Wittlake, Eugene B.

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Journal of the Arkansas Academy of ScienceVolume 23 Article 281969Fossil Phylloxerid Plant Galls From the LowerEoceneEugene B. WittlakeFollow this and additional works at: http://scholarworks.uark.edu/jaasPart of the Entomology CommonsThis article is available for use under the Creative Commons license: Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0). Users are able toread, download, copy, print, distribute, search, link to the full texts of these articles, or use them for any other lawful purpose, without asking priorpermission from the publisher or the author.This Article is brought to you for free and open access by ScholarWorks@UARK. It has been accepted for inclusion in Journal of the Arkansas Academyof Science by an authorized editor of ScholarWorks@UARK. For more information, please contact scholar@uark.edu, ccmiddle@uark.edu.Recommended CitationWittlake, Eugene B. (1969) "Fossil Phylloxerid Plant Galls From the Lower Eocene," Journal of the Arkansas Academy of Science: Vol. 23, Article 28.Available at: http://scholarworks.uark.edu/jaas/vol23/iss1/28164Arkansas Academy of Science Proceedings, Vol. 23, 1969•FOSSIL PHYLLOXERID PLANT GALLSFrom the Lower EoceneEugene B. WittlakeArkansas State UniverstiyFossil Plant galls were encountered in an investigation of sand-stone outcrops of the Upper Wilcox formation at Lafe, Arkansas.These deposits consist of very fine, buff-colored to whitish sand-stone. Great detail of a structural nature is recorded in individualspecimens. The deposit must represent the central area of a largelake or lagoon that was fed by streams with a moderate current. Allcoarse material in suspension had been filtered out or settled out,before the area of the deposit had been reached by the diminishingcurrent. Apparently, finer particles then slowly settled out afterbeing in suspension for some time. Individual sand grains are allof the same size. Another factor that lends support to the supposi-tion that the area of the deposits was some distance from land, isthe fact that the fossils are sparse and are chiefly the casts ofstems, leaves and fruits of aquatic, emergent plants. Dicotyledonousleaves exist here and there; most have entire margins. In the lastanalysis, monocotyledons dominate.Galls incited by insects are very scarce in the fossil recordto date. Berry (1916) cites two examples from the lower Eoceneand specifically locates them in the beds cf the Wilcox from Lagrangeformation near Puryear, Tennessee. Collins (1925) refers to themas found in the lower Eocene, and in another work Berry (1931)cites Wilcox specimens. Brooks (1955) received specimens collectedin the area near Puryear, Tennessee and reviews the literature onthe status of the fossil record of both galls and other types of in-jury to angiosperm fossil leaves. In fact, to the best knowledge ofthe author, these are the only papers there are on any reference tofossil insect galls from the lower Eocene. The remaining six papersin the Tertiary literature are in reference to galls found in theOligocene Epoch from Florissant, Colorado: Scudder (1886), Cock-rell (1908), Brues (1910) and Kinsey (1919); Hoffman (1932) hasdescribed Miocene galls and Brues (1946) refers to an observationof Cretaceous insect galls. None of the galls described in the pre-ceding works resemble the galls presented here. All evidence of thespecimens collected by the author, point to one insect family, namely,the Phylloxeridae.The Phylloxerids belong to the insect order, Homoptera, thefamily Phylloxeridae and the subfamily Phylloxeridinae. This family*This study was supported by a grant from the Faculty ResearchCommittee, Arkansas State University.164Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28Published by Arkansas Academy of Science, 1969165Fossil Phylloxerid Plant Gallsof aphids that attack modern hickories, for the most part produce agreat variety of plant galls. Felt (1940) states that 29 kinds of gallson Carya can be attributed to this family alone, and others appearon Quercus and Jug lands, not to mention the long known formsthat infest the leaves and roots of Vitis.Most phylloxerid galls have an orifice that is usually found onthe surface of the gall, exposed on the lower side of the leaf. Accord-ing to Comstock (1949) the orifice is produced by the emergenceof a young generation of aphids that hatch from the egg-packedgall. The eggs are laid by a wingless, agamic, stem-mother aphidwhose activity initiates the growth of the gall to its mature size.In some forms, galls of this type are armed around the orifice andrim of the gall, by spines or bristle-like plant hairs.The fossil galls in question, lie well within the limits of size,shape and other characteristics exhibted by present-day phylloxeridgalls that occur on plants belonging to the Junglandaceae andspecifically in the genus Carya. In fact, the fossil forms presentedhere have characteristics very close to the gall produced by Phylloxerarimosalis Perg. on hickory, described and illustrated by Felt.The mature fossil galls average 9mm. long and 6 to 7mm. wide.The conical body of the gall is 2-3mm. high. Young to older gallsrange in length from 2 to 12mm. Three to four of the larger gallsshow raised and rimmed margins with compression of equally-spaced spines or bristles, 14 in number and attached at their basesto the flange-like rim of the gall. The orifice at the summit of thecone in the mature galls, ranged in diameter from 0.75mm. to 2.5mm. The fossil galls are clustered in aggregates as are moderngalls.The leaf compressions on which the fossil galls are situated,are not detailed enough to show their complete outline, and themargins of the leaves are not too distinct. However, the leaf marginsappear to be serrulate and not entire. On the suface of the compres-sions are many casts of leaf hairs. Perhaps this indicates that theunderside of the leaf is exposed rather than the upper side. Mostof the species in the Juglandaceac have leaves that are clothed withlairs on the underside, at least when the leaves are young. Manyretain this hirsute nature in the mature leaves. Some species in themodern flora have minute glands along the veins and distributedamong the plant hairs. These fossil forms also show gland distributedllong the only midrib visible in these compressions. As a result ofthe above dscussion, the author makes no commitment as to thegeneric designation of the fossil leaf compressions because they aretoo fragmentary. In the light of the present day restriction of thisype of gall on a number of species in the family Juglandaceae,t seems likely in the author's opinion that these leaf compressions 165Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28http://scholarworks.uark.edu/jaas/vol23/iss1/28166do irepresent some genus in this plant family. It has been shownby past studies of other authors, i.e. Berry (191G). (1924) and(1930a), (1930b), Knowlton (1922), MacGinitie (1941) and Potbury(1935) that genera of the Juglandaceae were prevalent throughoutthe Eocene epoch.In Fig. 1, a complete restoration of the mature fossil gall isillustrated, as seen from above with its rim fringed with spines. Fig.2 is a diagrammatic vertical section of the fossil gall. Figs. 3 and 4are photographs of the total number of young and mature fossilgall specimens studied in this work. Specimens illustrated in Figs.3 and 4 (L 66031 and L 64101) are in the Paleobotanical Herbariumof Arkansas State University Museum.In conclusion, the author wishes to thank Dr. Harvey E. Bartonfor his analysis and comments on these fossil specimens.rigs. 1-4: Fig. 1, Restotration of Fossil Gall, Fig. 2, Vertical sectionof gall, Fig. 3 and 4, Fossil gall specimens. X1.25166Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28Published by Arkansas Academy of Science, 1969167Fossil Phylloxerid Plant GallsLITERATURE CITEDKinsey, A. C. 1919. Fossil Cynipidae: Psyche 26: 44-49.Knowlton, F. H. 1922. Revision of the Flora of the Green River For-mation, U.S. Geol. Surv. Prof. Paper 131f. pp. 133-176.MacGinitie, H D. 1941. A Middle Eocene Flora from the CentralSierra Nevada, Carnegie Inst. Wash. Pub. 534.Potbury, S. S. 1935. The LaPorte Flora of Plumas County, Cal. Car-negie Inst. Wash. Pub. 465, II., pp. 29-81.Scudder, S. H. 1886. Systematic Review of Fossil Insects, U.S. Geol.Surv. Bull., 31.Berry, E. W., 1016. The Lower Eocene Floras of Southeastern NorthAmerica, U.S. Geol. Surv. Prof. Paper 91.1924. The Middle and Upper Eocene Floras ofSoutheastern North America, U.S. Geol. Surv. Prof. Paper 92.1930a. Revision of the Lower Eocene Wilcox Floraof the Southeastern States, U.S. Geol. Surv. Prof. Paper 156.1930b. A Flora of Green River Age in the WindRiver Basin of Wyoming, U.S. Geol. Surv. Prof. Paper 165,pp. 55-79.1931. An Insect-cut Leaf from the Lower Eocene:Amer. Journ. Sci., 21: 301-304.Brooks, H. K. 1955. Healed Wounds and Galls on Fossil Leavesfrom Wilcox Deposits (Eocene) of Western Tennessee: Psyche62:1-9.Brues, C. T. 1910. The Parasitic Hymenoptera of the Tertiary ofFlorissant, Colorado: Bull. Mus. Comp. Zool. 54: 1-125.Cockrell, T. D. A. 1908. Fossil Insects from Florissant, Colorado:Amer. Mus. Nat. Hist., 24: 59-69.Collins, R. L. 1925. A Lower Eocene Termite from Tennessee: Amer.Journ. Sci. ser. 5, 9: 406410.Comstock, J. H. 1949. An Introduction to Entomology, ComstockPub. Co. p. 434. N.Y.Felt, E. P. 1940. Plant Galls and Gall Makers, Comstock Pub. Co.Hoffman, A. 1932. Miocene Insect Gall Impressions: Bot. Gazette93: 341-342.N.Y.167Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28http://scholarworks.uark.edu/jaas/vol23/iss1/28

Fossil Phylloxerid Plant Galls From the Lower Eocene

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Journal of the Arkansas Academy of Science Volume 23 Article 28 1969 Fossil Phylloxerid Plant Galls From the Lower Eocene Eugene B. Wittlake Follow this and additional works at: https://scholarworks.uark.edu/jaas  Part of the Entomology Commons Recommended Citation Wittlake, Eugene B. (1969) "Fossil Phylloxerid Plant Galls From the Lower Eocene," Journal of the Arkansas Academy of Science: Vol. 23 , Article 28. Available at: https://scholarworks.uark.edu/jaas/vol23/iss1/28 This article is available for use under the Creative Commons license: Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0). Users are able to read, download, copy, print, distribute, search, link to the full texts of these articles, or use them for any other lawful purpose, without asking prior permission from the publisher or the author. This Article is brought to you for free and open access by ScholarWorks@UARK. It has been accepted for inclusion in Journal of the Arkansas Academy of Science by an authorized editor of ScholarWorks@UARK. For more information, please contact scholar@uark.edu. 164Arkansas Academy of Science Proceedings, Vol. 23, 1969•FOSSIL PHYLLOXERID PLANT GALLSFrom the Lower EoceneEugene B. WittlakeArkansas State UniverstiyFossil Plant galls were encountered in an investigation of sand-stone outcrops of the Upper Wilcox formation at Lafe, Arkansas.These deposits consist of very fine, buff-colored to whitish sand-stone. Great detail of a structural nature is recorded in individualspecimens. The deposit must represent the central area of a largelake or lagoon that was fed by streams with a moderate current. Allcoarse material in suspension had been filtered out or settled out,before the area of the deposit had been reached by the diminishingcurrent. Apparently, finer particles then slowly settled out afterbeing in suspension for some time. Individual sand grains are allof the same size. Another factor that lends support to the supposi-tion that the area of the deposits was some distance from land, isthe fact that the fossils are sparse and are chiefly the casts ofstems, leaves and fruits of aquatic, emergent plants. Dicotyledonousleaves exist here and there; most have entire margins. In the lastanalysis, monocotyledons dominate.Galls incited by insects are very scarce in the fossil recordto date. Berry (1916) cites two examples from the lower Eoceneand specifically locates them in the beds cf the Wilcox from Lagrangeformation near Puryear, Tennessee. Collins (1925) refers to themas found in the lower Eocene, and in another work Berry (1931)cites Wilcox specimens. Brooks (1955) received specimens collectedin the area near Puryear, Tennessee and reviews the literature onthe status of the fossil record of both galls and other types of in-jury to angiosperm fossil leaves. In fact, to the best knowledge ofthe author, these are the only papers there are on any reference tofossil insect galls from the lower Eocene. The remaining six papersin the Tertiary literature are in reference to galls found in theOligocene Epoch from Florissant, Colorado: Scudder (1886), Cock-rell (1908), Brues (1910) and Kinsey (1919); Hoffman (1932) hasdescribed Miocene galls and Brues (1946) refers to an observationof Cretaceous insect galls. None of the galls described in the pre-ceding works resemble the galls presented here. All evidence of thespecimens collected by the author, point to one insect family, namely,the Phylloxeridae.The Phylloxerids belong to the insect order, Homoptera, thefamily Phylloxeridae and the subfamily Phylloxeridinae. This family*This study was supported by a grant from the Faculty ResearchCommittee, Arkansas State University.164Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28Published by Arkansas Academy of Science, 1969165Fossil Phylloxerid Plant Gallsof aphids that attack modern hickories, for the most part produce agreat variety of plant galls. Felt (1940) states that 29 kinds of gallson Carya can be attributed to this family alone, and others appearon Quercus and Jug lands, not to mention the long known formsthat infest the leaves and roots of Vitis.Most phylloxerid galls have an orifice that is usually found onthe surface of the gall, exposed on the lower side of the leaf. Accord-ing to Comstock (1949) the orifice is produced by the emergenceof a young generation of aphids that hatch from the egg-packedgall. The eggs are laid by a wingless, agamic, stem-mother aphidwhose activity initiates the growth of the gall to its mature size.In some forms, galls of this type are armed around the orifice andrim of the gall, by spines or bristle-like plant hairs.The fossil galls in question, lie well within the limits of size,shape and other characteristics exhibted by present-day phylloxeridgalls that occur on plants belonging to the Junglandaceae andspecifically in the genus Carya. In fact, the fossil forms presentedhere have characteristics very close to the gall produced by Phylloxerarimosalis Perg. on hickory, described and illustrated by Felt.The mature fossil galls average 9mm. long and 6 to 7mm. wide.The conical body of the gall is 2-3mm. high. Young to older gallsrange in length from 2 to 12mm. Three to four of the larger gallsshow raised and rimmed margins with compression of equally-spaced spines or bristles, 14 in number and attached at their basesto the flange-like rim of the gall. The orifice at the summit of thecone in the mature galls, ranged in diameter from 0.75mm. to 2.5mm. The fossil galls are clustered in aggregates as are moderngalls.The leaf compressions on which the fossil galls are situated,are not detailed enough to show their complete outline, and themargins of the leaves are not too distinct. However, the leaf marginsappear to be serrulate and not entire. On the suface of the compres-sions are many casts of leaf hairs. Perhaps this indicates that theunderside of the leaf is exposed rather than the upper side. Mostof the species in the Juglandaceac have leaves that are clothed withlairs on the underside, at least when the leaves are young. Manyretain this hirsute nature in the mature leaves. Some species in themodern flora have minute glands along the veins and distributedamong the plant hairs. These fossil forms also show gland distributedllong the only midrib visible in these compressions. As a result ofthe above dscussion, the author makes no commitment as to thegeneric designation of the fossil leaf compressions because they aretoo fragmentary. In the light of the present day restriction of thisype of gall on a number of species in the family Juglandaceae,t seems likely in the author's opinion that these leaf compressions 165Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28https://scholarworks.uark.edu/jaas/vol23/iss1/28166do irepresent some genus in this plant family. It has been shownby past studies of other authors, i.e. Berry (191G). (1924) and(1930a), (1930b), Knowlton (1922), MacGinitie (1941) and Potbury(1935) that genera of the Juglandaceae were prevalent throughoutthe Eocene epoch.In Fig. 1, a complete restoration of the mature fossil gall isillustrated, as seen from above with its rim fringed with spines. Fig.2 is a diagrammatic vertical section of the fossil gall. Figs. 3 and 4are photographs of the total number of young and mature fossilgall specimens studied in this work. Specimens illustrated in Figs.3 and 4 (L 66031 and L 64101) are in the Paleobotanical Herbariumof Arkansas State University Museum.In conclusion, the author wishes to thank Dr. Harvey E. Bartonfor his analysis and comments on these fossil specimens.rigs. 1-4: Fig. 1, Restotration of Fossil Gall, Fig. 2, Vertical sectionof gall, Fig. 3 and 4, Fossil gall specimens. X1.25166Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28Published by Arkansas Academy of Science, 1969167Fossil Phylloxerid Plant GallsLITERATURE CITEDKinsey, A. C. 1919. Fossil Cynipidae: Psyche 26: 44-49.Knowlton, F. H. 1922. Revision of the Flora of the Green River For-mation, U.S. Geol. Surv. Prof. Paper 131f. pp. 133-176.MacGinitie, H D. 1941. A Middle Eocene Flora from the CentralSierra Nevada, Carnegie Inst. Wash. Pub. 534.Potbury, S. S. 1935. The LaPorte Flora of Plumas County, Cal. Car-negie Inst. Wash. Pub. 465, II., pp. 29-81.Scudder, S. H. 1886. Systematic Review of Fossil Insects, U.S. Geol.Surv. Bull., 31.Berry, E. W., 1016. The Lower Eocene Floras of Southeastern NorthAmerica, U.S. Geol. Surv. Prof. Paper 91.1924. The Middle and Upper Eocene Floras ofSoutheastern North America, U.S. Geol. Surv. Prof. Paper 92.1930a. Revision of the Lower Eocene Wilcox Floraof the Southeastern States, U.S. Geol. Surv. Prof. Paper 156.1930b. A Flora of Green River Age in the WindRiver Basin of Wyoming, U.S. Geol. Surv. Prof. Paper 165,pp. 55-79.1931. An Insect-cut Leaf from the Lower Eocene:Amer. Journ. Sci., 21: 301-304.Brooks, H. K. 1955. Healed Wounds and Galls on Fossil Leavesfrom Wilcox Deposits (Eocene) of Western Tennessee: Psyche62:1-9.Brues, C. T. 1910. The Parasitic Hymenoptera of the Tertiary ofFlorissant, Colorado: Bull. Mus. Comp. Zool. 54: 1-125.Cockrell, T. D. A. 1908. Fossil Insects from Florissant, Colorado:Amer. Mus. Nat. Hist., 24: 59-69.Collins, R. L. 1925. A Lower Eocene Termite from Tennessee: Amer.Journ. Sci. ser. 5, 9: 406410.Comstock, J. H. 1949. An Introduction to Entomology, ComstockPub. Co. p. 434. N.Y.Felt, E. P. 1940. Plant Galls and Gall Makers, Comstock Pub. Co.Hoffman, A. 1932. Miocene Insect Gall Impressions: Bot. Gazette93: 341-342.N.Y.167Journal of the Arkansas Academy of Science, Vol. 23 [1969], Art. 28https://scholarworks.uark.edu/jaas/vol23/iss1/28

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Fossil Phylloxerid Plant Galls From the Lower Eocene

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