbed; legs long in relation to body (fig. 4A)…Superfamily Tydeoidae—Family Ereynetidae…Ereynetoides scutulis
10 / Legs IV without claws and terminating with two whiplike hairs (fig. 6)…Family Tarsonemidae…11
10’ / Legs II-IV terminating with two claws; legs I terminate with a single large claw (fig. 4B)…Family Pyemotidae…Pygmephorus bennetti
11 / Legs II and III each with two claws; claw of leg I short, stout, and straight (fig. 6A)…Heterotarsonemus lindquisti
11’ / Legs II and III each with two claws; claw of leg I single, not modified (fig. 6B, C)…12
12 / Cuticular thickenings anterior to bases of legs III extending laterally beyond bases of legs III; lobe between bases of legs IV not elongated (fig. 6B)…Tarsonemus krantzi
12’ / Cuticular thickenings anterior to bases of legs III not extending laterally beyond bases of legs III; lobe between bases of legs IV very elongated and extending behind bases of legs IV (fig. 6C)…Tarsonemus ips
13 / All legs short and stout; legs III and IV; leg III and IVoften directed backward; distal segments of legs III and IV short (fig. 5A)…Superfamily Acaroidea—Family Acaridae…Histiogaster arborsignis
13’ / Legs I and II stouter than legs III and IV; legs III and IV often directed forward and have long, slender distal segments (fig. 5B, C)…Superfamily Anoetoidea—Family Anoetidae…14
14 / Entire dorsal surface always ornamented; fused mouthparts project well beyond body outline; dorsal hairs short and slender (fig. 5B)…Anoetus sordida
14’ / Dorsal ornamentation variable; fused mouthparts do not usually project beyond anterior edge of body; dorsal hairs long and thick (fig. 5C)…Anoetus varia
Table 7.—Frequently encountered pathogens of southern pine beetle (total of 13,458 diseased specimens) in Mississippi and Alabama (1975-1977)